TFSSvol1 3 PDF

TREE FLORA
of
SABAH AND SARAWAK
Produced with the financial support of
GOVERNMENT OF MALAYSIA
INTERNATIONAL TROPICAL TIMBER ORGANIZATION
OVERSEAS DEVELOPMENT ADMINISTRATION, U.K.
Tree Flora of Sabah and Sarawak Editorial Committee Chairman: E. Soepadmo

(Forest Research Institute Malaysia). Members: Abdul Latiff Mohamed (Universiti
Kebangsaan Malaysia), Ruth Kiew (Universiti Pertanian Malaysia), H.S. Lee
(Sarawak Forestry Department), K.M. Wong (Sabah Forestry Department). Editorial
assistant (Volume One): R.C.K. Chung (Forest Research Institute Malaysia).
Advisors: P.S. Ashton (Harvard University, U.S.A.), J. Dransfield (Royal Botanic
Gardens, Kew, U.K.), C. Kalkman (Rijksherbarium, Leiden, the Netherlands), T.C.
Whitmore (University of Cambridge, U.K.).
TREE FLORA
of
SABAH AND
SARAWAK
Volume One
edited by
E. Soepadmo and K.M. Wong
A joint publication of
Sabah Forestry
Department, Malaysia
Forest Research
Institute Malaysia
1995
electronic version prepared by
L. G. Saw & B. Aslina (2007)
Sarawak Forestry
Copyright Forest Research Institute Malaysia (FRIM),

Sabah Forestry Department, and Sarawak Forestry Department 1995
Forest Research Institute Malaysia (FRIM), Kepong, 52109 Kuala Lumpur, Malaysia
First published 1995
Perpustakaan Negara Malaysia Cataloguing-in-Publication Data

Tree flora of Sabah and Sarawak / edited by E. Soepadmo and
K. M. Wong.
Includes bibliography and index
ISBN 9839592343
1.Botany-Sabah. 2. Botany-Sarawak. 3. Botany-Borneo.
4. Botany-Classification. 5. Plants-Identification. I. Soepadmo, E.
II. Wong, K. M. (Khoon Meng).
581.095952
Electronic version by L. G. Saw & B. Aslina, Forest Research Institute

Malaysia, 2007.
CONTENTS
Page
VII
IX
Foreword
Acknowledgements
INTRODUCTION
Background to the Tree Flora of Sabah and Sarawak Project (E. Soepadmo)
A Brief History of Botanical Collecting and Documentation in Borneo (K.M. Wong)
Biogeography and Ecology (P.S. Ashton)
XIII
XXI
XLIII
THE TREE FLORA PROPER

1.
Aceraceae (A. Noorsiha)
2.
Alangiaceae (A. Berhaman)
3.
Anisophylleaceae (K.M. Wong & L. Madani)
4.
Araucariaceae (P.C. Yii)
5.
Bignoniaceae (A. Berhaman)
6.
Burseraceae (K.M. Kochummen)
7.
Capparaceae (Deborah Kennard)
8.
Celastraceae (K.M. Kochummen)
9.
Chrysobalanaceae (Ghillean T. Prance)
10.
Clethraceae (A. Berhaman)
11.
Connaraceae (Lesmy Tipot)
12.
Cornaceae (R.C.K. Chung)
13.
Datiscaceae (E.J.F. Campbell-Gasis)
14.
Goodeniaceae (K.M. Wong)
15.
Hypericaceae (K.M. Wong)
16.
Illiciaceae (Richard M.K. Saunders)
17.
Juglandaceae (E.J.F. Campbell-Gasis)
18.
Monimiaceae (P.C. Yii & Lesmy Tipot)
19.
Nyssaceae (P.C. Yii)
20.
Ochnaceae (K.M. Kochummen)
21.
Olacaceae (Lesmy Tipot)
22.
Oxalidaceae (R.C.K. Chung)
23.
Pittosporaceae (John B. Sugau)
24.
Rhamnaceae (Carsten Schirarend)
25.
Rhizophoraceae (L. Madani & K.M. Wong)
26.
Rutaceae (David T. Jones)
27.
Simaroubaceae (Julius Kulip & K.M. Wong)
28.
Sonneratiaceae (Othman Bojo)
29.
Staphyleaceae (J.T. Pereira)
30.
Styracaceae (R. Kiew)
31.
Trigoniaceae (K.M. Wong)
1
5
15
27
33
45
101
107
155
181
187
199
209
213
219
227
233
245
253
257
271
287
297
305
321
351
421
443
453
463
467
Abbreviations of Frequently Cited References

Commonly Used Abbreviations for Localities
Glossary
Index to Scientific Names
Index to Vernacular Names
471
473
474
487
506
FOREWORD
roviding a foreword to the first volume of any flora is always a rare and pleasurable
task, but at the same time a conscience-demanding one. In a region like Borneo,
where the Malaysian states of Sabah and Sarawak are located, and where a complete
documentation of the flora of any one sizeable area is not available, a beginning
volume of a new flora is an exciting endeavour. Yet the work to continue with further
volumes must press on, and we can only congratulate ourselves within that first limit.
The plant biodiversity of Sabah and Sarawak is a daunting topic. Estimates by specialists
throughout the decades vary but are in the region of 10,000 species of vascular plants for the
whole of Borneo. This is not likely to differ very much from the total for the habitat-rich and
topographically varied territories of Sabah and Sarawak. To engage enough specialists to
carry out the immense responsibilities of preparing such a flora is therefore conceivably a
hugely expensive and time-consuming task. To divide the work into workable components
is an equally complicated affair: how does one select certain families of importance, or
determine which revisions should await more expert attention at some later time?
The Tree Flora of Sabah and Sarawak addresses those plant families that contain
arborescent species for the mere fact that trees form the main framework of the rain forests
and are of a fundamental ecological and conservation importance. The basic idea has always
been to be able to identify these species, so that work in conservation, management and
sustainable use of species and products have a sound scientific basis, without which no
systematic applied work can proceed with ease. Our task is a major national effort with the
Forest Research Institute Malaysia, the Sabah Forestry Department and the Sarawak
Forestry Department as the key partners. We have the support of the universities and other
research agencies in Malaysia, as well as key institutions involved with Southeast Asian
botany within our region, in Australia, Japan, the United Kingdom, Europe and North
America, without which would have made the work extremely tedious and time-consuming.
The bulk of the funding for this enormous project comes from the Malaysian Government
(especially through its funds for the IRPA, or the Intensification of Research by Priority
Areas programme, disbursed to the participating institutions), the Overseas Development
Administration (ODA) of the United Kingdom, and the International Tropical Timber
Organization (ITTO). Such support has been crucial to the initiation of the Flora on an
organized basis and will be the essential fuel for continuing work. The main testimony to the
successful production of this first volume of the Flora, and the research now going on for
further volumes, however, remains the local, regional and international cooperation among
scientists, research managers and other resource people who have given time and
importance to this project. During the projected ten-year period of the project, a new
generation of tropical botanists would have developed; in this, which is of paramount
VII
significance to the continuing and future undertaking of conservation and forestry

management, the Tree Flora project would have served an enriching and catalystic role.
We would like to congratulate Prof. Soepadmo the project leader, and the scientists involved
in the project for their dedication and commitment, and acknowledge all national and
international agencies for their support. A ten-year project is a long-term project which
needs commitment and dedicated support and we hope that both elements will continue to
be present to take this project to its ultimate successful end.
Dato Dr. Salleh Mohd. Nor

Director-General
Forest Research Institute Malaysia
Haji Awang Tengah bin Haji Awang Amin
Director
Sabah Forestry Department, Malaysia
Datuk Leo Chai Chia Liang
Director
Sarawak Forestry Department, Malaysia
VIII
ACKNOWLEDGEMENTS
he successful initiation and continuation of the Tree Flora of Sabah and Sarawak
project owes much to the financial and technical support provided by the Malaysian
Government, the Overseas Development Administration (ODA) of the United
Kingdom and the International Tropical Timber Organization (ITTO). In particular,
the foresight and managerial skills of Dr. Jeff Burley and Dr. Philip Bacon (Oxford Forestry
Institute), and Dato Dr. B.C.Y. Freezailah and Dr. Efransjah (ITTO) have been essential to
the good progress made thus far.
Numerous other institutions and individuals have also contributed invaluable advice, time
and effort to this undertaking. Foremost in mind are our collaborating institutions in
Malaysia (the Universiti Kebangsaan Malaysia, Universiti Malaya, Universiti Malaysia
Sarawak, Universiti Pertanian Malaysia, Malaysian Agriculture Research and Development
Institute, Sabah Parks Department) and abroad (the Royal Botanic Gardens, Kew, U.K.; the
Oxford Forestry Institute, U.K.; the Rijksherbarium, Leiden, the Netherlands; the Royal
Botanic Garden, Edinburgh, U.K.; the Herbarium Bogoriense, Indonesia; the Singapore
Botanic Garden; the Arnold Arboretum of Harvard University, U.S.A.; the Institute of
Biological Sciences, University of Aarhus, Denmark; the Botanical Garden and Museum in
Berlin-Dahlem, Germany). We extend our gratitude also to colleagues who have specially
helped with facilities and research matters, at Kew (its director Prof. G.T. Prance, Prof.
G.L.Lucas, Dr. J. Dransfield, M.J.E. Coode, Dr. D. Kirkup, Diane Bridson, P. Bygrave, N.
Martland, A.M. Smith), Leiden (its director Prof. P. Baas, Dr. M. Roos, Dr. C. Kalkman, Dr.
J.F. Veldkamp, Dr. W.J.J.O. de Wilde, Dr. Ding Hou, Dr.M.M. van Balgooy, Dr. P. van
Welzen, Dr. P. Kessler, Dr. E.F. de Vogel, Stans Kofman, H. Lut), Harvard (Prof. P.S.
Ashton, Prof. P.F. Stevens), Oxford (Dr. P. Bacon, Dr. D. Filer, Prof. D.J. Mabberley),
Vienna (Dr. C. Puff) and the C.S.I.R.O., Australia (Dr. M. Dallwitz, Dr. T.G. Hartley). The
Keepers and Curators of the herbaria at Aarhus (A), the Arnold Arboretum (AA), BerlinDahlem (B), Bogor (BO), British Museum (Natural History) (BM), Oxford (FHO), the
Philippine National Herbarium (PNH), Sabah Parks, Sandakan (SAN), Singapore (SING),
Universiti Kebangsaan Malaysia (UKMB), and the Universiti Pertanian Malaysia have
assisted with loans of specimens, the provision of working facilities and information.
At the Forest Research Institute Malaysia, we thank Dato Dr. Salleh Mohd. Nor, Dr. Abd.
Razak Mohd. Ali, L.C. Cheah, Dr. Roslan Ismail, Dr. H.T. Chan, Dr. N. Manokaran, Dr.
L.G. Saw, K.M. Kochummen, R.C.K. Chung, Noorsiha Ayop, Lesmy Tipot, Rusea Go, Mat
Asri Ngah Sanah, Kamarudin M. Saleh, Lucy Sigamoney V. Rajoo, Shahani Saad, Khartini
Ahmad, and Jamaluddin Osman for assistance in many and different ways with the Project.
Acknowledgement is also made here of the help provided in Sabah by past and present
Directors of Forestry, Datuk M. Munang and Tuan Haji Awang Tengah bin Haji Amin, and
colleagues from the Forest Research Centre, Sandakan (Anuar Mohamed, R.C. Ong, Y.F.
Lee, A. Berhaman, J.T. Pereira, J.B. Sugau, Pung Vui Lee, J. Tangah, L. Madani, Dewol
Sundaling); and in Sarawak by its Director Datuk Leo Chai and colleagues from the
Sarawak Forestry Department (H.S. Lee, Abang Mohtar, Runi S. Pungga, P.C. Yii, Haji
Othman). We are also grateful to Professor Abdul Latiff Mohamad and Professor Ruth
IX
Kiew for their help and cooperation. Illustrators who have assisted with the bulk of
botanical drawings used in the first volume are Rosemary M. Wise (who conducted an
illustrations workshop in Sabah in 1995), C.L. Chan (who supervised a number of
illustrations), M. Fogherty, Jamal Hassim, Mohd. Akip, Mohd. Nizam Isa, J. Pao, Yap Pak
Hau and Zainal Mustaffa.
As work was being finalised for Volume One, Sarawak herbarium staff member Rena
George succumbed to malaria during field work, and long-time Sabah tree-climbers Kumin
Muroh and Tuyuk Gangou passed away. We do not forget the valuable service of these
colleagues, and that of many others who through the years have provided the backbone of
the plant-collecting effort based in the forestry departments of Sabah and Sarawak, and
which, like the efforts of scientific collectors of earlier times, are the essence of any
biodiversity inventory.
Not least must be acknowledged the contributors of revisions to Volume One, and those
who have consented to give assistance to, or provide chapters for future volumes. Theirs is
the hard work presented here.
E. Soepadmo
K.M. Wong
INTRODUCTION
Background to the Tree Flora of Sabah and Sarawak Project
E. Soepadmo
A Brief History of Botanical Collecting and Documentation in Borneo

K.M. Wong
Biogeography and Ecology

P.S. Ashton
TREE FLORA OF SABAH AND SARAWAK
BACKGROUND TO THE TREE FLORA

OF SABAH AND SARAWAK PROJECT
E. Soepadmo
Forest Research Institute Malaysia,
Kepong, Malaysia
Why a Tree Flora of Sabah and Sarawak?
orneo, the third largest island in the world, of which Sabah and Sarawak are parts,
has been frequently acknowledged as one of the most important centres of plant
diversity in the world. The island, which occupies a total land area of approximately
740,000 sq. km, is conservatively estimated to harbour 10,00012,000 species of
flowering plants, representing about 56% of the world total (Merrill 1950; van Steenis
1950; Kiew 1984; Mat-Salleh et al. 1992). Of these, 4050% are endemic to the island, and
up to 80% of the endemic species in Borneo occur in Sabah and Sarawak.
In certain localities in Sabah and Sarawak, where botanical exploration has been carried out
more intensively, the species diversity is indeed extremely high. Beaman & Beaman (1990),
for instance, have found that the flora of the Mt Kinabalu Park, Sabah, encompassing an
area of about 700 sq. km, contains not less than 4,000 species of vascular plants in 180
families and 980 genera.
For a number of economically important families and genera of flowering plants, Borneo
(Sabah and Sarawak in particular), is also known as the centre of distribution and species
diversity. For example, of the 386 known species of the Dipterocarpaceae, 291 species or
about 75% are recorded from Borneo, of which 257 species or about 66% occur in Sabah
and Sarawak. Of the 291 species occurring in Borneo, 156 or about 54% are endemic, of
which 59 species or about 20% are restricted to Sabah and Sarawak (Ashton 1982). In the
genus Durio (Bombacaceae), of the 30 species which have been described to date, 20
species or about 66% are recorded from Borneo, with 16 species or about 53% occurring in
Sabah and Sarawak (Kostermans 1958; 1990). Nine or about 45% of the 20 species known
from Borneo are endemic, and all occur in Sabah and Sarawak. Similarly, the genus
Mangifera (Anacardiaceae), comprising about 50 species, also has its centre of distribution
and species diversity in Borneo, where 20 indigenous species have been recorded (Bompard
& Kostermans 1992; Kostermans & Bompard 1993; Kochummen, pers. comm. 1995). Of
these, 18 species are known from Sabah and Sarawak.
For non-woody components of the flora, e.g., orchids and rafflesias, Borneo and Sabah and
Sarawak in particular, also house the most number of species in the Malesian
phytogeographic region. In this region, the orchid family or the Orchidaceae is known by
3,0004,000 species, representing 1216% of the entire flora (Chan et al. 1994). Of these,
XIII
TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)
Lamb (1991) has estimated that 2,5003,000 species are found in Borneo, equivalent to
about 75% of the Malesian orchid flora. A great number of the Bornean species have been
recorded from Sabah and Sarawak. The genus Rafflesia (Rafflesiaceae) with the largest
flower in the world, also has its centre of species diversity in Borneo where 56 of the 14
species known to date have been recorded, and all occur in Sabah and Sarawak (Meijer
1984; Mat-Salleh 1991; Nais 1992).
Other Old World tropical families which have their centres of distribution and species
diversity in Borneo include the Anacardiaceae, Burseraceae, Celastraceae, Clusiaceae
(Guttiferae; Calophyllum), Euphorbiaceae, Fagaceae, Myrtaceae (Syzygium or Eugenia),
Rhizophoraceae, and several others (Airy Shaw 1975; Ding Hou 1958, 1962, 1978;
Kochummen 1995; Leenhouts 1956; Merrill & Perry 1939; Soepadmo 1972; Stevens 1980).
The presence of high species diversity in the natural forests of Sabah and Sarawak also
means that there is a wealth of forest products to be harvested. There is no doubt that, in the
past few decades, the harvesting and utilisation of these forest products, e.g., tropical
hardwood timbers and rattans, has contributed significantly toward the socio-economic
development of these two eastern Malaysian states. In recognising that of late, the
exploitation and conservation of biodiversity in Sabah and Sarawak has become the focus of
international attention and scrutiny, and the need to strike an acceptable balance between
development and conservation of natural resources in these two states, it is imperative that
up-to-date botanical inventories should be carried out without further delay. Such basic
information is of paramount importance to the understanding of the availability, distribution,
ecological and conservation requirements, and economic potential of the plant resources.
Without such information, it will be extremely difficult if not impossible to develop and
manage the available resources on a sustained basis.
The need for a Flora of Borneo
he flora of a given region provides an inventory of plant species occurring in that

region, facilitates a means of species identification, and provides a source of
information pertaining to up-to-date taxonomic and conservation status, distribution,
ecological amplitude, and economic potential of the treated species. It is, therefore,
unfortunate to note that despite the widespread national and international recognition of the
great economic and conservation value of Bornean rain forests, and that botanical
exploration and collection in Borneo has begun as early as 1822, and botanical accounts of
its flora have appeared since 1894 (cf. Wong 1995 for more details), to date this species-rich
island has neither a comprehensive flora of any kind nor even a concise checklist of plant
species. For this reason Kiew (1984) has highlighted the need and urgency of producing a
simple Flora of Borneo in order to facilitate the implementation of sustainable
management practices of forest resources of this species-rich island. She argued that by
using Merrills (1921) and Masamunes (1942) enumerations as a basis, and by adopting a
pragmatic approach, a concise Identification Flora of Borneo can be readied within 10
years. However, due mainly to the lack of or insufficient institutional and financial support,
as well as shortage of qualified man-power, the proposed project never materialised.
XIV
BACKGROUND (SOEPADMO)
The Tree Flora of Sabah and Sarawak Project
n recognising the urgent need of producing a Flora of Sabah and Sarawak of any kind,
that recent collections (more than 200, 000 numbers from Sabah and Sarawak up to
1990) necessitate revision and updating of taxonomic accounts in the Flora Malesiana
itself, and that trained scientific personnel is now available at various national
institutions, the Tree Flora of Sabah and Sarawak Project was officially launched by the
Director-General of the Forest Research Institute Malaysia on the 18th of November 1991.
The modest 10-year project is executed jointly by the Forest Research Institute Malaysia
(FRIM), the Sabah Forestry Department (FD Sabah), and the Sarawak Forestry Department
(FD Sarawak) with the collaboration of other research institutions and universities. The
main objectives of the project are:
To document and update the taxonomic status of all tree-species (taken as reaching at
least 5 m in height, and 10 cm in diameter) native to Sabah and Sarawak.
To publish 8 volumes (each containing 300400 species) of a concise Tree Flora of

Sabah and Sarawak within 10 years.
To upgrade Malaysian capability and expertise in plant taxonomic research and the
survey and documentation of tree diversity.
To develop and strengthen the management capability of herbaria and their related data
bases in the three participating institutions (FRIM, FD Sabah and FD Sarawak).
For the first 5 years, the project is jointly funded by the Malaysian Government, the
Overseas Development Administration (ODA) of the United Kingdom, and the International
Tropical Timber Organization (ITTO).
To prepare the manuscripts of the estimated 109 families comprising about 3,000 treespecies, botanists of the following national and international institutions are taking part:
MalaysiaForest Research Institute Malaysia, FD Sabah, FD Sarawak, Universiti
Kebangsaan Malaysia (UKM), Universiti Malaya (UM), Universiti Malaysia Sarawak
(UNIMAS), Universiti Pertanian Malaysia (UPM), Mt Kinabalu Park, Malaysian
Agricultural Research and Development Institute (MARDI), and the WWF-Malaysia. In all
36 botanists are involved.
OverseasInstitute of Botany, University of Vienna, Austria; University of Brunei
Darussalam, Brunei; University of Aarhus, Denmark; Botanischer Garten und Botanisches
Museum, Berlin, Germany; University of Hong Kong, Hong Kong; Rijksherbarium Leiden,
the Netherlands; Singapore Botanic Garden, Singapore; Oxford Forestry Institute, Oxford,
United Kingdom; Royal Botanical Gardens, Kew, United Kingdom; Arnold Arboretum,
Harvard University, USA; University of Florida, USA; and Everglades National Park,
Florida, USA. A total of 21 botanists are involved.
Coverage. Being an identification type of flora intended to be user-friendly to nonspecialist readers, the Tree Flora will be written in simple and easy-to-understand English.
The use of highly technical botanical terms is, therefore, to be avoided.
XV
In the Tree Flora, all dicotyledonous trees, here defined as woody plants with the main
upright stems measuring not less than 5 m tall and 10 cm diameter, will be treated in full.
Non-tree and introduced tree species known only in cultivation will be given cursory
treatment and annotated in the keys only. For each tree-taxon (family, genus, species),
treatment will be confined to the following aspectscorrect/accepted scientific name, major
references, description of diagnostic characters, vernacular names (if applicable),
distribution, ecology, uses, notes on taxonomy (if applicable), and key(s) to lower rank taxa.
For each genus, at least one line-drawing depicting important vegetative and reproductive
characters will be provided.
Format. To standardise the format of manuscripts to be published in the Tree Flora, a set of
guidelines has been prepared by the Editors (cf. Soepadmo & Wong 1995). This guideline is
obtainable from the project Secretariat at FRIM. Among the main guidelines which should
be adhered to by contributors of manuscripts are the following:
NomenclatureNames of families and genera will normally follow Brummitt (1992),

but in cases of potential confusion and controversy, authors of revisions for the Tree
Flora are requested to confirm the accepted names with the Chief Editor.
ReferencesOnly selected references most relevant to the taxonomy and distribution

of the family, genus, and species and which are relevant to Borneo in general and Sabah
and Sarawak in particular will be cited. Unless listed in the Guidelines, book names
should be cited in full. Names of journals and other serials are to be cited following van
Steenis-Kruseman (1956).
Derivation of genus and species namesLanguage and meaning of root words to be

indicated.
Basionym and synonymBasionym to be given if it indicates a new taxonomic

perspective of the taxon being treated. Only synonyms relevant to Borneo are to be
included.
TypificationWhenever possible, citation of type specimen(s) of accepted species

should be provided.
DescriptionRequired for any family with more than one genus worldwide, for
genera, and for species, and should account for the following aspects: habit; leaf
arrangement and type; stipules; inflorescence type and position; flower sexuality,
symmetry, merism, details of fusion of parts; fruit type; seed, embryo, cotyledons,
endosperm, aril and others if very relevant.
Vernacular namesOnly names commonly used in Sabah and Sarawak will be

included. The dialect or language of each entry of vernacular name should be indicated.
DistributionNumber of genera and/or species is to be mentioned, followed by global
and Malesian distribution, and followed by that in Sabah and Sarawak. If the taxon also
occurs in Brunei and/or Kalimantan, it should be indicated following the distribution in
Sabah and Sarawak.
XVI
EcologyFor a family or a genus to be given only if general trends can be

summarised. For a species, information on forest type, soil/rock type, and altitudinal
range is required.
UsesFor a family or a genus to be provided only if there are general or interesting

uses. For a species as much information as possible should be given.
TaxonomyTo be provided only if there are taxonomic controversies regarding the

taxon. Brief commentary could be given if this will clarify the taxonomic status of the
taxon under consideration.
Key(s) to lower rank taxaBracket key structure is to be used. Couplets to be

numbered once only, the individual leads in the couplet not further numbered. In
constructing the keys, only diagnostic vegetative and/or reproductive characters are to
be used. Non-tree taxa are to be annotated only.
Infraspecific taxaIn cases where two or more infraspecific taxa exist in Sabah and
Sarawak, a full description of the species, including all variations known in Sabah and
Sarawak, should be given in the species description. This should be followed
immediately by a key to the taxa in which each infraspecific taxon is annotated.
In cases where only one of the infraspecific taxa is known to occur in Sabah and
Sarawak, the species description should follow the following format: species name and
authority, derivation of species name, reference for the species; infraspecific taxon
name and authority, reference, basionym, type specimen, synonyms, description of the
sole infraspecific taxon which represents the species in Sabah and Sarawak, vernacular
name(s), distribution, ecology, uses, and taxonomy.
Other project activities
part from preparing and publishing eight volumes of the Tree Flora of Sabah and
Sarawak, the project also carries out a number of activities relevant to its overall
objectives. These activities include:
Organisation of collecting expeditions to a number of botanically little-known localities

in Sabah and Sarawak.
Establishment of a data base for specimens and taxonomic references using BRAHMS
and related softwares.
Conducting workshops and specialised training on flora writing, documentation
software, botanical illustration, editing, curation and management of herbarium
specimens, etc.
Postgraduate training to upgrade Malaysian capability in plant taxonomic research and
the inventory and documentation of plant/tree diversity.
Collaboration with other institutions and agencies with similar aims in the Malesian
region.
XVII
References
Airy Shaw, H.K. 1975. The Euphorbiaceae of Borneo. Kew Bulletin Additional Series IV.
245p.
Ashton, P.S. 1982. Dipterocarpaceae. Flora Malesiana 1, 9(2): 237552.
Beaman, J.H. & R.S. Beaman. 1990. Diversity and distribution patterns in the flora of
Mount Kinabalu. In: P. Baas, K. Kalkman & R. Geesink (eds.), The Plant Diversity of
Malesia, pp. 147160. Kluwer Academic Publisher, Dordrecht, the Netherlands.
Bompard, J.M. & A.J.G.H. Kostermans. 1992. The genus Mangifera in Borneo: Results of a
IUCNWWF/IBPGR Project. In: Ghazally Ismail, Mustedza Mohamed & Siraj Omar
(eds.), Forest Biology and Conservation in Borneo, pp. 6171. Centre for Borneo Studies,
Publication No. 2.
Chan, C.L., A. Lamb, P.S. Shim & J.J. Wood. 1994. Orchids of Borneo. Vol. 1.
Introduction and a Selection of Species. Sabah Society & Royal Botanic Gardens, Kew.
xviii + 402 p.
Ding Hou. 1958. Rhizophoraceae. Flora Malesiana 1, 5(4): 429493.
Ding Hou. 1962. Celastraceae I. Flora Malesiana 1, 6(2): 277291.
Ding Hou. 1978. Anacardiaceae. Flora Malesiana 1, 8(3): 395548.
Kiew, R. 1984. Towards a Flora of Borneo. In: Ismail Sahid, Zainal Abidin A. Hasan, A.
Latiff Mohamed & A. Salam Babji (eds.), Research Priorities in Malaysian Biology, pp. 73
80. Penerbit Universiti Kebangsaan Malaysian, Bangi, Malaysia.
Kochummen, K.M. 1995. Burseraceae. In: E. Soepadmo & K.M. Wong (eds.), Tree Flora of
Sabah and Sarawak, Vol. 1: 45106. Celastraceae, ibid.:107154.
Kostermans, A.J.G.H. 1958. The genus Durio Adans. (Bombacaceae). Reinwardtia 4(3):
47153.
Kostermans, A.J.G.H. 1990. Durio bukitrayaensis Kosterm. (Bombacaceae), a new species
from Borneo. Botanica Helvetica 100/1: 2931.
Kostermans, A.J.G.H. & J.M. Bompard. 1993. The Mangoes. IBPGR/Linnean Society
London, Academic Press; xvi + 233.
Lamb, A. 1991. Orchids of Sabah and Sarawak. In: R. Kiew (ed.), The State of Nature
Conservation in Malaysia, pp. 78-88. Malayan Nature Society & IDRC, Canada.
Leenhouts, P.W. 1956. Burseraceae. Flora Malesiana 1, 5 (2): 209296.
Masamune, G. 1942. Enumeratio Phanerogamarum Bornearum. 739 p.
Mat-Salleh, K. 1991. RafflesiaMagnificent Flower of Sabah, 49 p. Borneo Publishing
Company, Kota Kinabalu, Sabah.
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Mat-Salleh, K., J.H. Beaman & H. Beaman. 1992. Specimen database and their utilization
for the Flora of Borneo. In: Ghazally Ismail, Murtedza Mohamed & Siraj Omar (eds.),
Forest biology and Conservation in Borneo, pp. 117137. Center for Borneo Studies, Publ.
No. 2.
Meijer, W. 1984. New species of Rafflesia (Rafflesiaceae). Blumea 30: 209215.
Merrill, E.D. 1921. A Bibliographic Enumeration of Bornean Plants. J. Str. Br. Roy. As.
Soc., Special Number. 637 p.
Merrill, E.D. 1950. A brief survey of the present status of Bornean botany. Webbia 7: 309
324.
Merrill, E.D. & L.M. Perry. 1939. The Myrtaceous genus Syzygium Gaertn. in Borneo.
Mem. Am. Acad. Arts & Sci. 18(3): 135202.
Nais, J. 1992. Distribution, dispersal and some notes on Rafflesia around Kinabalu,
Malaysia. In: Ghazally Ismail, Murtedza Mohamed & Siraj Omar (eds.), Forest Biology and
Conservation in Borneo, pp. 97108. Center for Borneo Studies, Publ. No. 2.
Soepadmo, E. 1972. Fagaceae. Flora Malesiana 1, 7(2): 165403.
Soepadmo, E. & K.M. Wong. 1995. Guide to Preparing and Editing Manuscripts. 31p.
Forest Research Institute Malaysia, FD Sabah and FD Sarawak, Malaysia.
Steenis, C.G.G.J. van. 1950. The delimitation of Malaysia and its main geographical
division. Flora Malesiana 1, 1: LXX LXXV.
Steenis-Kruseman, M.J. van. 1956. Citation of serial and some books. Flora Malesiana 1,
5(2): CXLVCLXV.
Stevens, P.F. 1980. A revision of the Old World species of Calophyllum (Guttiferae). J.
Arn. Arb. 61 (2 & 3): 117699.
Wong, K.M. 1995. A brief history of botanical collecting and documentation in Borneo. In:
E. Soepadmo & K.M. Wong (eds.), Tree Flora of Sabah and Sarawak, 1: XXIXLI.
XIX
A BRIEF HISTORY OF BOTANICAL

COLLECTING AND DOCUMENTATION
IN BORNEO
K.M. Wong
Forest Research Centre,
Sabah Forestry Department,
Sandakan, Malaysia
uch of the information on early botanical collections in Borneo is summarised, by

collectors names, in the great Cyclopaedia of Collectors for the Flora Malesiana
(van Steenis-Kruseman 1950, 1958, 1974). The present account attempts to collate
this information, together with some information on collectors beyond van
Steenis-Kruseman (1974) so that the progress of plant exploration in Borneo may be
followed through each period and in as chronological an order as possible. Collectors of
lesser significance, in that they have only gathered very few numbers, are not normally
listed here. We give importance here to collections made up to only around the beginning of
the 1980s, as these are often less immediately known to unfamiliar botanists, because they
are not held in herbaria in the region (through historical factors) or because they have rarely
been referred to in the literature. Many collectors, though not all and not in every instance,
since the 1950s, have distributed good sets of their collections to regional herbaria serving
areas where the collections have been taken, and in general there is a better chance that the
more recent collections made (especially as the 1970s and 1980s are approached) may now
be found deposited in local herbaria and are easily accessible for study locally. The period
beyond this is not covered in the present account as it can be expected that such modern
collections will be easily found for reference in either the local herbaria (Bogor, Kepong,
Kuala Lumpur, Sandakan, Sarawak, Singapore) or in the large botanical institutions abroad
that have a long-standing association with the study of Bornean plants.
Early collectors and documentation
he earliest significant efforts at plant collecting in Borneo were those of the early to
middle 19th century, and were motivated initially either by an interest to study or
collect groups of horticultural interest, or by the interest of specialists in Europe, Java
(where the Dutch administration of the Indonesian islands was based) and Malaya
and Singapore (the seat of English administration in the region then) to document interesting
and rare life forms of a little explored, faraway tropical land that was virtually terra
incognito as far as scientific documentation is concerned.
As early as 1822, George Mller, the Acting Resident in Dutch West Borneo, made
explorations and plant collections in the Kapuas valley and Pontianak, and around Kutei and
the Mahakam River until 1826, when he is thought to have been murdered near the upper
XXI
Kapuas. Mller collected for Carl Ludwig Blume, Director of the Buitenzorg (Bogor)
Botanic Garden around this time.
In 1836, Dutch botanist Pieter Willem Korthals, a member of the Commission for Natural
Sciences for the Dutch East Indies who had explored parts of Java and Sumatra, collected in
southeast Borneo, mainly in the Banjarmasin, Barito River and Mt Pamatton areas, together
with Salomon Mller (a German member of the Commission) and Ludwig Horner (Swiss
member of the Commission, who apparently did not collect plants on this expedition).
Merrill (1921) and van Steenis-Kruseman (1950) point out it is likely that some of
Korthalss Borneo records are probably of Sumatran or Javanese origin, owing to confusion
in labelling after the collecting. Korthalss collections are principally at the Leiden
herbarium. In 1842, the United States Exploring Expedition under the command of
Charles Wilkes, which began its journey in 1838 and had travelled through the Philippines,
visited the Banggi and Balabac Islands off the northernmost tip of Borneo, during which
expedition botanists William Rich and J.D. Brackenridge, with the help of zoologist C.
Pickering, made some plant collections (now principally with the U.S. National Herbarium
and the Gray Herbarium at Harvard). Between 1852 and 1855, Cornelis de Groot, Chief of
the Mining Department of the Dutch East Indies collected in southeast and east Borneo,
including the Banjarmasin, Tanjung Batu, Mahakam and Samarinda areas.
Among the more significant early collectors of Bornean plants must count also the Rajah
James Brooke, who had helped the Sultan of Brunei in suppressing a revolt in 1840 and was
consequently given Sarawak to rule in the following year. He was also made Governor of
Labuan and Consul General of Borneo in 1847, and collected some plant specimens which
the Kew herbarium acquired in 18531855. Hugh Low, appointed by the Rajah Brooke first
as his secretary in 1845 and later as Colonial Treasurer of Labuan between 1848 and 1877,
and who made the first documented climb by a European to the summit of Kinabalu in
1851, was well known even at the time for his interest in orchids. Lows successful third trip
to Kinabalu in 1858, together with Spenser St. John, secretary to the Rajah Brooke,
included the first collections from Marai Parai, the ultramafic western spur of the mountain.
Low had also collected in the Kuching area, Mt Penrissen and Lawas. Low sent most of his
orchid specimens to John Lindley at the London University and Joseph Hooker at Kew.
Lows plant-collecting contemporaries included James Motley, a civil engineer at the
Labuan coal mine during 18511854 and who was later (18541859) at Banjarmasin in
south Borneo (Motleys Labuan collections were often sent to Kew by E. Barber Scott
labelled coll. Barber, Labuan, although his Kalimantan collections were distributed under
his own name), and Thomas Lobb, who made an unsuccessful attempt to reach the summit
of Kinabalu in 1856. Lobb, who was collecting plants of horticultural value for the firm of
Messrs Veitch in England, also collected some herbarium specimens from Sarawak and
Labuan. Lobb is thought to have intentionally falsified localities on some labels in order to
increase the value of his collections for the firm of Veitch; a specimen labelled Luzon in
one herbarium may be labelled Borneo or Singapore or Java in another (van SteenisKruseman 1950). The well-known zoogeographer Alfred Russel Wallace visited and
collected mainly ferns in Sarawak (Sarawak, Simunjon and Sadong Rivers) during
18541856, at the start of his travels through the Malay Archipelago.
XXII
BRIEF HISTORY (WONG)
Willem Hendrik de Vriese, a botanist at Leiden at some time during this period, collected in
Dutch West Borneo in 1860. Eduard von Martens, Custodian of the Berlin Zoological
Museum, also made plant collections in Dutch West Borneo in 1863. Between 1865 and
1867, Odoardo Beccari, the well known Italian naturalist, made many plant and animal
collections in Sarawak; many of his collections were made from around Kuching and
Matang, and the Sarawak River and Batang Lupar. He also collected from the Bintulu area
and the Rejang valley, and was also briefly in Labuan, Brunei and the then Dutch West
Borneo. His plant collections number around 20,000; he wrote many papers, especially on
palms. Perhaps his most well known book is Nelle Foreste di Borneo (Wanderings in the
Great Forests of Borneo) (Beccari 1902). Burtt (1964) points out that although Beccari
ascended the Poi range, he did not climb Mt Poi (Poe or Pueh) as that name is used on
modern maps, but a more south-easterly peak called Mt Berumput. Somewhat less
illustrious than Beccari but nevertheless a collector who during 18661869 sent several
cases of important plant specimens from Sarawak and Sambas in Dutch West Borneo to the
herbarium at Buitenzorg was a man documented only as a coffee planter in Java and
manager of an estate in Sarawak, called Martin.
In 18741875, the well-known botanical explorer Johannes Elias Teijsmann (also Curator
of the Buitenzorg Botanic Gardens in 18311869) explored Dutch West Borneo, making
numerous collections in the Pontianak, Kapuas River, Kenepei River, Penein Mts, Biang
Mts and other areas. His Borneo collections are mainly in the Bogor, Leiden, Kew and
Florence herbaria. In 18811884 Friederich Grabowsky, a German zoologist, collected a
few hundred plant specimens from southeast Borneo, mainly around the Banjarmasin and
Kapuas River areas. His collections are principally at the Berlin and British Museum
herbaria. During 18791880, the Danish explorer, ethnographer and zoological collector
Carl Bock made an expedition to northeast and southeast Borneo, collecting also some
plants along the Mahakam and Telen and in the Samarinda and Banjarmasin areas.
Another important plant collector about this time was Frederick William Burbidge, a
collector for the Veitch nurseries in England, who collected in Labuan and Kinabalu with
Peter Veitch in 1877 and in the Sandakan area with William Pryer, the founder of
Sandakan, in 1878. He also collected along the Lawas, Meropok, Limbang and Pandaruan
rivers in the northern part of Sarawak. Burbidges botanical plates and sketches from
Borneo are housed in the British Museum of Natural History. In 1880, two other Veitch
collectors, David Burke and Charles Curtis collected in Sarawak; Curtis later (1884)
became the Superintendent of Gardens and Forests in Penang.
The Hose family, comprising of the Reverend George Frederick Hose (Bishop of Singapore,
Labuan and Sarawak, 18811908), his nephews Charles Hose (district officer in the Baram
in 1884 and afterwards, Sibu) and Ernest Hose (a planter), and his daughter Gertrude
Hose all collected plants in Sarawak. The Rev. Hose collected on Matang and (with Charles
and Alfred Hart Everett, freshly retired Resident of Trusan and Honble of the 4th
Division) on Mt Dulit, and other places, taking especially grasses, sedges and ferns, as his
daughter did. Charles Hose made zoological studies as well as collected plants, in the
Baram, Mt Dulit, Mt Mulud, Kayang, Rejang and Niah areas. Their collections were sent
mainly to the Kew and Edinburgh herbaria.
XXIII
Between 1883 and 1885 when he was murdered, M. Fraser, Medical Officer of the Chartered
Company in North Borneo, collected several hundred plant specimens from Kudat, the
Marudu Bay, Balambangan and Banggi Islands, Gaya Island and Papar. These specimens
are at Kew. Around 1885 and 1890, a Singapore schoolmaster, R. Hullett visited Sarawak
and collected several hundred specimens, now mostly in the Singapore and Kew herbaria. In
1886, Ignatz F. Frstermann, in the employ of the English horticultural firm of Sander &
Sons, collected ferns and orchids in Sarawak. John Whitehead, an ornithologist, collected a
small number of plants during his famous expedition to Kinabalu in 18871888.
In 1892, George D. Haviland, Curator of the Sarawak Museum at Kuching, visited Kinabalu
(with his brother H.A. Haviland) and collected about 450 numbers on its upper slopes. Both
Burbidges and Havilands collections were studied by Otto Stapf of the Royal Botanic
Gardens, Kew, who wrote the first monograph on the Kinabalu flora (Stapf 1894),
enumerating 360 species of flowering plants, ferns and bryophytes. Haviland also collected
at Kuching, Mt Braang, Mt Penrissen, Limbang, Sibu, Ulu Tawaran, and (with Charles
Hose) Mt Lambia. van Steenis-Kruseman (1954) has compiled a list of Havilands
collections in Borneo as then known. Edward Bartlett, a zoologist who also collected a
small number of plant specimens in Sarawak around 1893, later succeeded Haviland in
1895. Robert W.C. Shelford, who in 1897 was the Curator of the Sarawak Museum,
collected Sarawak plants between 1897 and 1903, mainly in the Kuching and Trusan areas.
Charles van de Leur Creagh, the Governor of North Borneo during 18881895, collected at
least 1839 numbers on Kinabalu and also made collections from Pulau Gaya, Kimanis and
Labuan on the west coast to Cowie Harbour, Tawau and Sandakan on the east coast (Meijer
1969). Johannes Waterstradt, Danish zoological collector, collected mainly orchids as
well as insect specimens on his trips in Borneo. He went to the Padas River (North Borneo)
in 1891, the mouth of the Lawas River and Kinabalu (18921893), Kudat, Banggi,
Balambangan, Labuan , Kinabalu (N Borneo) and the Lawas and Limbang Rivers (Sarawak)
(1894), Kinabalu (1895), Brunei (1902), and again Kinabalu (1903, 1908 and 1912). Much
of his orchid collections before his return to Europe in 1904 went to the firm of Hugh Low
in London.
This was the time when the great Dutch explorer and ethnologist, Anton Willem
Nieuwenhuis, made his expeditions into Dutch Borneo, exploring with German botanist
Johann Gottfried Hallier (then an assistant at the Buitenzorg herbarium; he also used the
initial H for Hans) and Dutch geologist Gustaaf Frederik Molengraaff in 18931894 the
Mandai, Kenepai and Mendalam Rivers, the Kapuas delta and the Pontianak area; with
Indonesian plant collector Jaheri (of the Buitenzorg Botanical Garden) in 18961897 the
Mendalam and Mahakam areas; with Indonesian collectors Amdjah and Sakaran (of the
Buitenzorg Botanical Garden) in 18981899 the Kapuas, Mahakam and Samarinda areas;
and in 18991900 the Apo Kayan highlands. These collections (several thousand numbers)
are principally at the Bogor and Leiden herbaria. Amdjah collected again in northeast
Borneo in 1912.
Early 20th Century collectors
.N. Ridley, Director of Gardens and Forests, Straits Settlements during 18881900,
and Director of the Singapore Botanic Garden during 19011912, visited and
collected in Bau, Matang and Lundu in Sarawak in 1893. In 1897 he collected in
XXIV
Labuan, Kudat, Sandakan and the Labuk Bay area, and in Sarawak again. His subsequent
visits to Sarawak were in 1903, 1905 and 1915. Ridleys collections are principally at the
Kew and Singapore herbaria. John Hewitt, who became Curator of the Sarawak Museum
during 19051908, collected plants mainly in the Kuching, Saribas, Baram and Limbang
areas.
The German orchid specialist Friedrich R.R. Schlechter collected in Labuan, Kudat,
Sandakan in British North Borneo, and Banjarmasin, Balikpapan, Samarinda and the Kutei
region in Dutch Borneo in 1901, and later in Kuching in Sarawak during 19061907. H.
Witkamp, a mining engineer with the Royal Batavian Oil Company, made plant collections in
Kutei district in east Borneo in 1905, 1907, 1910 and 1928.
Around this time, Cecil J. Brooks, a chemist with the Borneo Company which exploited
gold in Sarawak, collected plants, mainly ferns, along the Sarawak River and on Mts
Santubong and Poe, and around Bau and the Bungo Range, between 1907 and 1910.
In 1908 the German botanist Hubert Winkler collected more than a thousand numbers
during his trip to east and southeast Borneo, covering the Banjarmasin, Barito River,
Balikpapan, Samarinda and Mahakam areas. His collections are principally at the herbaria at
Breslau, Berlin, Geneva and Manila (the last destroyed in the Second World War). During
19101914, the Dutch geologist Louis Martin Robert Rutten collected plants in east and
northeast Borneo in the Balikpapan and Samarinda areas, along the Kayan and Rapah
Rivers, and in the Sangkulirang Bay area.
Lilian Suzette Gibbs, an English botanist interested in tropical alpine floras, made
collections in the Beaufort area and the Tambunan plain, and also collected on Mount
Kinabalu in 1910 (she collected on the Gurulau and Marai Parai spurs and ascended to the
summit via Kamborangoh). Gibbs collected about 1000 numbers from North Borneo. Her
paper on Kinabalu (Gibbs 1914) was the second monograph on the Kinabalu flora after
Stapfs. The American botanist Frederick William Foxworthy, serving at the Bureau of
Science in Manila (19061911), collected in the Lundu, Niah, Mt Poe, Mt Santubong and
Kuching areas in Sarawak in 1908 and also climbed Kinabalu in 1910. John Coney
Moulton, Curator of the Sarawak Museum during 19051915, collected widely in Sarawak
between 1909 and 1920, and on Kinabalu in 1913. Moultons collections are largely in the
Edinburgh, Kew and Singapore herbaria. Sarawaks first Conservator of Forests in 1919,
John Phillips Mead, also collected plant specimens. Eric P. Mjberg collected in Mt
Murud, Mt Dulit, Mt Poe, the Bidi Caves, Mt Matang, Mt Penrissen and other areas in
Sarawak when he was Director of the Sarawak Museum in 19221924, and afterwards in
Dutch East Borneo and Southeast Borneo in 19251926. Forest Ranger D. Carroll collected
several hundred specimens, mainly from the Kuching, Lundu and Sadong areas in Sarawak
between 1922 and 1953.
The Forest Research Institute, Buitenzorg began sending its collectors to Dutch
(Indonesian) Borneo since about 1913. Johan Philip Pfeiffer, a Dutch chemical engineer,
made botanical collections in 1913 in south and southeast Borneo (in the Sampit and
Martapura areas), in 19151916 in northeast Borneo (in the Telok Selimau, Tanjung
Mangkalihat and Berouw River areas) and again in 1918 in northeast Borneo (Telok
Selimau); his collections are mainly at the Bogor and Leiden herbaria.
XXV
Johan Frederik Labohm, Forest Officer in southeast Borneo, collected many plant
specimens in the Banjarmasin, Sampit, Balikpapan and Samarinda areas during 19161920.
At this time (19171925), a Forest Overseer in the same district, Carel Nicolaas Johan
Delmaar, also collected many plant specimens from the Kuala Kapuas area. Mohamad
Dachlan, another overseer in southeast Borneo, too, made several hundred collections
around the Banjarmasin, Martapura and Kuala Kapuas areas between 1918 and 1939.
Frederik Hendrik Hildebrand of the Buitenzorg Forest Research Institute made some small
collections of plants in 1925 and 1928 in southeast Borneo. Another Forest Officer, Laurens
Verhoef, also collected in southeast Borneo in 19281929.
Joseph Clemens, a chaplain in the U.S. army, and his wife Mary Strong Clemens made
substantial collections during several trips to Kinabalu between 1915 and 1917 (collecting at
least 1,839 numbers) and between 1931 and 1933 (making more than 7,000 collections).
Most of these have been sent to the Arnold Arboretum in the U.S.A. and the British
Museum of Natural History. Their specimens were worked on by Merrill and others, and
subsequently enumerated by H. Heine (1951, 1953). David L. Topping, a government
official in the Philippines, collected more than 500 fern specimens on Kinabalu together
with Mary Clemens in 1915. A smaller collection of about a hundred numbers was also
made on Kinabalu by George A.G. Haslam in 1916.
The Chartered Company which administered British North Borneo formed its Forestry
Department based in Sandakan, with American forester D.M. Matthews as the first
conservator in 1914, who had an interest in Philippine dipterocarps. The North Borneo
Forest Department started botanical collecting around 1915 (when its collector Aniceto
Villamil was joined by Foxworthy to make the first collections for the Sandakan
herbarium), and sent its dipterocarp specimens to the Bureau of Science in Manila for study
by Foxworthy, who also served as advisor to the Forestry Department in North Borneo
during 19151916. In 1916, the Forest Department had obtained collections representing
537 species. In 1916 also, the first bulletin of the forest department, on the timbers and
minor forest products of North Borneo, was published (Foxworthy 1916). Another
American, Devillo D. Wood, transferred from the Bureau of Forestry in Manila in 1916 and
who became conservator in North Borneo in 1917, began encouraging active botanical
collecting in the east coast lowlands. The Forestry Department maintained close links with
its counterparts in the Philippines and sent plant specimens to the Bureau of Science in
Manila for identification. In 1920 Maximo Ramos, a trained collector borrowed from the
Bureau of Science in Manila, collected 1256 specimens in Sabah, mostly in the lowlands
around Sandakan. By 1921 Woods own collections had numbered at least 2555 and in
1922 together with Jose Agama, a Filipino forest ranger who joined the British North
Borneo forestry service in 1915, he collected on Balambangan Island off the north coast. In
1922 he sent forest officers P. Castro and F. Melegrito to collect on Balambangan and the
nearby Banggi island. Other forest officers who took part in this collecting effort during this
time were L. Apostol, P. Orolfo and Aniceto Villamil. These collections, and those of the
Clemenses from Kinabalu and of Hose from Sarawak, were the principal basis of a number
of botanical papers on the North Bornean flora by Elmer Drew Merrill, first a botanist and
later the director at the Bureau of Science in Manila (Merrill 1916, 1917a, b, 1918, 1922a, b,
1924, 1926). Merrill, who published a bibliographic enumeration of Bornean plants (Merrill
1921), also studied the collections of Adolph Daniel Edward Elmer, a botanist and collector
XXVI
for the Bureau of Science, who collected in the Sandakan and Labuk Bay area in 1921, and
the Tawau area in 19221923 (about 1,523 numbers) (Merrill 1929). The zoologist C.B.
Kloss, of the Federated Malay States Museum in Taiping, also visited the Sandakan area
and Balambangan and Banggi Islands in 1927 and Kinabalu in 1928, and collected a few
plant specimens. Merrill left for the University of California at Berkeley in 1924, where
many duplicates from North Borneo (including those of Elmer) were then sent, and moved
to the New York Botanical Garden in 1930.
The German botanist Hans Winkler, Director of the Botanical Garden at Hamburg,
collected in Dutch West and Central Borneo in 19241925; these collections are mainly in
the Hamburg, New York, Leiden and Bogor herbaria. Frederik Hendrik Endert of the
Buitenzorg Forest Research Institute made collecting expeditions to Dutch Borneo in 1925
(around Kutei in northeast Borneo), 1928 (Tanah Blomboe in southeast Borneo) and 1938
(Sanggau, Sambas and Paloh areas in West Borneo). The 1928 expedition was made
together with Dirk Fok van Slooten, an assistant of the Buitenzorg herbarium. Enderts
several thousand Borneo collections are principally at the Forest Research Institute in
Bogor, while the 200-odd collections of van Slooten are at the Bogor herbarium.
Between 1929 and 1932 Jan Pieter Schuitemaker, Forest Officer in Dutch West Borneo,
made a number of collections there. The Dutch botanist, Oene Posthumus, collected in the
Samarinda, Balikpapan and Tengarong areas in east Borneo in 1930; the collections, mostly
of ferns, are mainly in the Bogor herbarium. In 19311933, the Assistant Resident of Dutch
West Borneo, Louis Coomans de Ruiter likewise collected plant specimens there. Betje
Polak, temporarily attached to the Buitenzorg Botanic Gardens and later the Buitenzorg Soil
Science Institute, collected plant specimens in West Borneo (around Pontianak) in 1930,
southeast Borneo (around Banjarmasin) in 1939 and West Borneo in 1940.
Richard Eric Holttum, Director of the Singapore Botanic Garden, made a collection of
about 300 ferns, mosses and other plants on Kinabalu in 1931 together with the Clemenses.
In 1932, also together with the Clemenses, Caetano Xavier Furtado (Assistant Curator in
the Straits Settlements Gardens Department and a specialist in palms) collected on Kinabalu.
In 1933, Cedric Errol Carr, a retired rubber planter in Malaya, collected about 2,000
numbers (including 700 orchids) on Kinabalu. Carrs collections are in the Singapore
herbarium.
The next conservator in North Borneo, Henry George Keith, who assumed headship of the
Forest Department in 1931, continued to encourage botanical collecting and also collected
plant specimens himself after his arrival in North Borneo in 1925. Keith was interested in
pursuing a foresters flora and also compiled a preliminary list of North Borneo plant names
(Keith 1937); he based much of his work on identifications provided by H.K. Airy Shaw at
the Kew herbarium. Many of the duplicates of collections at this time were despatched to
the Kew, Singapore and Kepong herbaria, the last where dipterocarp specimens were identified
by the Forest Botanist of Malaya, Colin Fraser Symington (who visited the Kabili-Sepilok
Forest Reserve, Semporna and the Betotan and Segaliud areas in North Borneo, and Brunei
in 1938). By 1940, the Sandakan Herbarium had amassed 10,803 sheets.
During the Oxford University Expedition to Sarawak of 19321933, its botanist, Paul
Westmacott Richards, made about 2,800 collections, from Miri up the Baram to Marudi,
XXVII
the Dulit range, Bidi, Bau and Santubong. The specimens are principally at the Kew,
Oxford, British Museum of Natural History, Singapore and Sarawak herbaria. Patrick M.
Synge, who assisted Richards but fell ill, collected principally orchids and pitcher plants. At
about this time, C.O. Flemmich, who was appointed State Forest Officer in Brunei made a
substantial collection from trees in Brunei, which are mainly at the Kepong herbarium.
In 1937, the American ornithological collector John Augustus Griswold collected over a
hundred plants on Kinabalu during the Asiatic Primate Expedition of Harvard to Siam and
Borneo. Expedition leader H.J. Coolidge, who never went up Kinabalu, is sometimes
erroneously credited as having collected Kinabalu plants with Griswold.
In 1938, the Japanese Nobuhira Hanada collected specimens of woody climbers of the
Menispermaceae mainly from the Kuching area. He was probably assisting botanist
Yoshimatsu Yamamoto who was a specialist in the family and who collected in Tawau in
North Borneo and Pontianak and Banjarmasin in West and South Borneo in 1939.
In 1940, Pieter Buwalda of the Buitenzorg Forest Research Institute collected more than
300 numbers from the Sampit region in south Borneo.
Post-War Collectors
hen the Second World War intervened, Australian air raids on Sandakan in 1944
destroyed the herbarium. Still with Keith as conservator, the herbariums
collection programme resumed in 1947, assisted by a return of some 1,000 sheets
of specimen duplicates from the Singapore Herbarium. In the same year Keith
(1947) published his account of North Borneo timbers. Dipterocarp specimens, in particular,
which had been sent to Kepong for study and identification by Symington, were, after his
death in 1943, also sent to van Slooten in Bogor and later Amsterdam, until van Slooten also
died in 1953.
Java-born Dutchman Andre J.G.H. Kostermans, of the Botanical Division of the
Buitenzorg Forest Research Institute, made a number of expeditions to Borneo around this
time: in 1948 and 1953 around Sampit in south Borneo; in 1951, 1952, 19531954, 1955
and 1957 (and also later in 1963) in northeast and east Borneo; and in 1955 and 1956 in
central Borneo. Kostermans 1952 and 1953 trips were partly joined by Willem Meijer, an
assistant in the Bogor Herbarium then, and later in 1955 a lecturer at the Faculty of
Agriculture in Pajakumbuh in Sumatra before his repatriation in 1958. Anne Johnson,
demonstrator in botany at the University of Malaya in Singapore, collected some Sarawak
plants, mainly bryophytes and ferns, in 1951; these are found in the Singapore herbarium.
James Sinclair, Curator of the Singapore Herbarium, collected around Santubong in
Sarawak in 1949, on Kinabalu and around Sandakan in 1957, and in Sarawak and Brunei in
1960. In 1955, John William Purseglove, Director of the Singapore Botanic Garden,
collected in the 1st Division of Sarawak (with Mohamad Shah of the Singapore
Herbarium), and again in the Bako National Park in 1956 (with Hamish Boyd Gilliland of
the University of Malaya and Mohamad Shah). George Alphonso, Curator of the Singapore
Botanic Gardens, also collected in Sabah (on the Crocker range, near Tenom, Tambunan
and Patau, and around Keningau) in 1959; his further trips to Borneo (1965 to Sarawak,
XXVIII
with Samsuri Ahmad; 1966 to Kinabalu) did not yield many more herbarium specimens.
Hsuan Keng, botany lecturer at the University of Malaya in Singapore, made a short
collecting trip to Sarawak (Baram district) in 1961.
In Sarawak, James Aidan Robb Anderson, who became Assistant Conservator during
19511954 and Forest Research Officer in Kuching in 1955 before transfer to Brunei as
State Forest Officer in 1956, and back to Sarawak later as Forest Research Officer, made
substantial collections, especially of peat-swamp plants in the Rejang, Baram and
Kayangeran areas (Sarawak) and Badas (Brunei). His collections are mainly at the Sarawak
and Kepong herbaria. Anderson published a guide to the peat-swamp flora (Anderson
1963). At this time Francis George Browne was the Conservator, and Browne also made
some plant collections besides writing a compendium (Browne 1955) of common forest
trees. Andersons successor as Assistant Conservator, and later also State Forest Officer in
Brunei, was a German forester, F.W.O. Eberhard Brnig, who collected especially the flora
of podzolized soils in the 1st and 2nd Divisions in Sarawak and Brunei (Brunig 1968).
During the Oxford University Expedition to Borneo, 19551956, led by G. Arnold, forester
Gordon Harold Pickles collected plant specimens (lodged principally at Forest Herbarium,
Oxford, Singapore and Kuching) in the Plieran Valley along the Rejang River, the Usun
Apau Plateau, Mt Kalulong and along the Baram River.
In 1952, Keith was succeeded as conservator in North Borneo by A.B. Walton from the
Malayan forestry service, who continued to emphasize floristic inventory work. Dipterocarp
collection and inventory continued to dominate botanical work based at the forest
department when, upon the recommendation of John Wyatt-Smith (Forest Botanist of the
Federation of Malaya, at Kepong), Geoffrey Howorth Spencer Wood was appointed the
first Forest Botanist for North Borneo in 1954, when G.L. Carson succeeded Walton as
conservator. In the same year Wyatt-Smith and Wood collected in North Borneo together,
and afterwards Wood spearheaded the departments botanical collecting and began to
specialize in the dipterocarps. After a years home leave to Britain, Wood returned in 1957
and went on a collecting trip to Brunei with P.S. Ashton, Bruneis newly arrived and first
Forest Botanist. Wood died in Brunei from an accidental explosion in camp. Between 1948
and 1958, John Kidman Cox of the North Borneo Agriculture Department collected mainly
orchids there (including on Kinabalu); he made about 900 collections from North Borneo,
which were sent to Singapore, Kew, the British Museum, Berkeley and Leiden.
Bertram Evelyn Smythies, State Forest Officer in Brunei in 19521959 and thereafter
Conservator of Forests in Sarawak until 1964, when he joined the Royal Society Expedition
to Kinabalu for a short period, also added substantially to the botanical collections. He
published a simple guide to common Sarawak trees from 34 families (Smythies 1965). Peter
Frederick Burgess, Deputy Conservator at Sandakan during 19561965 (when North
Borneo became Sabah), and who wrote an excellent account of the timbers (Burgess 1966),
also made small numbers of collections, especially in the east coast areas.
Timothy Charles Whitmore collected in Sarawak (around Kuching), Brunei and North
Borneo (Labuan, Lungmanis, Sandakan, Kalabakan, Tawau areas) in 1957 when principally
studying dipterocarps in the field; he returned when serving as Colombo Plan botanist at
Kepong to collect in Sarawak (at Semengoh) in 1966. Whitmores Borneo collections are
mainly at Cambridge, Sandakan, Singapore and Leiden. During this period, Marius Jacobs
XXIX
of the Bogor herbarium made collecting trips to Sarawak, Brunei and Kinabalu in North
Borneo in 1958, taking nearly 800 numbers. University of Malaya plant physiologist John
Carrick collected in Sarawak (Bako) in 1959 (with I.C. Enoch) and 1961; his collections,
about 600, are in the university herbarium (Kuala Lumpur) and in the herbaria in Singapore
and Kuching.
A successor to Wood who would want to travel out to North Borneo was not easily found,
and the job was finally given in 1959 to Meijer, who had just been repatriated from
Sumatra. Meijer continued working on dipterocarps, building on manuscript notes left by
Wood. Shortly after Meijers arrival, the wooden Sandakan Herbarium building was razed
by fire that spread from an adjacent veneer factory in 1961, when almost all its 15,000
mounted specimens were destroyed. In 1961, Dan H. Nicholson, then a student at Cornell
University, collected mainly aroids in Sarawak (Semengoh, Matang, Bau, Bako, Setapok).
Peter Shaw Ashton, who left Brunei in 1961, became Forest Botanist in Sarawak between
1962 and 1966. Although there were suggestions for a herbarium in the Sarawak Forest
Department in 1947, a combined Sarawak Museum and Forest Department herbarium came
into being only by 1959, and Sarawak Museum specimens (including Haviland collections)
were then incorporated into a new herbarium in its own building in 1961. Some specimens
collected by Haviland, Hose and the Clemenses from the Brunei, Sandakan and Singapore
herbaria were also returned to the Sarawak herbarium. Ashtons numerous collections of
Brunei plants (as well as those of Hasan Pukul, with whom Ashton sometimes collected)
and Sarawak plants are mainly at the Sarawak, Brunei, Kew and Kepong herbaria. He
collected again in Brunei in 1965. Ashton wrote a manual to Brunei dipterocarps (Ashton
1964), published the same year as the one by Wood & Meijer (1964) for Sabah, followed by
a supplement (Ashton 1968), and also completed by 1973 the manuscript for a book on 25
non-dipterocarp families in Sarawak, which publication was delayed until 1988 (Ashton
1988). Based at the Sarawak herbarium, collectors of importance include herbarium
assistant Ilias Paie (from 1962, active until the 1980s), tree climber Banyeng anak
Nyudong, together with Ardzi Arshid, Bojeng Sitam, Galau, and Rashid Taggoi.
In Sandakan, the 1960s saw a new drive in collecting with herbarium assistants Leopold
Madani (from 1960 and still collecting in the 1990s) and Aban Gibot (from 1962, active
until the 1980s); there, others with substantial collections to their credit include George
Mikil, J. Ampuria, James Ah Wing, David Brand, Jaswir Singh, Muin Chai, David
Charington, Abdul Rahim, Masirom Rundi, and Henry Sinanggul. Through the massive
recollection programme some 17,200 specimens (including about 13,600 post-1961
collections) had been accessioned in 1964, when a new herbarium building was ready
(Meijer 1964). Meijer, Aban and Madani also collected with both Royal Society expeditions
to Kinabalu (1961, 1964), but used the SAN number series.
Interest in the Kinabalu flora had not waned during this period. Although there were very
few who collected frequently on the mountain apart from Forest Department staff, there was
also some notable exploration carried out, such as the exploration by Sheila Collenette
(formerly Iris Sheila Darnton) of Kinabalu from Kundasang over the Pinosuk Plateau in
1960. Her collections, at first largely within the Kota Belud area where she was
headquartered in 1954, and later more generally in North Borneo, number nearly 2,000 and
are mainly with the herbaria of the British Museum of Natural History and at Leiden.
University of Minnesota professor Ernst Cleveland Abbe and his wife Lucy B. Abbe
XXX
collected during 19591960 in Kuching, Brunei and (accompanied by his research assistants
Linn Bogle and Robert Bruce Kaul) on Kinabalu (which the Abbes revisited in 1962) and in
1964 in Sarawak; their collections, many of oaks, are mainly in the herbaria at the Arnold
Arboretum, Kew, Edinburgh, the University of Minnesota and Singapore.
With the encouragement of Carson, Edred John Henry Corner from the University of
Cambridge led two Royal Society expeditions to Kinabalu, in 1961 (Corner 1964) and 1964.
Corner was Assistant Director of the Singapore Botanic Garden during 19291941. The
other principal botanical collectors on these expeditions, which represented an important
advance in knowledge for one of the richest floras in the tropics, were John David Adam
Stainton, a barrister-at-law and assistant organizer of the expedition (in 1961) and Chew
Wee Lek (both expeditions), first botanist and later Keeper of the Singapore herbarium.
Chew also collected in Sarawak in 1962 (Niah, Baram, Mt Mulu, Mt Api and Mt Benarat),
1963 (the Bau limestones, Mt Gading, Priam River, Mt Mentawa, Mt Maja), 1966 (Lambir,
Marudi, Baram, Limbang, and the Tutoh and Melinau Rivers) and 1967 (Mt Api, Mt
Benarat, the Tiang Bekap limestones near Kuching, Mt Mentawa and Mt Maja). Corner had
previously collected in Brunei and Sarawak (Bako National Park) in 1959, and also
collected in the vicinity of Kuching in Sarawak after the first expedition to Kinabalu in
1961. These collections are principally at the Kew herbarium; Chews other collections
were lodged mainly with the Singapore herbarium, and duplicated at the Leiden, Kew and
the Arnold Arboretum herbaria. Kinabalu collections were also made during the 1964
expedition by Martin Edward Duncan Poore, Professor of Botany at the University of
Malaya, and student Ho Coy Choke and deposited mainly in the University of Malaya
herbarium in Kuala Lumpur.
Brian Laurence Burtt and Patrick James Blythe Woods, botanists from the Royal Botanic
Garden, Edinburgh, collected in Sarawak in 1962 (Kuching, Niah, Marudi, Mulu, Lambir,
Kapit, Lundu, Poi) and 1967 (Kuching, Hose Mountains, Kelabit Highlands). The
collections are mainly at the Edinburgh, Leiden and Sarawak herbaria. Swiss Hans Peter
Fuchs, a Shell Company research palynologist, collected in Sarawak (Santubong, Semengoh,
the Bau limestone, Baram, Marudi, Niah, Lambir), Brunei and Sabah (Kinabalu) in 1963;
parts of his Kinabalu trips were with Meijer, Sheila Collenette and Hermann Otto Sleumer,
a Leiden botanist who continued to Sarawak (Mt Matang) to collect. Duplicates of Fuchss
collections are also at Leiden, Kew and Sarawak. Eduardo Quisumbing, Filipino orchidologist, collected in 1963 in Sarawak (Bako, Santubong, Semengoh, Bau limestones), Brunei
and Sabah (Mt Kinabalu). In 1963, T.D. Pennington from Oxford University collected
mainly Meliaceae from Sarawak (Lundu, Rejang River, Kapit) and Sabah (Sepilok,
Kinabalu); these are mainly at the Forestry Herbarium, Oxford, Singapore, Kuching and
Sandakan.
The Japanese botanist Mitsuru Hotta collected, while on a Kyoto University expedition to
Borneo with Minoru Hirano of the Osaka City University as leader, in Brunei (Temburong)
and Sarawak (including the Tatau River and the Mt Mulu area, Limbang, Marudi) in
19631964 (Hotta 1964). About 10,000 collections were taken, deposited mainly at the
Kyoto, Kuching, Leiden and Kew herbaria. This resulted in a whole series of papers (Hotta
1964, 1965a, b, c, 1966a, b, 1967a,b; Iwatsuki 1965a,b; Tagawa 1965, 1967; Tagawa &
Iwatsuki 1966).
XXXI
Harold Emery Moore, Jr., Director of the Bailey Hortorium, collected palm specimens in
Sarawak (Matang, Bako, the Bau limestones, Bintulu, Nyabau) in 19631964 and Sabah
(Sandakan, Sepilok, Kimanis, Beaufort, Tenom, the Crocker Range, Kota Belud, Jesselton;
with Meijer) in 1964; the collections are mainly at the Bailey Hortorium, Kew, Kuching and
Sandakan herbaria. In Brunei, Joannes Petrus van Niel, a Shell Company palynologist,
made herbarium collections between 1964 and 1971; these are at Leiden.
Andries Kanis, a Dutch botanist stationed by the Dutch Ministry of Foreign Affairs at the
Sandakan herbarium in 1965, and acting head of that herbarium for about nine months
before returning to Leiden in 1966, collected about 250 numbers in Sabah (as North Borneo
became known then); he also collected in Brunei and Sarawak (Baram, Lambir Hills,
Nyabau and Segan Forest Reserves, Bintulu, Matang, Semengoh, Bako) in 1966. Bruce
Weber, an American botanist, was attached to the Sabah Parks and Forest Department
during 19651967 and collected some plants. In 1966, a Chinese botanist who came to be
settled in the Netherlands and employed at the Rijksherbarium in Leiden, Ding Hou, also
visited to collect in Sabah (mainly in the Sandakan and east coast districts, and on Kinabalu)
and Sarawak (mainly the Kuching, Bau, Bintulu, Sibu and Nyabau areas). Thomas Kenneth
Newell, a graduate student at the University of Hawaii, made a small collection, mainly of
monocots, in Sabah (Sandakan, Jesselton, Tamparuli and Kundasang) in 1966; his
collections are at the Bishop Museum, Sandakan, Paris, Leiden, Florence and Kew. Meijer
left Sabah in 1968, when also Ken Ogata from the Meguro Forest Experiment Station in
Tokyo collected a large number of herbarium vouchers together with wood samples from
Sabah. Shigeo Kurata of the Japanese Insectivorous Society collected pitcher-plant
specimens in Sabah (mainly Kinabalu) during a Mindoro/North Borneo expedition in
19671968. Others in Sabah collected actively during this period, including planter Jim B.
Comber, who took mainly orchids and ferns, principally from the Tenom area and the
Crocker Range; his collections were given to Kew. Engkik Soepadmo (University of
Malaya) collected with Anderson in the Simanggang and Bau areas in Sarawak in 1961, the
collections numbered by Anderson under the Sarawak herbarium series, and with Gordon
Smith (University of Malaya) and Paul Chai (Sarawak herbarium) in the Kapit and Bau
areas in Sarawak in 1969.
John Dransfield collected in Brunei and Sarawak (Bako, Matang, Bintulu) in 1968 (as a
research student), and Kalimantan Timor (east Borneo: Balikpapan, Samarinda, Kutei) and
Kalimantan Selatan (southeast Borneo: Djaro, and Mt Sarempaka; with Indonesian botanist
Kuswata Kartawinata and Dutch botanist Eduard Ferdinand de Vogel) in 1971 (while a
Colombo Plan botanist at Bogor); his collections were distributed mainly to Kew, Ithaca,
Bogor, Leiden, Kepong and Singapore. W. Soegeng Reksodihardjo, botanist of the Bogor
Herbarium, collected in the Kutei Nature Reserve in East Kalimantan in 1970; the
collections are mainly at Bogor.
Hotta collected again in Sabah in 19681969 with Shohei Kokawa, on Kinabalu and the
Trus Madi area, and in the Tawau Hills and on Mt Silam. In 1969, Peter Francis Cockburn
was appointed Forest Botanist at Sandakan and continued a statewide collecting drive. He
left in 1977. David Mabberley from Oxford collected in Sabah in 1974. Around this time,
there are some collections by Anthony Lamb, of the Sabah Agriculture Department, registered;
his collections were mostly orchids and although between 1977 and 1981 he used Sandakan
herbarium collecting numbers, the specimens are not always lodged with the herbarium
there. Lamb continued collecting into the 1990s, in the later part using his own numbers;
many of his collections were sent to Kew or Leiden. P.S. Shim, a plantation silviculturist in
XXXII
the Forest Department in Sandakan also collected (mostly ferns and orchids), using the
herbariums number series. Gary Shea, a Canadian University Service Overseas volunteer
attached to the herbarium in Sandakan, also collected widely in Sabah during 19711972;
Sheas collections are at Sandakan and distributed to Kew, Kepong and Leiden. Cockburn
organized the Trees of Sabah project, which saw the publication of two volumes (Cockburn
1976, 1980) that covered, in a very superficial and cursory manner, 44 angiosperm families
and two conifer families of tree. Shea assisted with the write-ups for several families in the
first volume. K.K. Tiong was appointed Forest Botanist in Sandakan in 1977, and continued
the collection in Sabah.
Paul P.K. Chai became Forest Botanist in Sarawak in 1970, collecting widely throughout
Sarawak. Ilias & Chai also collected with the 1964 Royal Society Expedition to Kinabalu in
Sabah. Hans Peter Nooteboom, Leiden botanist, made collecting trips to Sabah (Mt Alab,
Mt Lumarku, Mt Trus Madi, Tambunan) in 1969, and Sabah (Kinabalu, Sandakan) and
Sarawak (Bario, Mt Murud, Apo Batu Buli Range; with Paul Chai) in 1970. Nooteboom and
Chais collections number about 1600, distributed to the Sandakan, Kuching and Leiden
herbaria. In 1971 Anderson returned to collect in the Mt Mulu area in Sarawak. Chai again
collected in the Mulu area (Mt Api, Mt Benarat, Mt Buda, the Melinau Gorge) in 1975
together with B.L. Burtt.
De Vogel collected again in Kalimantan Selatan (Jaro, Mt Sarempaka) in 1972 and 1973;
his collections are principally at Bogor and Leiden. In 1972, University of Hull research
assistant R.J. Morley made a small collection of plants on Kinabalu, distributed to the
University of Hull, Kew and Leiden. In 1973, a New Zealander, David Wilson Ives,
consultant with the N.Z. Colombo Plan aid to Indonesias beef industry, collected plant
specimens in Kalimantan Selatan (Tandjung, Tabanio River, Djilatan, Djurong); his
specimens are at the Christchurch herbarium.
Benjamin Clemens Stone from the University of Malaya, who had collected in Sabah and
Sarawak in 1967, also collected in Sabahs Danum Valley in 1976 as a World Wildlife Fund
(WWF) consultant botanist, on Gaya Island and on Balambangan Island in 1977 (Stone
1980) in the same capacity. These several hundred collections were lodged with the
Sandakan, Kuching and University of Malaya herbaria. Earlier in 1970, Sabah Forest
Department ecologist J.E.D. Fox had also collected on Balambangan (about 80 numbers).
P.J. Martin and A. Clive Jermy of the British Museum (Natural History) collected on Mt
Mulu and along the Melinau and Melinau Paku Rivers in Sarawak in 1976.
In 1976, Hotta again collected in Sabah (Kota Kinabalu, Kinabalu, Ranau, Telupid,
Sandakan, Madai) and Sarawak, together with another botanist Michio Tamura on yet
another Kyoto University Expedition to Borneo (Kobayashi & Hotta 1978). Further Kyoto
University expeditions followed (Iwatsuki et al. 1983). In a 19781979 expedition to East
Kalimantan (Sekatak lowlands near Tarakan, Samarinda, Sebulu, Tabang, Balikpapan, Mt
Beratus) and South Kalimantan (Banjarmasin, Mt Besar), led by Kunio Iwatsuki and joined
by G. Murata, Masahiro Kato, J.P. Mogea and K. Kartawinata, some 4,650 numbers were
collected. Another expedition in 19801981 to East Kalimantan, led by H. Ogawa together
with Kato and Kunihiko Ueda and joined by Dedy Darnaedi of the Bogor Herbarium, and
which collected about 2,500 specimens, surveyed Sebulu, Tabang, the Menubar River, Mt
Kongkat, Mt Kongbotak, and the limestone Mt Njapa and Mt Buntung. In a further
expedition to East Kalimantan in 1981, led by Iwatsuki and joined by Kato, Motoharu
Okamoto and Ueda, Darnaedi and Eko Baroto Walujo of the Bogor Herbarium, and Rob
XXXIII
Geesink of the Rijksherbarium in Leiden, the Long Bawan region near the Sarawak border
and the Balikpapan and Samarinda areas were explored, netting some 4,750 numbers for the
Japanese collectors. Kalimantan collections were distributed mainly to the Kyoto, Bogor and
Leiden herbaria; Sabah specimens to the Kyoto, Sandakan and Leiden herbaria. Two other
Japanese collectors, Suehiro and Harigae, are credited with Kinabalu collections made in
1979, deposited in the Kyoto herbarium (Iwatsuki et al. 1983).
During the Royal Geographical Society Expedition to Mt Mulu in Sarawak in 19771978
(Jermy & Kavanagh 1982; Jermy 1984), several botanists collected in the area at different
times, including: Chai, Anderson, George Argent (Royal Botanic Garden, Ediburgh), J.A.R.
Kerby (Edinburgh), Ivan C. Nielsen (University of Aarhus in Denmark), Carlo K. Hansen
(Copenhagen Botanical Museum), J. Dransfield, B.C. Stone, Ruth Kiew (Agricultural
University of Malaysia), Barbara S. Parris (University of Cambridge), A. C. Jermy, A.
Touw (Leiden), Walter F.B. Julich (Leiden), B. Coppins (Edinburgh) and N. Sammy (a
Singapore-trained biologist working in Australia).
Dransfield collected rattans in Sabah in 1979 while on secondment from the Royal Botanic
Gardens, Kew, to the British Overseas Development Administration project in Sabah to
carry out a rattan survey. He was accompanied by his wife, Soejatmi Dransfield, who
collected bamboos. John and Soejatmi Dransfield were in Sarawak in 1981, collecting
rattans during a similar survey there.
Of check-lists and floras
ndoubtedly, the account by Stapf (1894) of the Kinabalu plant life must be the
earliest systematic account of the flora of any one place in Borneo. Stapf
enumerated 360 species of flowering plants, ferns and bryophytes. After Gibbs
(1914), which updated this Kinabalu account, it was largely people like Merrill (see
references here), Copeland (1917, 1935), Ames & Schweinfurth (1920), Fisher (1932),
Christensen & Holttum (1934), and others who produced numerous papers on Bornean plant
taxa. Hallier (1916) probably made the first attempt at a flora of Borneo (in German).
Merrills bibliographic enumeration of Bornean seed plants (Merrill 1921), which lists 4,924
species, not including the 120 orchid species listed by Ames & Schweinfurth (1920), giving
a total of 5,044 species (in 1,162 genera and 156 families), was strong motivation for
continuing work. Merrills 1921 estimate was a Bornean flora of some 9,000 flowering
plants including 3,000 species of tree. This work is essentially uncritical, as is the
subsequent enumeration by Masamune (1942), listing 7,201 species (1,310 genera, 165
families) of seed plants, that reproduces Merrills list with few changes and adds partial
information from the intervening period. Masamune also published a list of Bornean
pteridophytes (Masamune 1945), which lists 963 species of ferns and fern-allies in 118
genera.
To Masamunes enumeration, Merrill (1950) adds another 200 species of seed plants
published by Ridley, Airy Shaw and others and not consulted by Masamune; estimating the
size of the Bornean flora at 12,000 to 15,000 species (including seed plants, ferns and fernallies), he laments the lack of a descriptive flora.
While specialists in the large western herbaria continued to work on a taxonomic basis,
foresters check-lists of trees and dipterocarp floras (identification manuals) heralded a
XXXIV
next phase of botanical development in Borneo, beginning with those of Wood & Agama
(1956), Hasan & Ashton (1964), Wood & Meijer (1964) and Ashton (1964, 1968). van
Steenis (1950) pointed out that Borneo had the largest number of endemic genera among
islands in western Malesia and Airy Shaw (1975), who named many Bornean collections at
the Kew herbarium and continued an interest in Bornean plants over a long period,
considered Borneo to be ...floristically and phytogeographically...an area of prime
importancealmost the hub of Western Malesia... Forest floras and tree floras continued
to be in the ambitions of the forest departments of North Borneo (after 1963, Sabah) and
Sarawak, and early attempts include Ashtons manual of non-dipterocarp trees of Sarawak
(revising only 25 families of tree, completed by 1973 but published only in 1988) and the
two volumes of the Trees of Sabah (accounting for 46 families of tree: Cockburn 1976,
1980). Anderson (1980) published a check-list of Sarawak trees and a guide to the
Moraceae of Sarawak (Primack 1983) appeared to push on with what Ashton began, but the
writing of such manuals in the same series was not continued. Kiew (1984) called for a
concerted effort towards a flora of Borneo and in 1985, an attempt to continue the Trees of
Sabah project by B.C. Stone, then of the Academy of Natural Sciences, Philadelphia, never
materialised (Wong 1994).
For Indonesian Borneo, local check-lists and floras were longer in coming (Whitmore,
Tantra & Sutisna 1990a, b, c) and did not, in the case of some large and taxonomically
complex families, attempt to list beyond a few common taxa. However, keys by Ashton to
all known Bornean dipterocarp species and genera are given. One of the more recent
accounts includes that by Kessler & Sidiyasa (1994), of the trees of the BalikpapanSamarinda area in East Kalimantan.
Aside from the tree flora, modern identification manuals, amounting to floristic
enumerations, of the rattans of Sabah and Sarawak (J. Dransfield 1984, 1992), and the
bamboos of Sabah (S. Dransfield 1992) have been published. Modern compilations of the
orchids of Borneo (Chan et al. 1994, Vermeulen 1991) as well as a check-list for Borneo
(Wood & Cribb 1994) have also been published.
In the meantime, the incredibly high plant diversity of Kinabalu, Borneos highest
mountain, continues to attract systematic study. As part of a modern botanical enumeration
of Kinabalus vascular plant flora, John Beaman and associates have published annotated
check-lists of the ferns (Parris, Beaman & Beaman 1992) and orchids (Wood, Beaman &
Beaman 1993). Farther south, a new inventory of the Brunei flora towards compiling a
modern check-list, initiated in 1989 between the Brunei Forestry Department and the Royal
Botanic Gardens, Kew, is yielding more information of a very rich flora, with many
novelties.
Current estimates of the Bornean vascular flora hover between 9,000 (Merrills original
estimate in 1921) and 15,000 (the upper limit of Merrills subsequent estimate in 1950).
This state of uncertainty surely reflects the existence of large gaps still in our knowledge.
XXXV
References
Airy Shaw, H.K. 1975. The Euphorbiaceae of Borneo. Kew Bulletin Additional Series IV.
245 p.
Ames, O. & C. Schweinfurth. 1920. Orchids of Mount Kinabalu, British North Borneo.
Orchidaceae 6 (IXIV): 1233.
Anderson, J.A.R. 1963. The flora of the peat swamp forests of Sarawak and Brunei,
including a catalogue of all recorded species of flowering plants, ferns and fern allies. Gard.
Bull. Sing. 20: 131228.
Anderson, J.A.R. 1980. A Check List of the Trees of Sarawak. Dewan Bahasa dan Pustaka
for Forest Department, Sarawak, Kuching. 364 p.
Ashton, P.S. 1964. Manual of the Dipterocarp Trees of Brunei State. Oxford University
Press, London. xii + 242 p.
Ashton, P.S. 1968. A Manual of the Dipterocarp Trees of Brunei State and of Sarawak.
Supplement. Borneo Literature Bureau for Sarawak Forest Department, Kuching. viii + 129.
Ashton, P.S. 1988 Manual of the Non-Dipterocarp Trees of Sarawak. Vol. II. Dewan
Bahasa dan Pustaka for Forest Department Sarawak, Kuching. xix + 490.
Beccari, O. 1902. Nelle Foreste di Borneo. Firenze.
Browne, F.G. 1955. Forest Trees of Sarawak and Brunei and their Products. Government
Printing Office, Sarawak. 369 p + xviii.
Brunig, E.F. 1968. Der Heidewald von Sarawak und Brunei. Mitteilungen der
Bundesforschungsanstalt fr Forst- und Holzwirtschaft Nr. 68. Vol. 1: vi + 1152d, 28 fig.;
Vol. 2: 153431, 16 tables, 14 fig.
Burgess, P.F. 1966. Timbers of Sabah. Sabah Forest Records No. 6. xviii + 513 p.
Burtt, B.L. 1964. Beccaris ascent of Mount Poi, Sarawak. Flora Malesiana Bulletin 19:
11311132.
Chan, C.L., A. Lamb, P.S. Shim & J.J. Wood. 1994. Orchids of Borneo. Vol. 1.
Introduction and a Selection of Species. Sabah Society & Royal Botanic Gardens, Kew.
xviii + 402 p.
Christensen, C. & R.E. Holttum. 1934. The ferns of Mt Kinabalu. Gard. Bull. Str. Settlem.
7: 191324.
Cockburn, P.F. 1976. Trees of Sabah, Volume 1. Sabah Forest Records No. 10. Borneo
Literature Bureau for Forest Department Sabah, Kuching. xv + 261.
Cockburn, P.F. 1980. Trees of Sabah, Volume II. Sabah Forest Records No. 10. Dewan
Bahasa dan Pustaka for Forest Department, Sabah, Kuching. xiii + 124.
XXXVI
Copeland, E.B. 1917. New species and a new genus of Borneo ferns chiefly from the
Kinabalu collections of Mrs. Clemens and Mr. Topping. Philip. J. Sci., C, Bot. 12: 4565.
Copeland, E.B. 1935. Additional ferns of Kinabalu. Philip. J. Sci. 56: 471481.
Corner, E.J.H. 1964. A discussion on the results of the Royal Society Expedition to North
Borneo, 1961. Proc. Roy. Soc., B, 161: 191.
Dransfield, J. 1984. The Rattans of Sabah. Sabah Forest Records No. 13. Forest
Department, Sabah. v + 182.
Dransfield, J. 1992. The Rattans of Sarawak. Royal Botanic Gardens, Kew & Sarawak
Forest Department. vi + 233 p.
Dransfield, S. 1992 The Bamboos of Sabah. Sabah Forest Records No. 14. Forestry
Department, Sabah. xi + 94 p.
Fisher, C.E.C. 1932. Contributions towards a Flora of British North Borneo. 1. Kew
Bulletin of Miscellaneous Information, No. 4.
Foxworthy, F.W. 1916. I. Timbers of British North Borneo. II. Minor Forest Products and
Jungle Produce. Government of British North Borneo, Department of Forestry, Bulletin No.
1.
Gibbs, L.S. 1914. A contribution to the flora and plant formations of Mt. Kinabalu and the
highlands of British North Borneo. J. Linn. Soc., Bot. 42: 1240.
Hallier, H. 1916. Beitrge zur Flora von Borneo. Beih. Bot. Centr. Bl. 2e Abt. 34: 1953.
Hasan bin Pukul & P.S. Ashton. 1964. A Checklist of Brunei Trees. 132 p.
Heine, H. 1951. Pflanzen der Sammlung J. & M.S. Clemens von Mount Kinabalu in
Britisch Nord-Borneo. Fedde, Rep. 54: 223248.
Heine, H. 1953. Diagnoses novae plantarum in Borneo septentrionali a J. et M.S. Clemens
lectarum, Pars II. Mitt. Bot. Munchen 6: 208209.
Hotta, M. 1964. Notes on the Kyoto University Borneo Expedition 196364. Tanken 8:
18. (in Japanese)
Hotta, M. 1965a. Itinerary of the Borneo (notes on the vegetation) (sic!). Acta Phytotax.
Geobot. 21: 153160. (in Japanese, with English title)
Hotta, M. 1965b. Notes on Schismatoglottidinae of Borneo, I. Mem. Coll. Sci. Univ. Kyoto,
B, 32: 1930.
Hotta, M. 1965c. Note on the Nepenthes of Borneo. Trans. Jap. Insectiv. Pl. Soc. 34: 19.
(in Japanese)
XXXVII
Hotta, M. 1966a. Notes on Bornean plants, I. Acta Phytotax. Geobot. 22: 110.
Hotta, M. 1966b. Notes on Schismatoglottidinae of Borneo, II. Mem. Coll. Sci. Uni. Kyoto,
B, 32: 223238.
Hotta, M. 1967a. Notes on Bornean plants, II. Acta Phytotax. Geobot. 22: 153162.
Hotta, M. 1967b. Notes on the wild banana (sic!) of Borneo. J. Jap. Bot. 42: 344352.
Iwatsuki, K. 1965a. Ferns of Borneo, collected by M. Hirano and M. Hotta, 1. Acta
Phytotax. Geobot. 21: 91100.
Iwatsuki, K. 1965b. Ferns of Borneo, collected by M. Hirano & M. Hotta, 2. Acta Phytotax.
Geobot. 21: 165171.
Iwatsuki, K., M. Kato, M. Okamoto, K. Ueda & D. Darnaedi. 1983. Botanical expedition to
East Kalimantan, during 15th June and (sic!) 15th September 1981. In: T. Hidaka (ed.),
Taxonomical and Evolutionary Studies on the Biota in Humid Tropical Malesia, with
reference to Diversity of the Species, pp. 3169. Department of Zoology, Kyoto University,
Japan.
Jermy, A.C. (ed.). 1984. Studies on the Flora of Gunung Mulu National Park, Sarawak.
Forest Department, Sarawak, Kuching. xv + 233 p.
Jermy, A.C. & K. Kavanagh. 1982. Gunung Mulu National Park, Sarawak. An account of
its environment and biota being the results of the Royal Geographical Society/Sarawak
Government Expedition and Survey 19771978. Part I. Sarawak Mus. J., Special Issue No.
2. xxiv + 279 p.
Keith, H.G. 1937. A Preliminary List of North Borneo Plant Names. North Borneo Forest
Records No. 2. 528 p.
Keith, H.G. 1947. The Timbers of North Borneo. North Borneo Forest Records No. 3. 154
p.
Kessler, P.J.A. & K. Sidiyasa. 1994 Trees of the Balikpapan-Samarinda area, East
Kalimantan, Indonesia. Tropenbos Foundation, Wageningen. 446 p.
Kiew, R. 1984. Towards a Flora of Borneo. In: Ismail Sahid, Zainal Abidin A. Hasan, A.
Latiff Mohamed & A. Salam Babji (eds.) Research Priorities in Malaysian Biology, pp.
7380. Penerbit Universiti Kebangsaan Malaysia, Bangi, Malaysia.
Kobayashi, T. & M. Hotta. 1978. Biological expedition to the rain-forest of Sabah in 1976.
Contrib. Biol. Lab. Kyoto Univ. 25 (3): 255271.
Masamune, G. 1942. Enumeratio Phanerogamarum Bornearum. 739 p.
Masamune, G. 1945. Enumeratio Pteridophytarum Bornearum. ii + 124 p.
XXXVIII
Meijer, W. 1969. The contribution of Governor Charles Van de Leur Creagh to botanical
exploration in Sabah, 18881895. Sabah Soc. J. 5: 6768.
Merrill, E.D. 1916. Notes on the flora of Borneo. Philip. J. Sci. 11, Bot.: 49100.
Merrill, E.D. 1917a. Contributions to our knowledge of the flora of Borneo. J. Str. Br. Roy.
As. Soc. 76: 75117.
Merrill, E.D. 1917b. Alabastra Borneensia. J. Str. Br. Roy. As. Soc. 77: 189247.
Merrill, E.D. 1918. New species of Bornean plants. Philip. J. Sci. 13 (2), Bot.: 67122.
Soc., Special Number. 637 p.
Merrill, E.D. 1922a. New or noteworthy Bornean plants (Part 1). J. Str. Br. Roy. As. Soc.
85: 151201.
Merrill, E.D. 1922b Additions to our knowledge of the Bornean flora. Philip. J. Sci. 21 (6),
Bot.: 515534.
Merrill, E.D. 1924. Plants from Banguey Island. Philip. J. Sci. 24(1): 113116.
Merrill, E.D. 1926. The Flora of Banguey Island. Philip. J. Sci. 24(3): 341427.
Merrill, E.D. 1929. Plantae Elmerianae Borneenses. Univ. of California Publications in
Botany 15: 1316.
Merrill, E.D. 1950. A brief survey of the present status of Borneon botany. Webbia 7:
309324.
Parris, B.S., R.S. Beaman & J.H. Beaman. 1992. The Plants of Mount Kinabalu. I. Ferns
and Fern Allies. Royal Botanic Gardens, Kew.
Primack, R.B. 1983. Foresters Guide to the Moraceae of Sarawak. Forest Department,
Sarawak, Kuching. 140 p.
Smythies, B.E. 1965. Common Sarawak Trees. Borneo Literature Bureau.
Stapf, O. 1894. On the flora of Mount Kinabalu in North Borneo. Trans. Linn. Soc. Lond.,
Bot. 4: 69263.
Steenis, C.G.G.J. van. 1950. The delimitation of Malaysia and its main geographical
division. Flora Malesiana 1, 1: LXXLXXV.
Steenis-Kruseman, M.J. van. 1950. Malaysian plant collectors and collections being a
Cyclopaedia of Botanical Exploration in Malaysia and a guide to the concerned literature up
to the year 1950. Flora Malesiana, 1, 1: CLII + 1639.
XXXIX
Steenis-Kruseman, M.J. van. 1954. Numerical list of the collections of Haviland in Borneo.
Duplicated typescript: Flora Malesiana Foundation, Leiden.
Steenis-Kruseman, M.J. van. 1958. Malaysian plant collectors and collections. Supplement
I. Flora Malesiana 1, 5 (4): CCXXXVCCCXLII (superposed pagination 1108).
Steenis-Kruseman, M.J. van. 1974. Malaysian plant collectors and collections. Supplement
II. Flora Malesiana 1, 8 (1): ICXV (superposed pagination 1115).
Stone, B.C. 1980 The vegetation and plant communities of Pulau Balambangan, Sabah. J.
Malays. Br. Roy. As. Soc. 53 (1): 6889.
Tagawa, M. 1965. Ferns of Borneo, collected by M. Hirano and M. Hotta, 3. Acta Phytotax.
Geobot. 21: 173180.
Tagawa, M. 1967. Ferns of Borneo, collected by M. Hirano and M. Hotta, 5. Acta Phytotax.
Geobot. 22: 183191.
Tagawa, M. & K. Iwatsuki. 1966. Ferns of Borneo, collected by M. Hirano and M. Hotta, 4.
Acta Phytotax. Geobot. 22: 8794.
Vermeulen, J.J. 1991. Orchids of Borneo. Vol. 2. Bulbophyllum. Royal Botanic Gardens,
Kew & Toihaan Publ. Co., Malaysia. x + 342 p.
Whitmore, T.C., I.G.M. Tantra & U. Sutisna. 1990a. Tree Flora of Indonesia. Check List for
Kalimantan. Part I. Forest Research and Development Centre, Bogor. ix + 1181. (Date on
cover 1989)
Whitmore, T.C., I.G.M. Tantra & U. Sutisna. 1990b. Tree Flora of Indonesia. Check List for
Kalimantan. Part II.1. Forest Research and Development Centre, Bogor. 182429.
Whitmore, T.C., I.G.M. Tantra & U. Sutisna. 1990c. Tree Flora of Indonesia. Check List for
Kalimantan. Part II.2. Forest Research and Development Centre, Bogor. 431620.
Wong, K.M. 1994. A tribute to Benjamin C. Stone, 19331994. Sandakania 4: 129.
Wood, G.H.S. & J. Agama. 1956. Check List of the Forest Flora of North Borneo. North
Borneo Forest Records No. 6.
Wood, G.H.S. & W. Meijer. 1964. Dipterocarps of Sabah (North Borneo). Sabah Forest
Records No. 5. 344 p. (The cover differs from the full-title page in stating W. Meijer &
G.H.S. Wood)
XL
Wood, J.J., R.S. Beaman & J.H. Beaman. 1993. The Plants of Mount Kinabalu. 2. Orchids.
Royal Botanic Gardens, Kew.
Wood, J.J. & P.J. Cribb. 1994. A Checklist of the Orchids of Borneo. Royal Botanic
Gardens, Kew. xii + 409 p.
XLI
BIOGEOGRAPHY AND ECOLOGY

P.S. Ashton
The Arnold Arboretum,
Harvard University, Cambridge,
Massachusetts, U.S.A.
The physical setting
otanists from Peninsular Malaysia who visit Sabah and Sarawak for the first time,
and who are interested in floristic ecology, are startled to find that their traditional
reference points are missing. The customary altitudinal sequencelowland
dipterocarp, hill dipterocarp, upper dipterocarp, (lower) montane oak and (upper)
ericaceous forestis either unrecognisable as in the case of these categories of dipterocarp
forest, or does not consistently adhere to the expected altitudinal sequence. With closer
familiarity, a new set of forest types correlated to soil becomes recognisable, which more or
less obscures the altitudinal sequence. Furthermore, the varied coastline in the northwest,
which is mostly bordered by vast peat swamps for which there is no peninsular equivalent,
lacks any consistent coastal hill association. It was Symington (1943) who first put forward
the floristic classification of Peninsular Malaysian forests which is still adhered to, but up to
now no East Malaysian scheme has been proposed. Why is it that the floristic ecology of
East Malaysian forests is so different?
The hills of Peninsular Malaysia are thought to have remained above sea level since before
the origin of the flowering plants, in the Jurassic era perhaps 150 million years ago, and the
present landscape may be dated from perhaps the Cretaceous, c. 100 million years ago
(Richardson 1947; Gobbett and Hutchinson 1973). Their soil cover is extraordinarily deep:
as a rule, soils there exceed two meters over rotting rock through which occasional roots
may penetrate, and sometimes much more. By contrast, in much of northern Borneo it is
difficult to find soils as deep as one meter. This is because northern Borneo is at the
southeastern edge of the ancient Laurasian continent and continues, even now, to experience
coastal up- and downwarping, alternately sinking and accumulating sediments, then lifting
and rapidly eroding, and thereby forming series of anti- and synclinally folded sedimentary
rocks (Liechti et al. 1960). The result is a sharply dissected terrain of parallel, narrow
interior ridges and valleys, in which ridge-top, slope and valley bottom are sharply
differentiated. This landscape is much more rugged than the peninsular main range, which is
gentle by comparison and with a dendritic lattice of ridges.
Each successive period of uplift and erosion has led to a further loss of clay minerals and
nutrients, and to the creation of new hills to the north and east near the coast which each
time become sandier and with poorer soils, and whose rocks are successively softer. These
sandy, infertile soils are mostly yellow, but are so acidic that litter decomposition is slow
XLIII
and a mantle of peaty organic matter forms. They are known as humult ultisols. The organic
layer impedes surface erosion and, as a result, these young sandy hills alone quickly develop
a mature rounded topography and remarkably deep soils. This contrasts with coastal clay
hills which, where they occur, remain steep even if low, with clear signs of surface erosion;
there, the only deep soils are formed by slumping downslope. Superimposed on this
landscape in the lowlands, along the northwestern coast where the tectonic activity is
greatest, are the imprints of changing sea levels during and after the ice ages: the current
coastline, mostly a straight, sandy shore often bordered with Casuarina equisetifolia and its
associates, is backed by a vast peat swamp which originated as an embayed mangrove when
the sea level last declined, about 5,500 years ago. Behind again, and also perched on the
hills which in places push to the coast between Bintulu and Kota Kinabalu, are extensive
white sand terraces bearing giant podsol soils with deep, peaty organic surface layers
(except where they have been burned); these were former coastlines during high sea levels,
between the great ice ages.
Inland, the predominating sedimentary hills are intruded by volcanic masses, granite, and
limestone. The volcanic rocks, which are mainly tertiary, are as diverse as the sediments,
from acid rhyolites yielding similar humult ultisols to the sandy coastal hills, through dacites
and base-rich fertile basalts, to ultramafic rocks which carry diverse soil types and
consequently floras, and which are widespread in northern and eastern Sabah but absent
from Sarawak. The granites are young, rocky and covered with shallow soils. The
outstanding example of a granite formation in this territory is the great dome of Kinabalu,
the highest mountain between the Himalayas and New Guinea, which arose less than one
million years ago. There are other, tertiary, granite mountains in West Sarawak. The several
limestone massifs in eastern Sabah and northeastern and western Sarawak are generally of a
pure dolomitic limestone which erodes only by solution, yielding no mineral soil. What soils
there, are organic and surprisingly acidic.
The floristic composition of forests is most markedly differentiated where the soils are
limiting to plant growth and survival, by adverse water conditions or low nutrients. The
combination of sharply defined topography, extraordinarily diverse rock substrates, and
generally shallow, low nutrient soils in most of Sabah and Sarawak has led to a greatly
diversified forest cover in which soils-correlated differentiation of forest types dominates
over altitudinal and other influences. This is particularly so between Kota Kinabalu and the
west coast of Borneo south to Pontianak in Kalimantan, where sandy sedimentary soils and
the often deep and diversified coastal peat swamps create a diversified landscape quite
different from those found elsewhere in the country.
Impact on taxonomy
notable result, familiar to the experienced field botanist in Sabah and Sarawak but a
potential pitfall to the herbarium taxonomist, is a marked and complex pattern of
small-scale variation in the characteristics of widespread species which occur in the
many disjunct habitats of the region. Identification of countless species from
ecological plots scattered through Brunei and Sarawak persuade us that finely distinguished
species based on single specimens will rarely be maintained as knowledge increases, and
that van Steenis' (1957) admonition to hold to a conservative and broad species concept may
prove wise.
XLIV
XLV
Forest types
he overall ecology of the main forest types has been well summarised by Whitmore
(1984). Here, emphasis is given to the major floristic associations. The lowland as well
as mountainous forests of Sabah and Sarawak vary dramatically in stature and
canopy structure, from less than 10 m tall with dense, even canopy to over 70 m,
with scattered clumps of giant emergent dipterocarps or Koompassia, or sometimes lofty,
dense, even canopy as in the alan (Shorea albida) peat swamps. However, forest stature and
structure, which correlate with soil water availability, do not always correlate in Borneo
with species composition, which correlates most closely with soil chemistry. Floristic
variation is actually continuous, but there are also certain rather sharp environmental
boundaries which differentiate major floristic associations. The following main lowland
types can therefore be recognised:
Mixed dipterocarp forest (MDF). Although dipterocarps decline in abundance and in

number of species with altitude, and although there are some ten species which are confined
to higher altitudes, the change in flora with altitude is continuous and no altitudinally zoned
types can be recognised. Most strikingly seraya (Shorea curtisii), although scattered in the
coastal hills of Sarawak and southwestern Sabah, is local and never dominant there, and is
rare on the inland ridges. Therefore, there is no consistent coastal hill species association,
nor is there a recognisable "hill dipterocarp forest." In Borneo, the upland forests on yellowred soils which are dominated by dipterocarps in the emergent canopy are termed Mixed
Dipterocarp Forests (MDF) to distinguish them from others, such as the S. albida-dominated
peat swamp forests, in which one or only a few emergent dipterocarp species occur. The
major floristic division in these forests is between Mixed Dipterocarp Forests in which
either Dryobalanops lanceolata or Shorea parvifolia (often also with Shorea macroptera)
are among the commonest species, and which are confined to clay-rich soils lacking an acid
matted surface organic mat; and forests in which Anisoptera grossivenia and variously
Dryobalanops aromatica and Dipterocarpus globosus, or in western Sarawak Shorea
falcifera, are among the commonest emergent species, and which are confined to humult
ultisols. Probably fewer than one quarter of species are shared between these two major
types (cf. Ashton 1964: 34). There is also a widespread type on intermediate, sandy clay
soils of the lowlands and also inland ridges, of which Dipterocarpus crinitus is a
characteristic representative, and in which Shorea macroptera is also common.
These major types vary geographically and with habitat, but they are almost always sharply
distinguishable and separated in the landscape, whereas variation within them is continuous.
Within the first type, giant forests in which Dryobalanops lanceolata, Shorea superba, and
Dipterocarpus caudiferus were abundant, and Octomeles sumatrana, Pterocymbium
tubulatum and Pterospermum javanicum were characteristic pioneers, once clothed the
extensive fertile volcanic hills and sedimentary clay loams of lowland east Sabah as well as
the lower basalt massifs and low clay hills of Sarawak. These forests were the first to be
logged and their land converted to agriculture but some remain at Quoin Hill in Tawau,
Bukit Mersing in Sarawak, and elsewhere. The Shorea parvifolia subtype is the prevailing
forest of inland Sabah and Sarawak, on the shallower yellow clays and sandy clays which
cover much of the inland sediments.
The most distinctive forest type on humult ultisols is that which is confined to its deepest
soils, and therefore to low soft sandstone hills which are mostly but not always near the
coast. The most extensive areas are in the Lambir Hills and the Andulau and Labi Hills of
Brunei. Smaller islands occur at Beaufort Hill in southwest Sabah, in the Ulu Tutoh, Baram
XLVI
BIOGEOGRAPHY AND ECOLOGY (ASHTON)
and Belait in Brunei, in Similajau, Nyabau and Segan forests in Bintulu, on the Arip
rhyolite, in the northernmost hills between the lower Balingian and Mukah rivers, and in
some small patches west of the Lupar including the foothills of Gunong Gaharu, and Bukit
Undan in Bau. This type deserves special mention on account of the confinement of certain
well known peninsular species there (including Dryobalanops aromatica and Shorea
curtisii), because it appears to be the richest in species of all forest types in Malaysia,
because of its remarkable endemism, and because it still remains unfamiliar to many
foresters and biologists. The commoner dipterocarp species which are concentrated in or
confined to it include Anisoptera grossivenia, Dipterocarpus globosus, D. lowii, D. rigidus,
D. sarawakensis, Dryobalanops aromatica, Hopea beccariana, H. treubii, H. tenuinervula,
Shorea acuta, S. crassa, S. curtisii, S. cuspidata, S. elliptica, S. falcifera, S. flemmichii, S.
geniculata, S. kunstleri, S. ladiana, S. laxa, S. ovata, S. rubella, and S. slootenii; species
apparently confined to this type which appear in this first volume of the Tree Flora include
Anisophylla ferruginea, Canarium divergens, C. grandifolium, Santiria megaphylla,
Sarawakodendron filamentosum, Parinari metallica, Allantospermum borneense (often the
most abundant species, and very characteristic), and Quassia borneensis.
A variety of other MDF, floristically related to this but generally poorer in endemics but
more widespread in Sabah and Sarawak, occur on humult ultisols richer in clay, where up to
30% of the flora also occur in the S. parvifolia subtype. Dipterocarpus crinitus, Shorea
macroptera and on ridges Dryobalanops beccarii distinguish this association. There is also
a range of MDF on shallower, frequently more sandy soils, where there is an increasing
heath (kerangas) forest element.
Heath (kerangas) forest. These are forests, often but not invariably of low stature and
lacking emergents, that are confined to organic white sand podsol soils. Their flora is very
distinctive, with perhaps fewer than one half of species also occurring in MDF, usually on
humult ultisol soils. Although Brunig's monograph (1974) provides extensive insight into
the great floristic diversity of kerangas forest, the relationship between floristic and habitat
variation remains even more poorly understood in this type of environment than is the case
with MDF. Kerangas occurs in two principal habitats: on raised beach terraces where the
podsols are generally deep except in the center and along the lower edges where they are
often poorly drained; and on sandstone ridges and plateaux where they are generally shallow
and often rocky. There appear to be floristic differences betweenand to perhaps a lesser
extent withinthese habitats; lowland Agathis, for instance is almost entirely confined to
terraces. Certain families of kerangas are also particularly well represented in upper
montane forest ("elfin woodland"). They include Myrtaceae, Theaceae, and Podocarpaceae,
and to some extent Ericaceae. Clusiaceae (Guttiferae) and Ebenaceae are also well
represented in kerangas.
Peat swamps. There are extensive peat swamps up the northwestern coast, eastwards to
Papar in Sabah. Anderson (1963) recognised six phasic communities from the margins to
the centers of individual swamps, which he regarded as succeeding one another over the
time scale of the development of the dome of the raised bog. Anderson documented a total
vascular flora of only 317 species of flowering plants, including several which are confined
to certain districts and which otherwise are known from kerangas or MDF on humult
ultisols. The endemic flora is small, most species being found also in kerangas. Notable is
the tendency towards canopy dominance of Shorea albida and, towards the centers, Litsea
XLVII
palustris, Combretocarpus rotundatus and sometimes Dactylocladus stenostachys; but there

are no subcanopy dominants.
Other specialized lowland forests. There are major limestone formations in Bau District of
western Sarawak, Mulu National Park, and in East Sabah, and also many minor exposures.
The limestone flora is rich in endemics, particularly among herbaceous species; the general
characteristics are similar to the limestone outcrops of Peninsular Malaysia. The highest
limestone in Malaysia is at Gunong Benarat, Mulu NP, at 1,300 m, which bears upper
montane elfin woodland on its jagged upper slopes.
In several parts of Sabah, notably on the slopes of G. Kinabalu and in the northeast and east,
are extensive exposures of ultramafic rocks. These exposures can bear freely draining acid
soils with a surface organic layer. In this case the forest is of more or less short stature with
elements of both the MDF on humult ultisols, and kerangas. A small but important element
of Bornean affinities are present, such as the strictly endemic Borneodendron aenigmaticum
Airy-Shaw (Euphorbiaceae). In other areas, however, as at Kinabalu, the soil is a friable
red-brown loam and the flora bears affinities with that found on basalt.
The mangroves of Sabah and Sarawak do not compare in extent with those of the Peninsular
west coast, but are noteworthy because they are far richer in species than any others in the
world. The most extensive are at the mouths of the Kinabatangan River, and around the Bay
of Brunei where some magnificent primary stands still survive. The floristic formations are
similar to those of the Peninsula, as in most respects are those of the inland river banks and
alluvium.
Altitudinal zonation
he sharp topography and bold ridge lines of the mainly sedimentary mountains of
Sabah and Sarawak cast a stamp on vegetation differentiation which obscures the
zonation recognisable in the Peninsula. Below 1,200 m the primary differentiation is
between ridge and slope forests (cf. Ashton 1964: 66-70). Lower, broader shale
ridges bear the tallest forests, but those with the fewest endemics. Higher ridges are variably
supported by sandstone strata: Where the soils are deep, a variety of MDF on shallow
humult ultisols or yellow clay types will be present; Dryobalanops beccarii, Shorea laevis
and Shorea multiflora are characteristic species. The slopes bear S. parvifolia-type MDF
with pockets of Dryobalanops lanceolata on gentle lower surfaces. On shallow soils, which
increase on the ridges with the altitude, a distinct short-stature ridge forest is widespread
between 800 and 1,200 m, where Shorea flaviflora, Vatica umbonata and V. dulitensis are
characteristic of dipterocarps, and Engelhardia spp. are common. Where sandstone is
exposed, short-stature kerangas prevails, and this grades imperceptibly and continuously on
sandstone substrates into "upper montane" ericaceous elfin forest; this retains many
kerangas species, such as Eugenia (Syzygium) bankense and Calophyllum nodosum.
Nevertheless, true upper montane elements such as Weinmannia blumei, Leptospermum
flavescens, Xanthomyrtus spp., Rhododendron quadrasianum, Drimys piperita, Dacrydium
comosum and Phyllocladus hypophyllus can begin to appear as low as 600 m on the most
exposed peaks and skeletal soils, but only become common above 1,300 m.
XLVIII
On the slopes at about 1,300 m as in the Peninsula, there is an ill-defined transition on clayrich soils to oak-laurel dominated "lower montane forest" that is at approximately the same
altitude at which the upper montane elements become common on the sandstone ridges.
Again, it is the differentiation between ridge and slope, and between soils with and without a
surface organic horizon, which primarily correlate with forest type differentiation, although the
slopes adjacent to exposed summit ridges do become clothed in organic soils and ericaceous
forest.
Historical biogeography
his Tree Flora will for the first time provide a sound basis for historical
biogeographic analysis of the northern Borneo flora. For the present, two provinces
are recognised, Northwest Borneo from Pontianak in West Kalimantan northeast to
the Crocker Range and Kota Kinabalu, and Sabah north and east of the Crocker
Range.
The Northwest Borneo Province has phytogeographic connections with the Riau Archipelago
and eastern coastal Peninsular Malaysia, that is the Riau Pocket of Corner (Corner 1960,
Ashton 1992). Within it, two subprovinces are recognisable, being apparently divided east
and west of the Lupar Valleys in west Sarawak and the Kapuas Lakes in Kalimantan
(Ashton 1972, 1992). The western subprovince contains a number of endemics, and also a
number of Peninsular Malaysian species not found elsewhere in Borneo such as the
dipterocarps Shorea dasyphylla, S. dealbata, S. falcifera, and S. resinosa, and, in this first
volume of the Tree Flora, Dacryodes rubiginosa, Glyptopetalum quadrangulare, and
Lophopetalum pachyphyllum. The many endemic species include the dipterocarps
Dryobalanops fusca, Shorea alutacea, S. bakoensis, S. cuspidata, S. elliptica, S. induplicata,
S. lunduensis, S. pallidifolia, S. richetia, S. splendida, S. stenoptera, S. subcylindrica, Vatica
compressa, V. pedicellata and, in this volume, Alangium circulare, Haplolobus beccarii, H.
inaequifolius, Anisophyllea rhomboidea, Ellipanthus beccarii var. beccarii, Mastixia
glauca, Schuurmansiella angustifolia, Sarcotheca macrophylla, Glycosmis longisepala and
Maclurodendron parviflorum. Likewise, a number of species of the Province are absent
from this subprovince including, surprisingly, Dryobalanops aromatica and Shorea curtisii.
Ecological plots provide evidence that the habitats which occur in ecological islands,
notably the areas of the organic podsols and humult ultisols so characteristic of this Province
but also the areas of limestone and ultramafic rocks, experience significant between-locality
variation in the characters of many species, and also the apparent fortuitous absence of
species from suitable localities. The lack, to date, of any record of the usually common
humult ultisol species Upuna borneensis from the Lambir Hills is a striking example.
The Northeastern Province is not so clearly subdivided, but it includes two habitats unique
for all Borneo. Kinabalu, owing to its great height, is the home both of many endemics and,
on a smaller scale presumably owing to its youth, a regionally widespread upper montane
and alpine element. The extensive ultramafic extrusions support a number of endemics, both
in the lowlands and also on Kinabalu to which such notable species as Scaevola micrantha,
S. chanii and Pittosporum linearifolium are confined. But the Province is notable in addition
for its lowland endemic flora, which includes the dipterocarps Dryobalanops keithii, Hopea
badiifolia, Parashorea tomentella, Shorea symingtonii, S. waltonii, all of S. parvifolia type
XLIX
MDF, and, in this volume, Dacryodes elmeri, Kokoona sabahana, Microtropis sabahensis,
Parinari argento-sericea, Connarus agamae, Atuna cordata, Sarcotheca rubrinervis,
Maclurodendron pubescens, Melicope jugosa, M. sororia, M. subunifoliolata,
Monanthocitrus oblanceolata, and Turpinia nitida. Also, a number of Philippine species
including Dipterocarpus validus, Protium connarifolium, Pittosporum resiniferum,
Melicope bonwickii, M. denhamii, Turpinia borneensis and Scaevola micrantha, and species
such as Canarium asperum, C. decumanum and Kibara obtusa are only known from this
part of Borneo. Several of these are confined to ultramafic substrates.
The southward extent of this Province into East Kalimantan is as yet inadequately
understood. There is limited ultramafic rock in Kalimantan but some species of the zonal
clay soils, such as D. keithii, hardly reach Tidung while others, such as H. badiifolia, extend
as far as Balikpapan. This variable extent of ranges is not surprising in the Shorea parvifolia
type MDF on clay soils, which extends continuously over the hills of East and Central
Borneo. A number of other Philippine species such, for instance, as the dipterocarps
Parashorea malaanonan, Shorea almon and S. falciferoides, also extend westward into
north and central Sarawak.
Lastly, there is an element in the Shorea parvifolia type MDF, represented by such species
as Dipterocarpus mundus, D. pachyphyllus, Hopea bullatifolia, H. centipeda, H. dasyrrhachis,
H. fluvialis, H. megacarpa, Shorea agamii ssp. diminuta, S. asahi, S. collaris, S. iliasii, Vatica
endertii and V. granulata, which have been found from innermost lowland Sarawak across to
the Ulu Barito, and which may eventually delineate a distinct central Bornean trans-montane
floristic province.
References
Anderson, J.A.R. 1963. The flora of the peat swamp forests of Sarawak and Brunei
including a catalogue of all recorded species of flowering plants, ferns, and fern allies. Gard.
Bull. Sing. 20: 131228.
Ashton, P.S. 1964. Ecological Studies in the Mixed Dipterocarp Forests of Brunei State.
Oxford Forestry Memoirs 25.
Ashton, P.S. 1972. The quarternary geomorphological history of western Malesia and
lowland forest phytogeography. In P. & M. Ashton (eds.), Hull Geog. Dept. Misc. Series 13.
Transactions of the second Aberdeen-Hull symposium on Malesian ecology: The
quarternary era in Malesia. Pp. 3549
Ashton, P.S. 1992. Plant conservation in the Malaysian region. In S.K. Yap & S.W. Lee
(eds.), In Harmony with Nature. Proceedings of the International Conference of Tropical
Biodiversity. Kuala Lumpur: Malayan Nature Society. Pp. 8693.
Brunig, E.F. 1974. Ecological Studies of the Kerangas Forests of Sarawak and Brunei.
Kuching: Borneo Literature Bureau.
Corner, E.J.H. 1960. The Malayan flora. In R.D. Purchon (ed.), Proceedings of the
Centenary and Bicentenary Congress of Biology, Singapore. Pp. 2124.
Gobbett, D.J. & C.S. Hutchinson (eds.). 1973. Geology of the Malay Peninsula. New York:
Wiley.
Liechti, P., F.W. Roe & N.S. Haile. 1960. The Geology of Sarawak, Brunei and the Western
Part of North Borneo. British Borneo Geological Survey Department, Bulletin 3.
Richardson, J.A. 1947. An outline of the geomorphological evolution of British Malaya.
Geol. Mag. 84: 129144.
Steenis, C.G.G.J. van. 1957. Specific and infraspecific delimitation. Flora Malesiana 1, 5, 1:
CLXVII-CCXXIX.
Symington, C.F. 1943. Foresters' Manual of Dipterocarps. Malayan Forest Records 16.
Whitmore, T.C. 1984. Tropical Rain Forests of the Far East. (2nd edition). Oxford:
Clarendon.
LI
THE TREE FLORA PROPER
ACERACEAE
A. Noorsiha
Kepong, Malaysia
Koorders & Valeton, Bijdr. Booms. Java 9 (1903) 252; Bloembergen, FM 1, 4 (1948) 3; Backer &
Bakhuizen f., FJ 2 (1965) 143; Keng, OFMSP (1969) 416; Whitmore, TFM 2 (1973) 1; Anderson,
CLTS (1980) 133; Cockburn, TS 2 (1980) 14; Ashton, MNDTS 2 (1988) 1; Whitmore, Tantra &
Sutisna, CLK 1 (1989) 11.
Trees or shrubs. Buds with protective scales. Leaves opposite-decussate, simple, palmately
or pinnately veined, without stipules. Flowers in fascicles, panicles, racemes or corymbs,
regular, often unisexual; sepals and petals 45, rarely without petals; disc annular or lobed,
or reduced to teeth, rarely absent, intra- or extra-staminal; stamens (or staminodes) (4)8(
10), hypogynous or perigynous, filaments free; ovary often present in rudimentary form in
male flower, in female flowers 2-locular, superior, compressed, styles 2, free or connate at
base, ovules 2 in each locule, placentation axile. Fruit composed of 2 one-seeded, onewinged samaras. Seeds without endosperm; embryo with folded cotyledons.
Distribution. Two genera (Acer and Dipteronia) with about 200250 species distributed
mainly in mixed deciduous forests of the northern temperate zone. Only one genus (Acer)
with one species in western Malesia.
Taxonomy. Closely allied to the Sapindaceae, from which it can be distinguished by its
opposite/decussate simple leaves, samara-type fruit, and seed without endosperm.
ACER L.
(Latin name of the maple tree)
Sp. Pl. 1 (1753) 1054; Koorders & Valeton l.c. 253; Bloembergen l.c. 3; Backer & Bakhuizen f. l.c.
143; Whitmore l.c. 1; Anderson l.c. 133; Cockburn l.c. 14; Ashton l.c. 1; Whitmore, Tantra & Sutisna
l.c. 11.
Deciduous trees or shrubs with smooth bark. Leaves pinnately veined, more or less
distinctly 3-veined at the base, long-stalked. Flowers unisexual, male and female on the
same or different trees, organised in terminal or axillary racemes, corymbs, or panicles;
sepals and petals imbricate in buds. Other floral, fruit and seed characters as in the family.
Distribution. As for the family.
Ecology. The samaras are wind-dispersed.
ACERACEAE (NOORSIHA)
Uses. Many temperate species yield valuable timbers and a few others produce sugar/maple
syrup obtained by boring holes through the bark in February and March each year. A
number of shrubby species make excellent ornamental plants because of their strikingly
coloured (yellow, red and purple) aging foliage. The timber of the Malesian species is only
occasionally used for light construction.
Taxonomy. In a number of earlier publications (e.g., Blume, Rumphia 3 (1847) 193;
Miquel, Fl. Ind. Bat. 1, 2 (1859) 581; Bentham & Hooker f., Gen. Pl. 1 (1867) 409; Hooker
f., Fl. Brit. Ind. 1 (1875) 692) the genus was included in the Sapindaceae.
Acer laurinum Hassk.
Fig. 1.
(Latin, laurinum = resembling Laurus; the blue-grey colour of the leaves)

in Hoeven & de Vriese, Tijd. Nat. Gesch. & Phys. 10 (1843) 138; van Steenis, FM 1, 4 (1954) 592;
Backer & Bakhuizen f. l.c. 143; Anderson l.c. 133; Cockburn l.c. 14; Delendick, Reinwardtia 9, 4
(1980) 395; Ashton l.c. 1; Whitmore, Tantra & Sutisna l.c. 11. Type: Junghuhn, s.n., West Java,
Megamendong (holotype L; isotype BO). Synonyms: A. javanicum Jungh. apud Hoeven & de Vriese,
Monatsber. Verh. Ges. Erdk. Berlin 3 (1842) 96; A. niveum Blume, Jaarb. Kon. Ned Maatsch. Aanm.
Tuin (1844) 84; A. caesiaefolium Blume, Rumphia 3 (1847) 193; A. philippinum Merr., Philip. Govt.
Lab. Bull. 35 (1909) 36; A. curranii Merr., Philip. J. Sc. 4 (1909) 285; A. caesium (Reinw. ex Blume)
Kosterm., Reinwardtia 7 (1965) 141 (based on Laurus caesia Reinw. ex Blume, Bijdr. (1826) 553);
non A. caesium Wall. ex Brandis, For. Fl. NW. & C. Ind. (1874) 3).
Medium-sized to large tree up to 50 m tall; clear bole straight, cylindrical, up to 28 m tall

and 70 cm diameter; buttresses about 1.5 m high and 2 m out, thin, spreading, slightly
concave; crown dense, pale fresh green from below, more or less conical to hemispherical,
deciduous to semi-evergreen. Bark initially smooth, greenish grey, becoming rich red to
grey-brown and longitudinally fissured and flaky with age; inner bark c. 1.5 cm thick, firmly
fibrous, yellow-brown to red-brown, somewhat mottled, sometimes laminated. Heartwood
absent; sapwood white to pale yellow, with rather distinct concentric growth rings. Twigs
slender, terete or ribbed, drying nearly black, with numerous leaf-scars and minute lenticels.
Buds numerous, c. 4 mm long, covered by 411 pairs of decussate, caducous scales of c. 2
mm long. Leaves thinly leathery, glabrous, elliptic to ovate-lanceolate, 723 x 36 cm,
glaucous to grey beneath, upper surface dark green when fresh, drying dull red-brown; base
rounded to obtuse, margin entire, apex acute to acuminate-caudate; midrib slender, slightly
raised above, more or less prominent beneath; lateral veins 49 pairs, very slender, with a
pair at the base of the midrib; intercostal veins fine, reticulate; stalk 27.5 cm long, slender.
Inflorescences in the axils of fallen leaves, paniculate or corymbose, 310 cm long, 3050flowered; bracts and bracteoles fairly well-developed. Male flowers: sepals and petals (3
)5, free, obliquely erect, respectively 2.53 and 1.52.5 mm long; stamens (4)6(8),
arranged in one whorl on a flat, glabrous to woolly disc, filaments c. 5 mm long, anthers c. 1
mm long; pistillode strongly reduced. Female flowers with sepals and petals as in the
males; staminodes strongly reduced; ovary densely hairy, c. 2 mm across; styles 2, c. 1.5 mm
long, stigmas 2, sessile. Fruits red, wings 47.5 x 12.5 cm, asymmetrical, narrowed towards
the base, pubescent. Young seedling leaves coarsely and distantly toothed; sapling leaves
whorled.
Vernacular name. Sarawakperdu (Malay).
Distribution. Burma, possibly Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, the
Phillipines, Celebes and Lesser Sunda Islands. Rare in truly non-seasonal parts of
Peninsular Malaysia, Sumatra and Borneo, but relatively common elsewhere. In Sabah
uncommon, known only from 2 collections from Mt. Kinabalu (SAN 38438) and Nabawan
(SAN 124720). In Sarawak, it has a very local distribution, but is frequent where it occurs,
e.g., in the Hose Mts. (3rd Div.), Mt. Meluku (2nd Div.), and Usun Apau Highlands (4th
Div.).
Ecology. Apparently confined to soils of relatively high nutrient status, on igneous rocks at
2001500 m in the upper limits of Mixed Dipterocarp Forest and on granodiorite rocks in
the oak-laurel lower montane forests around 12001600 m. Flowering has been recorded in
AprilAugust and fruiting in JulyNovember. In Sabah, fruiting has been rarely recorded.
ALANGIACEAE
A. Berhaman
Sandakan, Malaysia
Merrill, PEB (1929) 232; Masamune, EPB (1942) 517; Backer & Bakhuizen f., FJ 2 (1965) 159; Eyde,
J. Arn. Arb. 49 (1968) 167; Kochummen, TFM 1 (1972) 56; Anderson, CLTS (1980) 133; Cockburn,
TS 2 (1980) 15; Ashton, MNDTS 2 (1988) 4; Whitmore, Tantra & Sutisna, CLK 1 (1989) 11.
The family consists of only one genus, Alangium, distributed in tropical Africa, and E Asia,
Malesia, eastern Australia and some South Pacific Islands.
Taxonomy. Alangium has in the past been included in the Cornaceae. Eyde l.c. investigating
the anatomy of the flower and discussing the relationships of Alangium, concluded that this
genus was not closely related to the Cornaceae. Instead, he suggested that the alkaloid
characteristics and floral morphology indicate that Alangium has its closest relatives in the
Rubiaceae. The evidences have not been further examined and the most recent treatments
accept the Alangiaceae as a monogeneric family, usually placed in the Cornales.
ALANGIUM Lam.
(after the Malabar plant name, alangi)
kondolon (Dusun, Sabah), midong (Iban, Sarawak)
Encycl. Meth. Bot. 1 (1783) 174; Merrill, EB (1921) 459 (as part of Cornaceae), l.c. (1929) 232;
Bloembergen, Blumea 1, 2 (1935) 241, Bull. Jard. Bot. Btzg. 3, 16 (1939) 139; Masamune l.c. 517;
Kochummen l.c. 56; Anderson l.c. 133; Cockburn l.c. 15; Ashton l.c. 4; Whitmore, Tantra & Sutisna
l.c. 12; Berhaman, Sandakania 4 (1994) 31.
Trees, more rarely shrubs or woody climbers, often with thin buttresses or stilt-roots. Bark
smooth, thin, grey-brown, often with pale lichen patches; inner bark thick, yellowish brown.
Leaves simple, alternate, entire, pinnately veined, or sometimes 35-veined at the base;
stalk terete or slightly grooved or flattened on upper side; stipules none. Inflorescence an
axillary cyme, few- to many-flowered. Flowers bisexual, sessile or stalked (the stalks, if
present, articulate); calyx-tube connate with the ovary wall, truncate or with 410 teeth or
lobes; petals 410, valvate, linear, becoming recurved, alternate with the calyx-teeth,
usually white; stamens as many as or 26-times the number of petals; anthers linear,
dehiscing laterally or introrsely; disc intrastaminal, well-developed; ovary inferior, 12celled, style single, usually as long as the corolla, ovule 1 in each locule, anatropous. Fruit a
drupe, globose to ellipsoid and somewhat bilaterally compressed, often longitudinally
ribbed, crowned by the persistent calyx-teeth and disc. Seed 12 per fruit with a copious
endosperm; cotyledons foliaceous, flat, palmately veined at base; radicle straight.
Distribution. 21 species, distributed as noted for the family. 10 species in Sabah and
Sarawak, including A. kurzii Craib, a new record for Borneo.
Ecology. Primary and secondary forests, lowlands to 1500 m.
Taxonomy. Bloembergen l.c. (1939) recognised four sections, Angolam Baill. (which Eyde
l.c. points out should be called section Alangium as it contains the type species), Marlea
Baill., Rhytidandra Baill., and Conostigma Bloemb.; all except Rhytidandra are represented in Sabah and Sarawak. Eyde l.c. has found that differences in style and stigma structure,
stamen number, the pattern of floral vasculature, pollen morphology, and fruit endocarp
characteristics support Bloembergen's designation of the four sections.
Key to Alangium species

1.
Woody climbers......................................................................................................................2
Trees........................................................................................................................................3
2.
Leaf coriaceous with domatia in the axils of main veins. Inflorescence almost sessile,
with less than 15 flowers...............................................................................................
Alangium sp. 1.
Woody climber. Leaves obovate-elliptic, 1215 x 4.56 cm, coriaceous; base 3veined, cuneate; apex acuminate; lateral veins 45 pairs; stalk 1.21.5 cm long,
glabrous.
Only one specimen, SAN 79310, from Telupid in Sabah known, with young flower
buds.
Leaf thinly chartaceous, without domatia. Inflorescence with stalk up to 4 cm long, and
1730 flowers................................................................................................................
A. scandens Bloemb.
l.c. (1935) 264, l.c. (1939) 193; Masamune l.c. 518; Whitmore, Tantra & Sutisna l.c. 12;
Berhaman l.c. 32. Type: Endert 4076, Sarawak, 4th Division, Long Petah (lectotype BO).
Woody climber. Leaves ovate to ovate-elliptic, 815.5 x 3.56 cm, chartaceous to

thinly coriaceous; base subcordate to rounded, apex acuminate; lateral veins 67
pairs; stalk hairy, 11.5 cm long.
Sumatra and Borneo. Hill to lower montane forest, common in Sabah and Sarawak.
3.
Leaf base 35-veined...................................................................................................4

Leaf base pinnately veined...........................................................................................9
4.
Leaves markedly obovate with narrowed base, or if elliptic and base rounded then
lateral veins 67 pairs........................................................................6. A. longiflorum
Leaves rounded to broadly ovate with rounded to cordate base, or if (rarely) elliptic and
base narrowed then lateral veins 45 pairs...................................................................5
5.
Leaf length at least twice the width.........................................................2. A. griffithii

Leaf length not reaching twice the width.....................................................................6
6
ALANGIACEAE (BERHAMAN)
6.
Leaf-stalk densely hairy; lateral veins 912 pairs. Inflorescence-stalk less than 8 mm
long. Flowers almost sessile........................................................................7. A. nobile
Leaf-stalk glabrous to sparsely hairy; lateral veins less than 8 pairs. Inflorescence-stalk
more than 10 mm long. Flowers distinctly stalked ......................................................7
7.
Leaf-apex blunt to emarginate; lateral veins 45 pairs. Petals c. 28 mm long and 5 mm

wide at base, covered by tiny stellate scales............................................1. A. circulare
Leaf-apex acute to acuminate; lateral veins 57 pairs. Petals less than 15 mm long and
12 mm wide at base, covered by straight hairs..........................................................8
8.
Leaf lower surface velvety (seldom sparsely) hairy; midrib and lateral veins on upper
surface densely hairy. Calyx-tube densely hairy, teeth to only about 0.5 mm long
..................................................................................................................5. A. kurzii
Leaf lower surface glabrous to sparsely hairy, never velvety; midrib and lateral veins on
upper surface glabrous. Calyx-tube sparsely hairy, teeth more than 1 mm long
.......................................................................................................8. A. rotundifolium
9.
Plants of mixed swamp forest. Leaf-base typically asymmetric. Calyx-limb with distinct
triangular teeth. Style thickly yellow hairy. Stamen filaments broadened and thickly
woolly hairy at base..............................................................................3. A. havilandii
Plants of lowland mixed dipterocarp forest. Leaf-base typically symmetric (rarely asymmetric). Calyx-limb subtruncate. Style sparsely white-hairy. Stamen filaments uniformly thick throughout, subglabrous to sparsely short-hairy at base....4. A. javanicum
1. Alangium circulare Stone & Kochummen

(Latin, circularis = circular; the round-shaped leaves)
Blumea 22 (1975) 219; Ashton l.c. 4. Type: Salleh ak Nantah S. 24325, Sarawak, 1st Division, Bukit
Siol, Kuching (holotype SAR; isotypes A, BO, K, KEP, L, SAN, SING).
Small to medium-sized tree to 15 m tall. Bark smooth, grey-brown; inner bark pale brown.
Sapwood pale yellow. Twigs grey to brown. Leaves broadly ovate-elliptic-obovate to
subcircular, 8.510.5 x 810 cm, coriaceous; base rounded to cordate, 3-veined, apex blunt
to emarginate; midrib raised on both surfaces; lateral veins 45 pairs, slightly raised on
both surfaces, glabrous; stalk glabrous, 710 mm long, 23 mm diameter, grooved on upper
surface. Inflorescence a short cyme, 12-times branched, stalk 1114 mm long, finely
greyish pubescent, 13-flowered. Flowers 5-merous, subsessile to shortly stalked, stalks
23 mm long; calyx funnel-shaped, 34 mm long, 3 mm across, finely stellate-pubescent,
teeth 5, triangular, to 1 mm high; petals linear-lanceolate, c. 28 mm long, c. 5 mm wide at
base, abaxial side covered with minute stellate hairs; stamens as many as petals, linear,
1011 mm long; ovary 1-celled, style hairy, 2226 mm long. Fruit and seed unknown.
Distribution. Endemic to Sarawak, recorded from Bukit Siol, Sempadi Forest Reserve and
Gunung Pueh in the 1st Division. Apparently a species of kerangas forest.
2. Alangium griffithii (Clarke) Harms

(W. Griffith, 18101845, doctor and botanist in India and Malacca)
7
in Engl. & Prantl., Pfl. Fam. 3, 8 (1898) 262; Bloembergen l.c. (1935) 266, l.c. (1939) 194; Masamune
l.c. 517; Kochummen l.c. 58; Cockburn l.c. 16; Anderson l.c. 134; Ashton l.c. 5; Whitmore, Tantra &
Sutisna l.c. 11; Berhaman l.c. 32. Basionym: Marlea griffithii Clarke in Hooker f., Fl. Brit. Ind. 2
(1879) 742. Type: Griffith 3387, Malacca (lectotype K; isolectotypes B, BM). Synonym: Marlea
densiflora Koord. & Valeton, Bijdr. Booms. Java 5 (1899) 84.
Medium-sized tree to 20 m tall, 30 cm diameter, sometimes with spreading buttresses to 50

cm wide. Bark smooth, grey to dark brown; inner bark pale yellow. Sapwood white to pale
yellow. Twigs dark brown, smooth to hairy. Leaves chartaceous, elliptic to narrowly ovate,
5.518.5 x 28 cm, length at least twice the width, hairy on the lower surface, mainly on the
main veins, grey to dark brown when dry; base cuneate, asymmetric, 35-veined, apex
acuminate; midrib flat on upper side, hairy on the lower surface; lateral veins 45(6) pairs,
usually glabrous between the veins on upper surface; stalk 515 mm long, sparse to densely
hairy. Inflorescence hairy, 34-times branched, 36.5 cm long, many-flowered, stalk 12.2
cm long. Flowers 5-merous, white to cream, fragrant; calyx-tube densely hairy, with teeth
0.250.5 mm long, limb spreading; petals linear, swollen at base, glabrous to sparsely hairy
outside, cream, 8.515 mm long; stamens as many as petals, 814 mm long; ovary 1-celled,
style glabrous, cream, 811 mm long. Fruit ovoid, 1421 x 810 mm, dark brown when
dry, faintly grooved, crowned by the persistent calyx-limb, dark blue when ripe.
Vernacular names. Sabahgadong hutan (Brunei Malay). Bruneimayam kampong
(Malay).
Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, Java, and Borneo. In
Sabah, quite common but in Sarawak only found in the Tinjar Forest Reserve. Also in
Brunei and Kalimantan.
Ecology. Primary and secondary forest, 60400 m.
3. Alangium havilandii Bloemb.

(J.D. Haviland, 18571901, first Sarawak Medical Officer and plant and insect collector)
l.c. (1935) 277, l.c. (1939) 213; Masamune l.c. 518; Anderson l.c. 134; Ashton l.c. 8; Whitmore,
Tantra & Sutisna l.c. 11. Type: Omar 54, Sarawak, 1st Division, Gunung Sedilu Forest Reserve
(holotype SING).
Small to medium-sized tree to 25 m tall, 30 cm diameter, with tall prominent thin flying
buttresses. Bark smooth, greyish; inner bark yellow. Sapwood pale whitish to yellowish.
Leaves ovate to elliptic, 515 x 3.56 cm, chartaceous or thinly coriaceous; base typically
asymmetric, rarely symmetric, pinnately veined, apex long acuminate to acute; midrib flat
on the upper surface; lateral veins, 69 pairs, raised on the lower surface; stalk 512 mm
long, slender. Inflorescence a short cyme, 12-times branched, 16-flowered, stalk 11.5
cm long. Flowers 45-merous; calyx-tube tomentose, c. 2 mm long, limb with distinct
triangular teeth; petals 1520 mm long, prominently dilated at base; stamens as many as
petals, 1417 mm long, filaments broadened and thickly woolly hairy at base; ovary 1celled, style thickly yellow hairy, 1113 mm long. Fruit ellipsoid-ovoid, finely hairy, 10.5
18 x 69 mm, flattened when dried, crowned by the persistent calyx-limb, ripening pink.
8
Vernacular names. Sarawakdadam or jadam paya (Melanau), jenangan (Melanau,

Bintulu), sisit (Malay).
Distribution. Sarawak, Brunei and Kalimantan, not yet recorded in Sabah.
Ecology. Frequent and locally common in mixed peat swamp forest at low altitude.
4. Alangium javanicum (Blume) Wangerin
Fig. 1.
(of Java)
in Engl. & Prantl., Pfl. Fam. 4, 220b (1910) 14; Bloembergen l.c. (1935) 281, l.c. (1939) 218; Merrill
l.c. (1921) 458, l.c. (1929) 232, in syn.; Masamune l.c. 518; Cockburn l.c. 16, in syn.; Kochummen l.c.
57, in syn.; Ashton l.c. 10; Anderson l.c. 133, in syn.; Whitmore, Tantra & Sutisna l.c. 12; Berhaman
l.c. 33. Basionym: Styrax javanicum Blume, Bijdr. 13 (1825) 671. Type: Blume, s.n., Java (holotype
BO; isotype NY).
Medium-sized tree to 30 m tall and 35 cm diameter, often with flying buttresses. Bark
smooth, yellowish to pale brown; inner bark yellowish to reddish. Sapwood yellowish.
Leaves ovate-elliptic to obovate, (8)15(35) x (2.5)5.5(15.5) cm, chartaceous to
coriaceous, lower surface glabrous to short-hairy on the midrib, or sometimes short velvety
all over, drying pale olive-brown to purplish brown; base cuneate to rounded, symmetric
(rarely asymmetric), pinnately veined, apex acute to acuminate; midrib flat to raised, rarely
sunken on the upper surface; lateral veins 69(19) pairs; stalk 0.53.8 cm long, slender,
sometimes grooved on the upper surface and densely hairy. Inflorescence 13-times
branched, with up to 35 flowers, stalk 28 mm long. Flowers (4)6(7)-merous, 22.5 cm
long; calyx-tube campanulate, limb subtruncate, glabrous to densely covered by silky long
hairs; corolla usually swollen at base, densely covered by long silky hairs or densely or
sparsely stellate-hairy; petals 8.517 mm long, stamens as many as petals, filaments
uniformly thick throughout, subglabrous to sparsely short-hairy at base; ovary 1-celled,
style 410 mm long, sparsely white-hairy. Fruit ellipsoid-ovoid, variable in size, smooth to
strongly ridged, crowned by the persistent calyx, 15 mm high and 210 mm wide.
Vernacular names. Sabahsatu inchi (colloquial Malay). Sarawakjadam (Malay).
Key to varieties
1.
Corolla densely covered by silky long hairs (velutinous). Lower side of leaf short-hairy
on midrib only...............................................................................................................
var. javanicum
Synonyms: A. bogoriense Wangerin, Fedde. Repert. 4 (1907) 338; A. borneense Merr., J.
Str. Br. R. As. Soc. 86 (1922) 10; A. javanicum "form B" of Ashton l.c. 11; A. ebenaceum
"var. C" of Cockburn l.c. 18.
Sumatra, Java and Borneo. Common in all districts in Sabah and Sarawak. Primary
mixed dipterocarp and secondary forest.
Corolla sparsely to densely covered with stellate hairs, not long-hairy. Lower side of
leaf glabrous, or sometimes short-velvety all over........................................................2
9
2.
Leaves drying pale olive-brown, typically smaller, 812 cm long, but exceptionally
reaching 27 cm long. Calyx-limb of mature fruit smaller, 12 mm high and 25 mm
wide..............................................................................................................................
var. meyeri (Merr.) Berhaman
l.c. 34. Basionym: A. meyeri Merr., Publ. Govern. Labor. 35 (1906) 54. Type: Meyer F.B.
2284, Phillipines, Luzon, Cagayan Province (lectotype BO; isolectotypes B, K, NY, S).
Synonyms: A. tutela Ridl., J. Str. Br. R. As. Soc. 61 (1912) 10; A. ebenaceum var. tutela
(Ridl.) Kochummen, Fed. Mus. J. 13 (1970) 133; A. javanicum "form C" (pro parte) &
"form D" of Ashton l.c. 11; A. ebenaceum "var. B" (pro parte), "var. D", "var. E", and "var.
G" of Cockburn l.c. 18 (pro parte).
Peninsular Malaysia, Java, Borneo, and the Philippines. Very common in all districts
in Sabah and Sarawak, in primary and secondary mixed dipterocarp forest.
Leaves drying purplish brown, (at least markedly so on the lower side), typically longer
or larger (2439 cm long). Calyx-limb of mature fruit larger, 25 mm high and 310
mm wide.......................................................................................................................
var. ebenaceum (Clarke) Berhaman
l.c. 33. Basionym: Marlea ebenacea Clarke in Hooker f., Fl. Brit. Ind. 2 (1879) 742. Type:
Griffith 3383, Malacca (lectotype K). Synonyms: A. ebenaceum (Clarke) Harms in Engl. &
Prantl l.c. (1898) 262; A. ridleyi sensu Ashton l.c. 14, non King (1902); A. mezianum
Wangerin, l.c. 338; A. sessiliflorum Merr. l.c. (1929) 232; A. javanicum "form C" of Ashton,
l.c. 11 (pro parte); A. ebenaceum "var. B" (pro parte) and "var. G" of Cockburn l.c. 18 (pro
parte).
Peninsular Malaysia, Java, Borneo, and the Philippines. Common in all districts in
Sabah and Sarawak. All these forms appear to be confined to clay-rich soils in
mixed dipterocarp and associated secondary forest.
5. Alangium kurzii Craib

(W.S. Kurz, 18341878, German soldier and naturalist, Bogor and Calcutta)
Kew Bull. (1911) 60; Bloembergen l.c. (1935) 262, l.c. (1939) 183; Kochummen l.c. 58; Berhaman l.c.
31. Type: Kerr 1172, Thailand, Chiangmai, Doi Sutep (holotype K). Synonyms: Alangium chinensis
var. tomentosum Merr., Philip. J. Sci., 21, 5 (1922) 505; Alangium begoniaefolium Ridl., FMP 1
(1922) 894.
Small tree to 15 m tall, 20 cm diameter, with short buttresses. Bark smooth, dark grey,
lenticellate; inner bark orange and cream mottled. Sapwood soft, pale brown. Leaves thin
coriaceous to chartaceous, lower surface velvety (seldom sparsely) hairy, ovate to broadly
ovate, 820 x 510.5 cm; base markedly asymmetric, broadly rounded to cordate, 35veined, apex acuminate-acute; midrib and major veins on upper surface densely hairy;
10
11
lateral veins 57 pairs; stalk 2.53 cm long, glabrous to sparsely hairy. Inflorescence
hairy, 24-times branched with 315 flowers per branch, stalk 1115 mm long. Flowers
pale greenish to yellow creamy; calyx-tube densely hairy, 1.252.5 mm long, teeth hairy,
0.250.5 mm long; petals (5)9(10), 5.510 mm long, swollen at base for 23 mm,
covered by straight hairs; stamens as many as petals, 510 mm long, densely covered with
long straight hairs; ovary (1)2-celled, style glabrous, 58 mm long. Fruit ellipsoid 11.5 x
0.50.7 cm, crowned with the persistent calyx-limb.
Vernacular name. Sabahmarapangi (Dusun Tambunan).
Distribution. Myammar (Burma), China, Indo-China, Thailand, Sumatra, Peninsular
Malaysia, Java, and Borneo. In Sabah, recorded only from Kota Marudu (SAN 99531),
Ranau (SAN 62010) and Tambunan (SAN 11355). Not recorded for Sarawak.
Ecology. Lowland to submontane forest to 1300 m. Flowering from January to July; fruiting
from May to August.
6. Alangium longiflorum Merr.

(Latin, longus = long, florum = flowers)
Philip. J. Sci. Bot. 7 (1912) 319; Bloembergen l.c. (1935) 253, l.c. (1939) 159; Cockburn l.c. 16;
Anderson l.c. 134; Ashton l.c. 11; Whitmore, Tantra & Sutisna l.c. 12; Berhaman l.c. 32. Type:
Darling FB 14773, Philippines, Malueg, Cagayan Province (holotype L; isotype BO). Synonyms:
Alangium salvifolium subsp. hexapetalum sensu Merr. l.c. (1912) 321; A. lamarckii Thwaites, Enum.
Pl. Zeyl. (1850) 133; A. hirsutum Bloemb. l.c. (1939) 161.
Medium-sized tree to 20 m tall and 25 cm diameter. Bark smooth, dark brown; inner bark
yellowish. Sapwood white. Leaves markedly obovate with narrowed base and 47 pairs of
lateral veins or if elliptic and base rounded then with 67 pairs of lateral veins, velvety
hairy on lower surface, 812 x 2.58 cm, chartaceous; base 3-veined, apex long-acuminate;
midrib raised and sparsely hairy on the upper surface; stalk slender, hairy, 813 mm long.
Inflorescence a short cyme with 15 flowers, densely hairy, almost sessile or with stalk to
1.5 mm long. Flowers white, 3050 mm long; calyx-tube campanulate, densely hairy, teeth
0.250.5 mm long, limb 11.5 mm long; petals 5, white, 3550 mm long, densely hairy
outside, swollen at the base; stamens 3050 mm long, 26-times as many as petals; ovary 1celled, style glabrous, 3045 mm long. Fruit ellipsoid to ovoid in dry state, rounded at both
ends, 2025 cm long, crowned by the persistent calyx-limb, ripening pinkish.
Distribution. Borneo and Philippines. Very uncommon in Sabah (e.g., SAN 30390, SAN
83891) and Sarawak (e.g., S. 21704, S. 43342), also known in Kalimantan.
Ecology. Primary mixed dipterocarp and associated secondary forests to 1200 m, on high
nutrient clay soils. Flowering in March to July; fruiting in August to December.
One specimen, SAN 14167 (Sabah, Ranau) bears spines, but the leaf characters are very
close to that for A. longiflorum.
12
7. Alangium nobile (Clarke) Harms

(Latin, nobilis = distinguished, noble, probably referring to the growth habit)
in Engl. & Prantl l.c. (1898) 262; Bloembergen l.c. (1935) 275, l.c. (1939) 211; Masamune l.c. 518;
Kochummen l.c. 58; Anderson l.c. 134; Ashton l.c. 12; Whitmore, Tantra & Sutisna l.c. 12; Berhaman
l.c. 32. Basionym: Marlea nobilis Clarke in Hooker f. l.c. 743; Ridley, Agr. Bull. Str. Settl. & Fed.
Mal. St. 1 (1902) 181. Type: Griffith 3385, Malacca (lectotype K; isolectotypes B, BM).
Medium-sized tree to 20 m tall, 30 cm diameter. Bark smooth, brown; inner bark thin,
yellow to pale brownish. Sapwood medium-hard, whitish. Leaves rounded to broadly
obovate, 923 x 518 cm, chartaceous to coriaceous, upper surface hairy to glabrous, lower
surface hairy to densely hairy; base rounded to cordate, 5-veined, apex rounded to acute,
rarely acute to acuminate; midrib flattened to raised on the upper surface, densely hairy on
the lower surface; lateral veins 912 pairs, raised and hairy below, flattened on the upper
surface; stalk densely hairy, 2.54.5 cm long and 35 mm thick. Inflorescence a short
cyme, stalk 57 mm long, 12-branched with 15 flowers. Flowers 45-merous, almost
sessile; calyx-tube campanulate, hairy, lobes 57; petals swollen at base, 1015 mm long,
densely hairy outside; stamens as many as petals, 914 mm long, with filaments broadened
at base; ovary 1-celled, style hairy, 712 mm long. Fruit ellipsoid-ovoid, 2530 x 1520
mm, grooved, velvety hairy, crowned by the persistent calyx-limb.
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah and Sarawak, uncommon
in mixed dipterocarp forest (e.g., SAN 73205 and S. 38479).
Ecology. Lowland to submontane forest to 1500 m.
8. Alangium rotundifolium (Hassk.) Bloemb.

(Latin, rotundus = rounded in outline, folium = leaves)
l.c. (1935) 258, l.c. (1939) 179; Masamune l.c. 518; Kochummen l.c. 60; Cockburn l.c. 16; Whitmore,
Tantra & Sutisna l.c. 12; Berhaman l.c. 38. Basionym: Diacaecarpium rotundifolium Hassk.,
Bonplandia 7 (1859) 172. Type: Sine coll., s.n., Java, probably from Tjibodas, Cult. Bogor Botanic
Garden (holotype BO; isotype L). Synonyms: Marlea rotundifolia (Hassk.) Teijsm. & Binn., Cat. Pl.
Hort. Bot. Bogor. (1866) 238; Alangium begoniifolium Harms in Engl. & Prantl l.c (1898) 261;
Marlea begonifolia (Harms) Ridl., J. Fed. Malay. St. Mus., 8, 4 (1917) 44; Alangium rotundatum Ridl.
ex Burkill & Henders., Gard. Bull. Str. Settl. 3 (1925) 380.
Tree to 28 m tall, 40 cm diameter. Bark smooth to slightly scaly, white to pale grey; inner
bark yellowish. Sapwood white to pale yellow. Twig dark brown to pale brown. Leaves
glabrous to sparsely hairy on the lower surface, never velvety, rounded to broadly ovate or
triangular-ovate, 9.522 x 7.514 cm, chartaceous; base rounded to cordate, sometimes
cuneate, 35-veined, apex acuminate to acute; midrib and lateral veins on upper surface
glabrous; lateral veins 57 pairs; stalk 2.53.5 cm long, glabrous to sparsely hairy.
Inflorescence a cyme, 34-times branched with 415 flowers, covered by short hairs, stalk
1220 mm long. Flowers fragrant, white or cream to yellow; calyx-tube sparsely hairy,
1.53 mm long, limb flared with teeth 11.5 mm long; corolla swollen at the base; petals
white or cream, 614(18) mm long; stamens as many as petals, 618 mm long; ovary
usually 1-celled, rarely 2-celled, style glabrous, 515 mm long. Fruit glabrous or thinly
hairy, ovate-ellipsoid, cuneate or rounded at base, crowned by the flared persistent calyxlimb 11.5 mm long, reddish when ripe.
13
Vernacular name. Sabahmarapangi (Dusun Ranau).

Distribution. Sumatra, Peninsular Malaysia, Java, and Borneo. In Sabah only recorded from
Mt. Kinabalu (e.g., SAN 46772, SAN 48064) and Crocker Ranges (SAN 83987). Not
recorded for Sarawak.
Ecology. Primary and secondary forest on ultramafic soils, at 4001600 m.
Excluded species
A. kinabaluense W.W. Sm., Not. Bot. Gard. Edinb. 8 (1915) 315; Merill l.c. (1921) 459, is
based on a specimen collected from Mt. Kinabalu, Native Coll. 49 (E, K), which is
Polyosma hookeri Stapf.
14
ANISOPHYLLEACEAE
K.M. Wong & L. Madani
Sandakan, Malaysia
Merrill, EB (1921) 420 (as part of Rhizophoraceae); Ridley, FMP 1 (1922) 692 (as Anisophylleae at
family rank); Masamune, EPB (1942) 515 (as part of Rhizophoraceae); Ding Hou, FM 1, 5 (1958) 429
(as part of Rhizophoraceae); Burgess, TBS (1966) 431 (as part of Rhizophoraceae); Corner, WSTM 1
(1988) 131 (as Anisophylleaceae); Ashton, MNDTS 2 (1988) 342 (as part of Rhizophoraceae); Juncosa &
Tomlinson, Ann. Missouri Bot. Gard. 75 (1988) 1278 (as Anisophylleaceae); Kochummen, TFM 4 (1989)
302 (as part of Rhizophoraceae); Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 295 (as part of
Rhizophoraceae).
Trees or shrubs, very rarely (in Combretocarpus) with small stilt-roots or pneumatophore
(breathing) roots. Bark invaginating finely and regularly into wood in most species.
Branches highly differentiated architecturally from the trunk, twigs solid, branch nodes not
swollen; supernumerary axillary buds present (in Anisophyllea) or not. Leaves alternate (2ranked in Combretocarpus, basically 4-ranked in Anisophyllea), simple, often (in Anisophyllea)
dimorphic, 37-veined from the base or (in Combretocarpus) pinnately veined; stipules
none. Inflorescence fundamentally paniculate or racemose, axillary, multiflorous, bisexual
or (rarely) unisexual. Flower unisexual (Anisophyllea, but plants monoecious) or bisexual
(Combretocarpus), radially symmetrical, 35-merous; calyx valvate; petals free, lobed or
laciniate, rarely entire; stamens free or (in Anisophyllea) epipetalous and episepalous, 2times the number of petals, anthers 4-celled, dorsifixed, splitting lengthwise; ovary inferior,
35-locular, ovules 12 per carpel, style single or (mostly) several and free; disc annular,
lobed. Fruit a drupe or (in Combretocarpus) a dry, 3-winged structure. Seed one; without
endosperm; embryo a solid structure with reduced or no cotyledons; germination hypogeal.
Distribution. 4 genera and c. 35 species, tropical Africa to S America and Malesia. In
Sabah and Sarawak, 2 genera, Anisophyllea and Combretocarpus, with 10 species.
Ecology. Combretocarpus is found mainly in peat-swamp forest, and Anisophyllea species
occur mainly in primary moist lowland forest and sometimes in secondary lowland forest.
Uses. Several Anisophyllea species and Combretocarpus grow to timber size and are exploited
for logs.
Taxonomy. The members of the Anisophylleaceae have traditionally been treated as part of
the Rhizophoraceae. Ridley (l.c.) appears to have been the first to consider the group (only
Anisophyllea as was known then for Malaya) at family rank, equivalent to Bentham &
Hookers designation of the tribe Anisophylleae within the Rhizophoraceae in 1865 (in their
Genera Plantarum 1, 2). Although Melchior (1964) (Englers Syllabus der Pflanzen-
15
familien, 12th Edition, Vol. 2) and Cronquist (1981) (An Integrated System of Classification of
Flowering Plants) both adopted the Anisophylleaceae, this has not been widely adopted until
multidisciplinary studies demonstrated the distinctness of the Anisophylleaceae as a family
(Juncosa & Tomlinson l.c. and Dahlgren, Ann. Missouri Bot. Gard. 75 (1988) 1259). Basically,
the Anisophylleaceae is distinct from the Rhizophoraceae by the following characteristics:
alternate leaves, absence of stipules, simple (not scalariform) vessel perforation, mostly
paniculate or racemose (not cymose) inflorescences, inferior ovary with separate styles (the
Rhizophoraceae have a superior to inferior ovary and a single style), relatively thin pollen
wall tapetum, lack of subepidermal floral laticifers (in the ovary and calyx), and lack of
endosperm in the seed.
Key to genera
Leaves basically 4-ranked on lateral branches (although in many species the reduced leaves
of the two upper ranks are hard to notice or fall away early), with 37 veins from the leaf
base. Leaf axils each with several axillary buds in a row. Fruit a drupe........1. Anisophyllea
Leaves 2-ranked on lateral branches, pinnately veined. Leaf axils each with only one distinct
axillary bud. Fruit a 3-winged structure.....................................2. Combretocarpus
1. ANISOPHYLLEA R. Br. ex Sabine

(Greek, anisos = unequal, phullon = leaf; the dimorphic leaves)
menengang (Murut, Sarawak), mertama (Iban, Sarawak),
mopu (Bidayuh, Sarawak), sireh sireh (Malay, Sabah)
Trans. Hort. Soc. 5 (1824) 466; Merrill l.c. 422; Ridley l.c. 701; Masamune l.c. 515; Ding Hou l.c.
474; Corner l.c. 131; Anderson, CLTS (1980) 290; Ashton l.c. 344; Kochummen l.c. 304; Whitmore,
Tantra & Sutisna l.c. 295; Madani, Sandakania 3 (1993) 49. Synonyms: Anisophyllum G. Don ex
Hook., Niger Fl. (1849) 342, 575; Tetracrypta Gardn. & Champ. in Hooker, J. Bot. Kew Misc. 1
(1849) 314.
Trees and shrubs, trunk not buttressed but sometimes fluted at base. Bark smooth or
minutely cracked or flaky; inner bark invaginating finely and regularly into the wood.
Sapwood pale to reddish brown, with conspicuous pale radiating lines. Branches
developing on the young stem in widely separated tiers; axillary buds several in a series in
each leaf axil. Leaves basically 4-ranked, often dimorphic (but reduced leaves of the two
upper ranks hard to notice or fall away early in most species), 37-veined from the base.
Flower unisexual (but plants monoecious), 35-merous; petals entire, lobed or laciniate;
stamens epipetalous and episepalous, usually unequal in length; ovary 35-celled, ovule 1
per cell, styles 34, free. Fruit a drupe, ellipsoid or pear-shaped, smooth or ridged, fruitwall usually hard and fibrous (fleshy in A. disticha). Seed solitary, globose to ellipsoid.
Distribution. c. 30 species, tropical Africa, Sri Lanka, India, South East Asia, and West
Malesia (Sumatra, Peninsular Malaysia and Borneo). 9 species in Sabah and Sarawak.
16
ANISOPHYLLEACEAE (WONG & MADANI)
Ecology. Primary or old secondary lowland and hill forests to c. 1000 m.
Key to Anisophyllea species

1.
Leaves along branches of two markedly different sizes, arranged in four rows, the
larger ones on the two lower rows trapezoid.................................................................2
Leaves of only one type easily discernible, basically elliptic or ovate, arranged in two
rows (reduced leaves scantily developed and falling early, and usually not noticed)......3
2.
Larger leaves not longer than 33.5 cm. Fruiting racemes 0.52.7 cm long, typically
bearing a solitary fruit.................................................................................4. A. disticha
Larger leaves typically 510 cm long. Fruiting racemes 410 cm long, typically bearing
several fruits........................................................................................9. A. rhomboidea
3.
Lateral veins all arising from the very base of the leaf-blade.........................................4
One or both pairs of lateral veins arising from the midrib away from the base of the
blade............................................................................................................................6
4.
Intercostal veins on the lower leaf surface conspicuously raised, forming a fine tessellated
pattern.................................................................................................................3. A. corneri
Intercostal veins on the lower leaf surface not conspicuously raised, forming a loose
reticulation and not distinctly tessellated......................................................................5
5.
Leaves coriaceous, broadly ovate, upper surface drying glossy, the lateral veins raised.
Inflorescence axes persistently densely covered with velvety purple-brown hairs............
....................................................................................................................8. A. nitida
Leaves chartaceous, elliptic, upper surface drying dull or matt, the lateral veins sunken
or flat. Inflorescence axes subglabrous or only minutely pale-brown hairy......................
..................................................................................................................2. A. chartacea
6.
Leaves rusty hairy on lower surface, the remains of the hairs always visible with 10X
magnification, even on old leaves; intercostal veins on lower surface finely tessellate
.......................................................................................................................5. A. ferruginea
Leaves glabrous, intercostal veins on lower surface indistinct or forming only a loose
network............................................................................................................................7
7.
All veins, including the intercostal veins, on upper leaf surface sunken. Mature fruit
globose............................................................................................7. A. impressinervia
Veins flat or raised on upper leaf surface. Mature fruit ellipsoid or globose.................8
8.
Leaves thinly coriaceous; apex rather abruptly caudate; midrib conspicuously thicker
than lateral veins. Mature fruit ellipsoid................................................1. A. beccariana
Leaves chartaceous; apex acute; midrib similar to lateral veins in thickness. Mature fruit
globose..............................................................................................6. A. globosa
17
1. Anisophyllea beccariana Baill.

(Odoardo Beccari, Italian explorer and botanist, 1843-1920)
Adansonia 11 (1875) 311; Merrill l.c. 422; Masamune l.c. 515; Ding Hou l.c. 474; Ashton l.c. 345;
Whitmore, Tantra & Sutisna l.c. 295. Type: Beccari PB 1001, Sarawak (lectotype K, here chosen;
isolectotypes G, P).
Small to medium-sized tree to 25 m tall. Bark scrolled to scaly; inner bark brownish.
Sapwood yellowish. Leaves of one form mainly, in two distinct ranks along the branches,
elliptic or ovate, 7.510.5 x 35 cm, thinly coriaceous, drying matt or slightly glossy on
upper surface, lower side with scattered tiny but conspicuous dot-glands; base cuneate, apex
abruptly caudate; midrib conspicuously thicker than lateral veins, flat to slightly depressed
on the upper side; lateral veins (one or both pairs) arising from the midrib away from the
base of the blade, flat or elevated on upper side, elevated on lower side; intercostal veins
forming only a loose network or somewhat obscure; glabrous; stalk 48 mm long.
Inflorescence a raceme or basally branched panicle to 10 cm long, axes minutely palebrown pubescent. Flowers unisexual; calyx pubescent outside, lobes 4; petals 0.60.7 mm
long; stamens in male 0.50.7 mm long, replaced by staminodes (stamen rudiments) in the
female; pistil in female 0.50.7 mm long, with 4 styles, replaced by pistillode (pistil
rudiments) in the male. Fruit ellipsoid, c. 3 x 2.5 cm, smooth on the surface when dry.
Vernacular name. Sarawakpei (Kayan).
Distribution. CE Sumatra and Borneo. In Sabah, recorded from Lahad Datu and Sandakan
districts; in Sarawak from the Kuching and Bintulu districts. Also in Brunei and Kalimantan.
Ecology. Mixed dipterocarp forest on leached, often shallow humult soils, and kerangas
(heath) forest in the lowlands, to c. 570 m.
2. Anisophyllea chartacea Madani

(Latin, chartaceus = papery; the leaves)
Sandakania 3 (1993) 51. Type: Othman S. 29055, Sarawak, 3rd Division, Kapit, Ulu Balleh, Ng.
Mengiong (holotype KEP; isotypes A, BO, K, L, MEL, SAR, SING).
Medium-sized tree to 33 m tall, 80 cm diameter. Leaves of one form mainly, in two distinct
ranks along the branches, elliptic, 5.57 x 2.53.5 cm, chartaceous, drying matt on upper
surface, lower side without conspicuous dot-glands; base cuneate, apex acuminate; midrib
not conspicuously thicker than lateral veins, sunken on upper side; lateral veins arising from
the very base of the leaf-blade, sunken on upper side, elevated on lower side; intercostal
veins forming only a loose network; glabrous; stalk 57 mm long. Inflorescence a raceme
1.55 cm long, axis minutely pale-brown pubescent. Flowers unisexual; calyx pubescent
outside, lobes 4; petals 0.60.7 mm long; stamens in male 0.50.7 mm long; female
unknown. Fruit unknown.
Distribution. Known only from the type collection from Ulu Balleh in the Kapit district in
Sarawak.
Ecology. Lowland mixed dipterocarp forest.
18
3. Anisophyllea corneri Ding Hou

(E.J.H. Corner, 20th Century eminent tropical botanist)
l.c. 478; Corner l.c. 132; Ashton l.c. 347; Kochummen l.c. 305; Whitmore, Tantra & Sutisna l.c. 295.
Type: Corner SFN 25927, Malay Peninsula, Trengganu (holotype SING; isotypes BO, KEP).
Small to medium-sized tree to 30 m tall, 55 cm diameter. Bark smooth; inner bark dark
brown. Sapwood reddish brown. Leaves of one form mainly, in two distinct ranks along the
branches, elliptic or oblong, 7.513.5 x 2.55.5 cm, thinly coriaceous, drying matt on upper
surface, lower side without conspicuous dot-glands; base cuneate, apex acute; midrib
conspicuously thicker than lateral veins, raised on the upper side; lateral veins arising from
the very base of the leaf-blade, elevated on both sides; intercostal veins forming a dense and
tessellated network of raised fine veins; glabrous; stalk 510 mm long. Inflorescence a
raceme-like panicle with reduced side-branches, 49 cm long, axes minutely pale-brown
pubescent. Flowers unisexual; calyx pubescent outside, lobes 45; petals 0.10.2 mm long;
stamens in male 0.30.5 mm long, replaced by staminodes in the female; pistil in female 0.5
mm long, with 4 styles, replaced by pistil rudiments in the male. Fruit broadly ellipsoid,
613.5 x 46 cm, smooth on the surface when dry.
Vernacular names. Sarawakmertama (Iban), sireh sireh (Malay).
Distribution. Peninsular Malaysia, Borneo. In Sabah and Sarawak in all districts. Also in
Ecology. Locally common in mixed dipterocarp forest in the lowlands on leached yellow
humult soils, and to submontane forest at 1200 m.
4. Anisophyllea disticha (Jack) Baill.

(Latin, distichus = 2-ranked; the leaf arrangement on branches)
l.c. (1875) 311; Merrill l.c. 422; Masamune l.c. 515; Ding Hou l.c. 479 (excl. A. rhomboidea in syn.);
Corner l.c. 132; Ashton l.c. 347 (excl. A. rhomboidea in syn.); Kochummen l.c. 307; Whitmore, Tantra
& Sutisna l.c. 295. Basionym: Haloragis disticha Jack, Mal. Misc. 2 (1822) 19. Type: Jack, s.n.,
Sumatra (lectotype L, here chosen; isolectotype FI). Synonyms: Anisophyllum trapezoidale Baill.,
Adansonia 3 (1862) 24, 36.
Shrub or small tree to 7 m tall, 5 cm diameter. Bark smooth; inner bark dark brown.
Sapwood pale brown. Leaves of two forms, in four distinct ranks along the branches, those
of the lower 2 ranks oblong-rhomboid, those of the upper 2 ranks tiny and subulate to
falcate, the larger leaf form 1.53.5 x 0.51.5 cm, the smaller leaf form 0.50.7 x 0.10.3
cm, chartaceous to thinly coriaceous, drying matt on upper surface, lower side without
conspicuous dot-glands; base cuneate, apex acute; midrib slightly thicker than lateral veins,
flat on the upper side; lateral veins one pair arising from the very base of the leaf-blade and
a third (unpaired) lateral vein arising from the midrib away from the base of the leaf-blade
19
20
on the acroscopic side of the midrib, flat on upper side, elevated on lower side; intercostal
veins scalariform or forming a dense and tessellated network of flat to slightly raised pale
fine veins; glabrous; stalk none or very short, 01 mm long. Inflorescence a raceme,
0.52.7 cm long, in the axils of the large leaves, axes pale-brown fine-hairy. Flowers
unisexual; calyx pubescent outside, lobes (3)4(5); petals 1.21.3 mm long; stamens in
male 0.30.5 mm long, replaced by staminodes in the female; pistil in female 0.50.8 mm
long, with 1 style, replaced by pistil rudiments in the male. Fruit usually one per raceme,
ellipsoid, 1.52.5 x 0.51 cm, ripening bright red, smooth on the surface when dry.
Vernacular names. Sarawakambun ambun (Kedayan), kayu runap (Punan Tutoh),
mertama (Iban).
Distribution. Sumatra, Peninsular Malaysia, Borneo. In Sabah and Sarawak in all districts.
Also in Brunei and Kalimantan.
Ecology. Understorey of mixed dipterocarp forest and old secondary forest, lowlands to
ridges to 1000 m, recorded on alluvial, sandstone, granite and ultramafic soils.
5. Anisophyllea ferruginea Ding Hou

(Latin, ferrugineus = red-brown; the hairs on the leaves)
l.c. 477; Ashton l.c. 349; Whitmore, Tantra & Sutisna l.c. 295. Type: bb. 25118, E Borneo (holotype
BO; isotype L).
Medium-sized tree to 30 m tall, 40 cm diameter. Bark smooth to cracking or slightly

fissured; inner bark dark brown. Sapwood pale brown. Leaves of one form mainly, in two
distinct ranks along the branches, ovate to elliptic, 1216 x 46 cm, coriaceous, drying matt
on upper surface, lower side without conspicuous dot-glands; base cuneate to rounded, apex
acute to shortly cuspidate; midrib slightly thicker than lateral veins, raised on the upper side;
lateral veins (one or both pairs) arising from the midrib away from the base of the leafblade, elevated on both sides; intercostal veins forming a dense and tessellated network of
raised fine veins; glabrous on upper side, rusty hairy on lower side (the remains of the hairs
always visible even on old leaves under 10X magnification); stalk 1017 mm long.
Inflorescence a panicle, 49 cm long, axes minutely pale brown pubescent. Flowers
unisexual; calyx pubescent outside, lobes (3)4(5); petals 0.10.2 mm long; stamens in
male c. 0.3 mm long, replaced by staminodes in the female; pistil in female 0.5 mm long,
with 4 styles, replaced by pistil rudiments in the male. Fruit ellipsoid, to 8 x 5.5 cm,
smooth on the surface when dry.
Vernacular names. Sarawakbelian landak, dajak, kepajang, kepajang landak, sangkuak,
tengoda (Malay).
Distribution. W and SE Borneo. In Sarawak recorded from the Semengoh (Kuching) and
Balingian, Bintulu (Nyabau FR) and Miri (Lambir NP) areas. Not recorded for Sabah. Also
in Brunei and Kalimantan.
21
Ecology. A characteristic species of deep sandy humult ultisol soils in mixed dipterocarp
forest to 600 m; uncommon but locally frequent.
6. Anisophyllea globosa Madani
Fig. 1.
(Latin, globosus = round; the fruits)

l.c. 53. Type: Shea & Minjulu SAN 76131, Sabah, Kudat, Bengkoka Peninsula (holotype SAN;
isotypes K, L).
Small tree to 10 m tall, 10 cm diameter. Bark smooth; inner bark red-brown. Sapwood
white. Leaves of one form mainly, in two distinct ranks along the branches, narrowly
elliptic, 57 x 1.52.5 cm, chartaceous, drying matt on upper surface, lower side with
scattered tiny but conspicuous dot-glands; base cuneate, apex acute to acuminate; midrib
similar to lateral veins in thickness, flat or slightly sunken on the upper side; lateral veins
(one or both pairs) arising from the midrib away from the base of the leaf-blade, flat on the
upper side, elevated on the lower side; intercostal veins forming a loose network of fine
veins; glabrous; stalk 47 mm long. Inflorescence and flower not known. Fruit globose,
55.5 cm across, irregularly rough-lumpy on the surface when dry.
Distribution. Probably endemic to Sabah, known only from the type collection from
Bengkoka in Kudat district.
Ecology. Open secondary vegetation at edge of steep gully, lowlands at 50 m.
7. Anisophyllea impressinervia Madani

(Latin, impressus- = sunken, nervus = veins; the impressed leaf veins)
l.c. 53. Type: Shea & Minjulu SAN 76094, Sabah, Kudat, Bengkoka Peninsula (holotype SAN;
isotypes A, K, L, SAR, SING).
Medium-sized tree to 26 m tall, 30 cm diameter. Bark irregularly rough-cracking, finely

flaky at the base; inner bark red grading to yellow-orange. Sapwood pale yellow. Leaves of
one form mainly, in two distinct ranks along the branches, elliptic or ovate, 44.5 x 23 cm,
chartaceous to thinly coriaceous, drying matt on upper surface, lower side with scattered
tiny but conspicuous dot-glands; base cuneate to rounded, apex acute; midrib similar to
lateral veins in thickness, sunken on the upper side; lateral veins (one or both pairs) arising
from the midrib away from the base of the leaf-blade, sunken on the upper side, elevated on
the lower side; intercostal veins forming a loose network of fine veins; glabrous; stalk 36
mm long. Inflorescence and flower not known. Fruit globose, c. 4.55 cm across,
irregularly rough-lumpy on the surface when dry.
Distribution. Possibly endemic to Sabah, known only from the type collection from Bengkoka
in Kudat district.
Ecology. Secondary forest, lowlands at c. 30 m.
22
8. Anisophyllea nitida Madani

(Latin, nitidus = polished; the shiny dry upper leaf surface)
l.c. 53. Type: Ampuria SAN 40249, Sabah, Kuala Penyu (holotype SAN).
Medium-sized tree to 23 m tall, 25 cm diameter. Bark smooth to scaly. Sapwood brownish.

Leaves of one form mainly, in two distinct ranks along the branches, broadly elliptic-ovate
to almost round, 5.510 x 45.6 cm, thickly coriaceous, drying glossy on upper surface,
lower side without conspicuous dot-glands; base obtuse-rounded, apex acute to slightly
cuspidate; midrib slightly to conspicuously thicker than lateral veins, raised on the upper
side; lateral veins arising from the very base of the leaf-blade, raised on both sides;
intercostal veins forming a loose network of fine veins; glabrous; stalk 1015 mm long.
Inflorescence and flower not known. Infructescence axis covered in persistent velvety
purple-brown hairs. Fruit broadly ellipsoid, to 5 x 2.5 cm, smooth on the surface when dry.
Vernacular name. Sabahengkop engkop (Kedayan Bundu).
Distribution. Possibly endemic to northwest Borneo and highly localised. Known only
from Kuala Penyu in SW Sabah and the Niah area in NE Sarawak.
Ecology. Lowland secondary and disturbed mixed dipterocarp forest on sandy soils.
9. Anisophyllea rhomboidea Baill.

(Latin, rhomboideus = rhombic; the leaf shape)
l.c. (1875) 310; Merrill l.c. 422; Masamune l.c. 515. Type: Beccari PB 1514, Sarawak (lectotype here
chosen: G). Synonyms: A. disticha sensu Ding Hou l.c. 479, pro parte, sensu Ashton l.c. 347, pro
parte, non (Jack) Baill.
Shrub or small tree to 4 m tall. Leaves of two forms, in four distinct ranks along the
branches, those of the lower 2 ranks oblong-rhomboid, those of the upper 2 ranks much
smaller and falcate-elliptic to falcate-ovate, the larger leaf form 3.510 x 1.23.5 cm, the
smaller leaf form 0.61.4 x 0.30.8 cm, chartaceous to thinly coriaceous, drying matt on
upper surface, lower side without conspicuous dot-glands; base cuneate to right-angled,
apex acute; midrib slightly or not thicker than lateral veins, flat on the upper side; lateral
veins one pair arising from the very base of the leaf-blade and a third (unpaired) lateral vein
arising from the midrib away from the base of the leaf-blade on the acroscopic side of the
midrib, flat on upper side, elevated on lower side; intercostal veins scalariform or forming a
loosely tessellated network of flat to slightly raised pale fine-veins or obscure; glabrous to
slightly hairy on the main veins on the lower side; stalk none or very short, 03 mm long.
Inflorescence a raceme or panicle with short side-branches, 1.58(10) cm long, in the axils
of the large leaves, axes rusty to pale brown fine-hairy. Flowers unisexual; calyx pubescent
outside, lobes 4; petals 1.52 mm long; stamens in male 11.5 mm long, replaced by
staminodes in the female; pistil in female 11.5 mm long, with 1 style, replaced by pistil
rudiments in the male. Fruit usually several per raceme, ellipsoid, 1.52.5 x 0.51 cm,
smooth on the surface when dry, ripening pinkish.
23
24
Distribution. Endemic to W Borneo, in Sarawak and Kalimantan. In Sarawak, apparently

restricted to the Datu Permatang, Gunong Pueh, Gunong Selang and Lundu areas, all in the
1st Division.
Ecology. Mixed dipterocarp forest on shallow brown clay soils, lowlands to c. 650 m.
Excluded species
Anisophyllea glandulifolia Madani, Sandakania 3 (1993) 51, previously known from poor
flowering material, is now shown to be Driessenia microthrix Stapf (Melastomataceae),
through the availability of further flowering and fruiting material. Its extreme anisophylly
and supernumery axillary buds are remarkably like that found in Anisophyllea species.
2. COMBRETOCARPUS Hook. f.
(Greek, karpos = fruit; fruit like that of Combretum)
Bentham & Hooker f., Gen. Pl. 1 (1865) 683; Masamune l.c. 516; Browne, FTSB (1955) 303; Ding
Hou l.c. 480; Burgess l.c. 432; Anderson l.c. 291; Ashton l.c. 360; Kochummen l.c. 317; Whitmore,
Tantra & Sutisna l.c. 297.
Trees, very rarely with small stilt-roots or pneumatophore (breathing) roots. Bark
invaginating finely and regularly into wood. Leaves alternate, 2-ranked on the branches,
pinnately veined. Inflorescence paniculate, bisexual. Flower bisexual, radially symmetrical,
34-merous; petals free, linear, entire or sometimes laciniate; stamens free, inserted
between the lobes of the disc; ovary inferior, 3-sided, 3(4)-locular, ovules 2 per carpel,
styles 34 and free; disc annular, lobed. Fruit a dry, 3(4)-winged structure. Seed solitary,
narrow, spindle-shaped.
Distribution. Monotypic. Sumatra, Riouw, Banka, Billiton, Peninsular Malaysia and Borneo.
Combretocarpus rotundatus (Miq.) Danser
Fig. 2.
(Latin, rotundatus = rounded; the leaf shape)

Bull. Jard. Bot. Btzg. 3, 10 (1929) 345; other references as for genus. Basionym: Macrosolen
rotundatus Miq., Sumatra (1861) 138, 346. Type: Teijsmann, s.n., Sumatra (L, U). Synonym: C.
motleyi Hook. f. in Bentham & Hooker f. l.c. 683, Masamune l.c. 516.
Medium-sized or large tree to 40 m tall, 1 m diameter; trunk base sometimes with small stiltroots or (in swampy or water-logged sites) with mats of rust-brown air roots or (in drier
sites) with short, slender, upright, rootlike pneumatophores. Bark grey-brown to brown,
deeply fissured; inner bark yellowish to reddish brown. Sapwood pale yellow. Leaves
broadly elliptic to ovate or nearly round, 814 x 5.59.5 cm, coriaceous; base cuneate to
obtuse, apex rounded; midrib flat to slightly raised on upper side, raised on lower side;
lateral veins 610 pairs, flat on upper side, raised on lower side; intercostal veins forming a
loose network of veins, often obscure; glabrous; stalk 714 mm long. Inflorescence of one
or more panicles to 9 cm long arising from the leaf-axil, the base becoming tuberculate with
25
age. Flowers yellow; calyx-cup conical with 3 sides, 2.54 x 2 mm, glabrous, lobes ovate,
2.53 x 1.52 mm, outside glabrous, inside scantily pale long-hairy; petals c. 2 mm long,
linear, entire or laciniate (the divisions also linear); stamens with filaments 22.5 mm long,
anthers ovoid, c. 0.3 mm long; styles 11.5 mm long. Fruit 3(4)-winged, 23 cm long,
1.52 cm wide; wings semicircular, membranous. Seed solitary in each fruit, elongate to
spindle-shaped.
Vernacular names. Sabahperepat paya (Brunei Malay), perepat perepat (Bajau).
Sarawakkayu tom (Kayan, Kenyah), keruntum (Iban), mutun (Pusa Malay), perepat paya
(Kayan, Kenyah), sabutun (Melanau Oya). Bruneibalak bekatan (Iban), keruntum
(Dusun, Iban), perepat hutan (Brunei Malay, Belait, Kedayan).
Distribution. As for genus. In Sabah and Sarawak found in all types of peat swamp forest.
Ecology. It is often found in alan (Shorea albida) swamp forest in Sarawak and Brunei and
mixed peat swamp forest in general. In the more open conditions on the domed centres of
the largest swamps, and on kerangas (heath forest) sites on groundwater podsols it occurs as
small trees, whereas in the wetter swampy sites truly big trees occur. In the periodically
drier sites in the centre of the big peat swamp complexes, gregarious stands of small
Combretocarpus trees have been called padang keruntum. Keruntum trees coppice easily.
Medium-sized or large trees of it are often stag-headed (with sparse, large-limbed crown
portions) and hollow (Browne l.c.).
Uses. The timber is suitable for railway sleepers and heavy construction if impregnated with
preservative. When well-seasoned, it can be used for flooring and panelling. The wood is
moderately hard and moderately heavy (keruntum logs sink in water). The texture is coarse
and uneven, due to broad rays which produce an oak-like figure (Burgess l.c.; Wong (1982)
DMT).
26
ARAUCARIACEAE
P.C. Yii
Sarawak Forestry Department,
Kuching, Malaysia
Hooker f., Fl. Br. Ind. 5 (1890) 650; Merrill, EB (1921) 32; Ridley, FMP 5 (1925) 277; Masamune,
EPB (1940) 2; Backer & Bakhuizen f., FJ 1 (1963) 87; Keng, OFMSP (1969) 12, TFM 1 (1972) 39; de
Laubenfels, Philip. J. Biol. 7, 2 (1978) 143, FM 1, 10 (1988) 419; Kochummen, Mal. For. Rec. 17
(1979) 54; Cockburn, TS 2 (1980) 2; Anderson, CLTS (1980) 347; Corner, WSTM 2 (1988) 762;
Whitmore, Tantra & Sutisna, CLK 1 (1989) 27.
Monoecious trees or rarely shrubs. Exudate resinous, colourless or slightly whitish. Leaves
simple, spiral, opposite, sub-opposite or in whorls, scale- or needle-like or broad leathery
blades with many faint closed longitudinal veins. Male and female cones on separate branches.
Male cones solitary, lateral or rarely terminal, cylindrical, consisting of numerous spirally
arranged microsporophylls, each bearing a few to numerous microsporangia at the abaxial or
lower side of an enlarged shield-like apex. Female cones solitary, terminal on robust shoots or
pedunculate with bracts at the base, ovoid or round, consisting of an axis with numerous
spirally arranged seed-scales (megasporophylls), each bearing a solitary seed (ovule) on the
adaxial (upper) surface; mature female cones large, woody; cone-scales wedge-shaped. Seed
winged or not; cotyledons either in 2 fused pairs or of 4 independent units.
Distribution. 2 genera (Agathis and Araucaria) and 40 species, chiefly in countries bordering
the South Pacific Ocean (Indo-China, Malesia, New Caledonia, Australia, New Zealand,
Melanesia, and South America). In Sabah and Sarawak represented by a single genus (Agathis)
with 5 species.
AGATHIS Salisb.
(Greek, agathis = a clew; the shape of the cone)
Trans Linn. Soc. London 8 (1807) 311; Hooker f. l.c. 650; Merrill l.c. 32; Ridley l.c. 277; Meijer
Drees, Bull. Jard. Bot. Btzg. 3, 16 (1940) 455; Backer & Bakhuizen f. l.c. 87; Keng l.c. (1969) 12, l.c.
(1972) 39; Cockburn l.c. 2; Anderson l.c. 347; Whitmore, Pl. Syst. Evol. 41 (1980) 46, New Phytol. 84
(1980) 407; de Laubenfels, Blumea 24 (1978) 499, Blumea 25 (1979) 531, l.c. (1988) 419; Whitmore,
Tantra & Sutisna l.c. 27. Synonym: Dammara Link, Enum. Hort. Berol. Alt. 2 (1822) 411.
Usually large trees. Bole straight and cylindrical, often swollen at the base or with large
superficial roots. Bark light-grey to greyish brown, usually smooth and lenticellate when
young, gradually peeling off into thin irregular flakes, leaving behind irregular pock-marks
on larger trees; inner bark granular, reddish brown. Twigs glabrous, greenish. Terminal
buds globular, usually covered with several pairs of overlapping scales. Petioles usually
very short, hardly distinguishable from the blades, slightly channelled above. Leaves entire,
27
spirally arranged on the older branches and decussate on the young twigs, size and shape
extremely variable, lanceolate, ovate, to elliptic, without distinct midrib; juvenile leaves
distinctly larger than the adult leaves and often have more acuminate apices. Male cones
subtended by several pairs of bracts, sessile or shortly pedunculate. Female cones with
numerous closely appressed more or less triangular seed-scales. Seed flattened ovoid, with 2
unequal lateral wings; cotyledon 2, opposite and leaf-like.
Distribution. About 21 species; Indo-China, Malesia, Australia, New Zealand and Melanesia
(except Solomon Islands). 5 species occur in Sabah and Sarawak.
Ecology. Large and emergent trees with 2 species, Agathis borneensis and A. endertii,
widely distributed from lowland peat swamp, kerangas (heath), mixed dipterocarp and
montane forest to about 2400 m. Young trees often produce female cones several years
ahead of males.
Uses. Agathis produces a beautiful softwood which fetches a very good price. Its pleasing
yellow timber has a natural sheen which is very suitable for furniture and cabinet making,
wall and ceiling panelling. It is not durable for outdoor use. The white crystalline resin is
also collected for sale as copal. For more information, see Whitmore, Econ. Bot. 34 (1980) 1.
Taxonomy. The species of Agathis occurring in Sabah and Sarawak are characterised by the
decussate leaves with close faint longitudinal veins without prominent midrib, and by the
male and female cones which are borne on separate branches. For a detailed account of the
systematy of the genus see papers by de Laubenfels, Blumea 24 (1979) 531 and Whitmore,
New Phytol. 84 (1980) 407 and Pl. Syst. Evol. 41 (1980) 41.
Key to Agathis species

1.
Leaf lower surface glaucous.........................................................................................2

Leaf lower surface not glaucous...................................................................................4
2.
Adult leaves distinctly convex-lens-shaped, asymmetric; apex and base sharply acute
...............................................................................................................4. A. lenticula
Adult leaves not so................................................................................................................3
3.
Adult leaves ovate or orbicular, apex obtuse or rounded, usually less than 4 cm. Male
cone usually less than 6 mm in diameter, shortly pedunculate. Seed-scales without
distinct lip...............................................................................................5. A. orbicula
Adult leaves elliptic, apex acute or rarely blunt, usually longer than 5 cm. Male cones
about 10 mm in diameter, sessile. Seed-scales with distinct protruding lips at the apical
margin.....................................................................................................2. A. endertii
4.
Adult leaves elliptic, usually longer than 5 cm, apex acute. Male cones oblong,
pedunculate, usually larger than 20 mm in diameter.................................................1. A.
borneensis
Adult leaves ovate, usually less than 5 cm long, apex usually acuminate or round and
blunt with a distinct tip. Male cones about 10 mm in diameter........3. A. kinabaluensis
28
ARAUCARIACEAE (YII)
1. Agathis borneensis Warb.
Fig. 1.
(of Borneo)
Monsunia 1 (1900) 184; Merrill l.c. 32; Masamune l.c. 2; Meijer Drees l.c. 459; de Laubenfels l.c.
(1979) 532, l.c. (1988) 433; Whitmore l.c. (1980) 54. Type: Beccari 491 (B) & 596 (B, K), Sarawak
(syntypes). Synonyms: Pinus dammara Lamb., Descr. Pinus 1 (1803) 61; A. dammara (Lamb.)
Richard, Comm. Bot. Conif. & Cycad. (1826) 83; A. loranthiifolia Salisb. l.c. 312; Dammara loranthiifolia
(Salisb.) Link l.c. 411; A. beccarii Warb. l.c. 184; A. macrostachys Warb. l.c. 183; A. rhomboidalis
Warb. l.c. 184; A. alba Foxw., Philip. J. Sc. 4 (1909) 442; A. latifolia Meijer Drees l.c. 459.
Emergent tree to 55 m tall and 100 cm diameter. Bark warty lenticellate, papery scaly or
flaky, maurish grey; inner bark granular, pale brown. Leaves when juvenile ovate to lanceolate,
up to 14 x 4 cm; when adult elliptic to ovate, 612 x 23.5 cm, leathery, glabrous; base
cuneate rounded, apex acute. Male cones oblong, 47 cm long, 2025 mm across, on short
peduncles, 210 mm long; microsporophylls 67 x 4.55 mm, with broad, rounded apex.
Female cones ovoid, 68.5 x 5.56.5 cm; seed-scales 3040 x 2528 mm, roughly triangular
in shape with well rounded apical margin. Seeds flattened ovoid, c. 12 x 9 mm, with one
well-developed wing, c. 20 x 16 mm.
Vernacular names. Sabahmanggilan (Dusun), salang (Kedayan). Sarawakbindang
(Malay), bulu (Iban), kayu jadi (Malay, Iban). Bruneitulong (Brunei, Malay, Dusun).
Distribution. Sumatra, Peninsular Malaysia and throughout Borneo. In Sabah and Sarawak
widespread; also in Brunei and Kalimantan.
Ecology. Infrequent in the lowlands, although found in sandy margins of some peat swamps
and kerangas, and lower montane forests to 1200 m. Generally prefers more acidic soils;
often forming dense, nearly pure stands on certain areas of low-lying wet kerangas forest
and dry kerangas forest on sandstone formations at higher altitude, elsewhere uncommon
and scattered.
2. Agathis endertii Meijer Drees

(F.H. Endert, 18911953, forester at the Forest Research Institute, Bogor, Indonesia)
l.c. 470; Masamune l.c. 3; de Laubenfels l.c. (1979) 534, l.c. (1988) 439. Type: bb. 21694, Southern
Borneo (holotype BO).
Emergent tree to 60 m and 100 cm diameter. Bark smooth with shallow pock-marks and
scales, grey-purplish brown; inner bark granular, straw-brown. Exudate whitish, opaque.
Leaves when juvenile lanceolate, with blunt or acute apex; when adult elliptic, 59 x 1.8
3.5 cm, leathery and slightly glaucous on undersurface; base cuneate rounded, apex broadly
acute or rarely blunt. Male cones cylindrical, 2.63.8 cm long, 710 mm across, sessile;
microsporophylls spoon-shaped, c. 25 x 20 mm, with spreading and slightly angled apical
margin. Female cones shortly ovoid, c. 4.5 x 7 cm; seed-scales c. 32 x 42 mm, more or less
triangular with well rounded apical margin, and a distinct protruding lip (c. 8 x 3 mm) at
the apical margin. Seed flattened ovoid, c. 11 x 8 mm; wing one, c. 18 x 14 mm.
29
Vernacular names. Sarawakbulok (Iban).

Distribution. Endemic to Borneo. In Sabah and Sarawak scattered.
Ecology. Scattered throughout lowland rain forests from sea-level to sandstone kerangas
forest at 1600 m.
3. Agathis kinabaluensis de Laub.

(of Mt. Kinabalu)
l.c. (1979) 535, l.c. (1988) 439. Type: de Laubenfels P625, Sabah, Mt. Kinabalu (holotype L; isotypes
A, K, SAN).
Tree up to 36 m tall, becoming relatively stunted on exposed ridges and summits. Bark dark
brown, with numerous lenticels, peeling off into irregular flakes; inner bark granular,
reddish brown. Exudate white. Leaves when juvenile ovate, up to 9 x 4.4 cm, apex strongly
acuminate; when adult ovate, 3.57 x 1.83.2 cm; base cuneate rounded, apex slightly
acuminate or more or less round and blunt with distinct tips on smaller leaves. Male cones
cylindrical, 1.83 cm long, 810 mm across, nearly sessile or on very short peduncles;
microsporophylls spoon-shaped, c. 1.7 x 1.4 mm, with very slightly angled apical margin.
Female cones ovoid, c. 8 x 11 cm; seed-scales 2832 x 4045 mm, with distinct narrow
ridges along apical margin and broadly rounded at the upper end. Seed c. 11 x 7 mm,
distinctly acute at one end and with a broad wing c. 20 x 12 mm at the other.
Vernacular name. Sarawaktumu (Kelabit, Murut).
Distribution. Only known from Mt. Kinabalu, Sabah, and Mt. Murut in Sarawak.
Ecology. Upper mossy montane forest at 15002400 m.
4. Agathis lenticula de Laub.

(Latin, lenticularis = a double convex lens; the leaf shape)
l.c. (1979) 537, l.c. (1988) 436. Type: de Laubenfels P619, Sabah, Mt. Kinabalu, Park HQ (holotype
L; isotypes A, K, SAN).
Tree to 45 m tall. Bark greyish brown, with numerous small lenticels; young bark distinctly
covered with thin exfoliating flakes, gradually breaking into irregular plates; inner bark
granular, reddish brown. Exudate whitish, gradually turning yellow. Leaves when juvenile
lenticular, up to 11 x 4.7 cm, tapering toward both ends; when adult lenti-cular, 57 x 1.7
2.4 cm, leathery, glaucous on the undersurface, apex and base more or less acute. Male
cones cylindrical, 34 cm long, 910 mm in diameter, on short peduncles,
30
ARAUCARIACEAE (YII)
31
26 mm long; microsporophylls spoon-shaped, 22.5 x 1.52 mm, with blunt spreading

apical margin. Female cones spherical, c. 7 x 6 cm; seed-scales c. 40 x 27 mm. Seeds c. 11
x 7 mm, with a broadly rounded wing, c. 8 x 14 mm at one end.
Vernacular names. Sabahtangilan, tungilan (Dusun).
Distribution. So far only known from the Crocker Range, including Mt. Kinabalu in Sabah.
Ecology. Emergent trees in mossy montane rain forest, at 11401680 m.
5. Agathis orbicula de Laub.

(Latin, orbicularis = circular; the leaf shape)
l.c. (1979) 540, l.c. (1988) 437. Type: de Laubenfels P614, Sarawak, Lawas (holotype L; isotypes A,
K, SAN, SAR).
Tree to 40 m tall. Bark dark brown, peeling off into irregular flakes, exposing scattered
lenticels; inner bark granular, reddish brown. Exudate resinous, light yellow and produced
in abundance. Leaves when juvenile ovate, up to 6.5 x 2.8 cm; apex bluntly acute; when
adult ovate to orbicular, 2.44 x 1.22.4 cm, broadly rounded to slightly angled at the apex,
tapering toward the base. Male cones minute, cylindrical, 0.81.4 cm long, 46 mm
through; microsporophylls helmet-shaped, 1.21.5 x 11.2 mm, with blunt apex. Female
cones ovoid, c. 7 x 4.5 cm; seed-scales ovate, c. 20 x 33 mm. Seeds similar to that of A.
lenticula.
Vernacular names. Sarawakbulok (Iban), tubu (Kenyah & Kayan), tumuh (Murut).
Distribution. Endemic to Borneo; known from southern parts of Sabah to Central Sarawak.
Ecology. Scattered in sandstone kerangas forest at 4501050 m.
32
BIGNONIACEAE
A. Berhaman
Sandakan, Malaysia
Merrill, EB (1921) 525; van Steenis, Rec. Trav. Bot. Neerl. 24 (1927) 787, Bull. Jard. Bot. Btzg. 3, 10
(1928) 173, FM 1, 8 (1977) 114; Masamune, EPB (1942) 652; Kochummen, TFM 3 (1978) 36;
Anderson, CLTS (1980) 152; Cockburn, TS 2 (1980) 19; Santisuk, Flora of Thailand 5 (1987) 32;
Corner, WSTM 1 (1988) 172; Whitmore, Tantra & Sutisna, CLK 1 (1989) 28.
Trees, shrubs or woody climbers; twigs often lenticellate; stipules absent. Leaves generally
opposite and 124-times pinnate, or simple and whorled, often with glands on the lower
surface of the leaflets; domatia sometimes present. Inflorescences generally terminal,
sometimes borne on the trunks and branches. Flowers bisexual, usually very showy; calyx
tubular, subtruncate or with 5 lobes or teeth, closed in bud and later splitting into lobes,
often glandular outside; corolla in most genera bilaterally symmetrical, in some genera
somewhat radially symmetrical, basally tubular, upper part expanded, generally campanulate
or salver-shaped, with 5 lobes, 2 on the upper side, 3 on the lower; stamens 5, almost equal,
or (mostly) 4 and didynamous (2 long and 2 short), attached to the corolla (epipetalous); ovary
superior, 2-celled, style long, stigma 2-lobed, nectary usually present and ring-like; ovules
several to many per cell, arranged longitudinally in 12 rows on the septum. Fruit a pod
divided by a longitudinal partition into 2 compartments, splitting open when ripe. Seeds
many, flat; embryo near the centre; peripheral wing very thin and transparent or thickened
and opaque; germination epigeal.
Distribution. About 120 genera and 650 species in the tropics and subtropics. In Sabah and
Sarawak, 10 species in 5 genera including 5 apparently unnamed species (4 woody climbers
and 1 tree). A number of exotic species have been introduced as ornamental plants:
Arrabidaea magnifica, Jacaranda rhombifolia (jambul merah, formerly J. filicifolia), Mansoa
hymensala, Millingtonia hortensis, Spathodea campanulata (African tulip tree), Tabebuia rosea,
and Tecoma stans (Yellow bells).
Ecology. Mostly evergreen but Oroxylum indicum and Dolichandrone spathacea have been
noted as leafless for many months during the dry season of seasonal parts of Asia.
Dolichandrone spathacea is confined to the mangroves, Deplanchea bancana is found in
kerangas (heath) and peat swamp forest, some woody climber species appear restricted to
limestones, and some Radermachera species may be restricted to ultramafic soils. In many
species, the flowers open at night, and some have fragrant white flowers with long tubes
which are pollinated by moths. O. indicum has flowers with lurid colour, musky odour,
fleshy funnel-shaped corollas, which are pollinated by nectarivorous bats. The winged seeds
are dispersed by wind.
33
34
BIGNONIACEAE (BERHAMAN)
Uses. The species generally have soft and light wood, at present without commercial value.
A few traditional uses of the wood or leaves have been recorded, although in Sabah and
Sarawak there appears to be no documented uses. Javanese fishermen in Indonesia use
Dolichandrone spathacea wood for floating their nets. In the Philippines, the wood of D.
spathacea has been used for making matches and wooden shoes (clogs). In Peninsular
Malaysia, a decoction of Oroxylum indicum leaves may be drunk for stomach ache, by the
Malays, causing eructations and so bringing relief.
Taxonomy. The family is closely related to the Scrophulariaceae. In biochemical

characteristics, it shares a number of similarities with the Labiatae, Scrophulariaceae, and
Verbenaceae. For a detailed discussion, see van Steenis l.c. (1977).
Key to genera
1.
Woody climbers.............................................................................................................
Nyctocalos Teijsm. & Binn.
in Miquel, J. Bot. Neerl. 1 (1862) 366; van Steenis l.c. (1927) 805, l.c. (1928) 178, l.c.
(1977) 123; Santisuk l.c. 34.
8 species in Assam, Burma, Yunnan, Thailand, and Malesia; 4 unnamed species in

Sabah and Sarawak.
Woody climbers without tendrils. Leaves pinnately 13- or 35-foliolate. Flowers
several in a short terminal raceme; calyx cup-shaped, subtruncate or with 5 hornlike teeth; corolla campanulate or salver-shaped, with 5 imbricate lobes; stamens 4,
didynamous. Fruit large, flat. Seeds hyaline-winged, rounded.
Trees...........................................................................................................................2
2.
Leaves simple, in whorls. Inflorescence a terminal thyrse with a somewhat flat-topped

structure, branches more than 20 and held horizontally at almost right angles to the
main inflorescence axis. Flowers yellow.................................................1. Deplanchea
Leaves compound, opposite. Inflorescence a raceme or a thyrse with few (less than 20)
ascending branches, never with a flat-topped structure. Flowers white to reddish or
purplish, or if yellow then borne on racemes on the bare parts of branches...................3
3.
Leaves once pinnate (trees of mangrove forest)..................................2. Dolichandrone

Leaves 14-times pinnate (trees of inland forests)........................................................4
4.
Leaves 12(3)-times pinnate, leaflets with more than 5 pairs of lateral veins, lower
surface always with scattered glands or glands grouped at the base. Corolla up to 5 cm
long, white to pale lilac or yellow. Pods to 2050 cm long, twisted or straight..............
.................................................................................................4. Radermachera
Leaves 24-times pinnate, leaflets with 45 pairs of lateral veins, lower surface with
glands only in young and some mature leaves, and then only at the basal parts of
leaflets. Corolla about 10 cm long, reddish purple. Pods 60120 cm long, never twisted.............................................................................................................3. Oroxylum
35
1. DEPLANCHEA Vieill.
(E.F. Deplanche 18241874, French physician & naturalist)
Bull. Soc. Linn. Normand. 7 (1862) 96; van Steenis l.c. (1927) 906, l.c. (1977) 135; Kochummen l.c.
37; Anderson l.c. 152; Cockburn l.c. 19; Corner l.c. 175; Whitmore, Tantra & Sutisna l.c. 28.
Small to medium-sized trees. Bark often lenticellate. Sapwood soft. Leaves simple, in whorls,
quite large, elliptic to obovate, chartaceous to coriaceous; base with two large saucershaped glands on upper surface. Inflorescence terminal, a thyrse with a somewhat flattopped structure, the branches many (more than 20) and held horizontally at almost right
angles to the main inflorescence axis, large and conspicuous. Flowers large; calyx 5-lobed;
corolla irregular (zygomorphic), funnel-shaped, tube with a short constricted basal part and
a flared upper part, lobes 5, yellow; stamens strongly exserted, 4 (rarely 5), didynamous;
disc annular to lobed; ovary 2-celled; style long exserted; stigma 2-lobed. Fruit a 2-valved
capsule. Seeds many, thinly winged.
Distribution. 5 species, Malesia to N Australia. 1 species in Borneo (Sabah, Sarawak, Brunei,
and Kalimantan).
Ecology. Being light-demanding, species of the genus are mainly found in secondary rain
forests (including kerangas forest), woodland savannahs, and grasslands, from sea-level to
1000 m.
Deplanchea bancana (Scheff.) Steenis
Fig. 1.
(of Banka Island)

l.c. (1927) 921, l.c. (1928) 221, l.c. (1977) 137; Kochummen l.c. 37; Anderson l.c. 152; Cockburn l.c.
19; Corner l.c. 175; Whitmore, Tantra & Sutisna l.c. 28. Basionym: Diplanthera bancana Scheff.,
Nat. Tijd. Ned. Ind. 31 (1870) 334. Type: Teijsmann, H.B. 9666, Banka (BO). Synonym: Deplanchea
coriacea Steenis l.c. 224.
Small to large trees 1045 m tall, 2570 cm diameter; bole fluted at base, buttresses small or
large. Bark white to brown, lenticellate and slightly fissured; inner bark yellow to pale
brown. Sapwood white to pale yellow, soft. Young twigs lenticellate, glabrous to pale hairy,
pale brown to yellow. Leaves obovate to elliptic, 934 x 5.520 cm, chartaceous to
coriaceous; base cordate to cuneate, apex rounded, rarely acuminate-acute; midrib glabrous
to yellow-hairy on lower surface; lateral veins 78 pairs, raised and glabrous to yellow-hairy
on the lower surface; stalk 36 cm long, glabrous to yellow-hairy. Inflorescences showy,
stalk 520 cm long. Flowers showy; calyx 1216 mm long, with scattered glands, teeth 5;
corolla c. 3.5 x 2.5 cm, yellow, 5-lobed, the upper 2 lobes recurved; stamens 4, filaments
erect and curved over the upper corolla-lobes, base glandular hairy; ovary subsessile, each
locule with two placentas; style greenish yellow, exserted and curved like the stamens;
stigma bilobed. Fruits oblong, 1014 x 3.5 cm. Seeds flat, c. 3 x 2 cm; wing broad, very
thinly transparent, irregularly lobed.
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah and Sarawak uncommon.
Ecology. In Sabah and Sarawak, it occurs in heath forests on white sands or in peat swamps,
in primary and secondary forests, from sea-level to 1000 m.
36
2. DOLICHANDRONE Seemen
(Greek, dolichos = long, andron = male; the long stamens )
Ann. Mag. Nat. Hist. Ser. 3, 10 (1862); van Steenis l.c. (1927) 928, l.c. (1928) 227, l.c. (1977) 141;
Kochummen l.c. 39; Anderson l.c. 152; Cockburn l.c. 19; Corner l.c. 176; Whitmore, Tantra & Sutisna
l.c. 28.
Medium-sized trees. Leaves once pinnate, leaflets elliptic, entire. Inflorescence a terminal
raceme. Flowers fragrant, nocturnal; calyx spathaceous, somewhat curved; corolla funnelshaped, equally 5-lobed; stamens 4, didynamous, included; disc annular; ovary with many
ovules in many rows; style slender. Fruit a subcylindric capsule. Seeds hyaline-winged.
Distribution. 9 species, from East Africa through tropical Asia to Australia; 4 species in
tropical SE Asia; 1 species in Sabah and Sarawak.
Dolichandrone spathacea (L. f.) K. Schum.
Fig. 2.
(Latin, spathaceus = resembling a spathe; the calyx form)

Fl. Kais. Wilh. Land. (1889) 123; Merrill, l.c (1921) 525; van Steenis l.c. (1927) 937, l.c. (1928) 227,
l.c. (1977) 142; Masamune l.c. 652; Kochummen l.c. 39; Anderson l.c. 152; Cockburn l.c. 19; Santisuk
l.c. 56; Corner l.c. 176; Whitmore, Tantra & Sutisna l.c. 28. Basionym: Bignonia spathacea L. f., Suppl.
(1781) 283. Type: Konig, Herb. Linn. No. 776. 8 (LINN). Synonyms: Dolichandrone rheedii Seemen,
J. Bot. 8 (1870) 380; Dolichandrone longissima K. Schum. in Engl. & Prantl, Pfl. Fam. 4, 3b (1894)
240.
Tree to 25 m tall, 1040 cm diameter. Bark pale brown, smooth to fissured; inner bark
laminated, reddish pink. Sapwood white, soft. Young twigs lenticellate. Leaves once pinnate,
opposite; rachis smooth, brown; leaflets usually 34 pairs with one terminal leaflet, ovateoblong to lanceolate, 5.515 x 2.56 cm, drying dark to brown-black; base oblique, apex
acuminate pointed; lateral veins 58 pairs, lower surface with hairy domatia; stalks short in
lateral leaflets but 24 cm in terminal leaflet. Inflorescences 28-flowered, with one flower
opening at a time. Flowers fragrant; calyx conical, closed and with hooked apex in bud,
with scattered glands, splitting along one side; corolla white, with a tube to 14.5 cm long,
bell-shaped near the mouth, 34 cm wide, with many scattered glands outside, lobes 5 with
frilled margin; stamens 4, didynamous with a fifth rudiment, included; style long, exserted,
persistent in young fruit; anthers glabrous, bilocular. Fruits 2033.5 x 1.53 cm, generally
curved, hanging in clusters from the ends of the branches. Seeds many, dark grey, in many
rows, c. 15 x 9 mm including the thick corky wings.
Vernacular names. Sabahkelaju, tui (Dusun). Sarawaktuih (Iban).
Distribution. From the coasts of Malabar to tropical SE Asia and the whole of Malesia.
Ecology. Restricted to the back mangroves and along banks of tidal rivers and estuaries. In
Sabah, common in the mangrove forests of the west coast; in Sarawak occasional on clay
soils at inland limits of mangroves.
37
Uses. The wood is soft and light with density of about 480 kg/m3, easy to saw and work
with and seasons well although it must be protected against sapstains. So far, there is no
record of its use in Sabah and Sarawak but in various parts of its geographical range, the
wood is used for wooden clogs, net-floats, scabbards and household utensils. Due to its
anatomical and physical characteristics the wood may also be suitable for pattern making
and matches.
3. OROXYLUM Vent.
(Greek, oros = mountain, xylon = wood or tree; the natural habitat of the tree)
Dec. Gen. Nov. (1808) 8; van Steenis, l.c. (1927) 816, l.c. (1928) 181, l.c. (1977) 128; Masamune l.c.
652; Kochummen l.c. 40; Anderson l.c. 152; Cockburn l.c. 19; Corner l.c. 178; Whitmore, Tantra &
Sutisna l.c. 28.
Small to medium-sized trees. Leaves opposite, 24-times pinnate; leaflets with 45 pairs of
lateral veins, glands on the lower surface present only in young and some mature leaves,
and then only at the basal part of leaflets. Inflorescence a terminal raceme, erect, long.
Flowers fragrant, showy; calyx closed in bud, with an apical pore opening as a somewhat
bell-shaped structure; corolla purplish, with 5 subequal lobes; stamens 5; ovules in many
rows in each cell. Fruit a long, flattened woody capsule. Seeds quite large, with thin wings
all around, in many rows.
Distribution. Monotypic genus distributed in India, Sri Lanka, and throughout SE Asia.
Oroxylum indicum (L.) Vent.
Fig. 3.
(of India)
l.c. 8; van Steenis l.c. (1927) 816, l.c. (1928) 181, l.c. (1977) 128; Masamune l.c. 652; Kochummen
l.c. 40; Anderson l.c. 152; Cockburn l.c. 19; Santisuk l.c. 36; Corner l.c. 166; Whitmore, Tantra &
Sutisna l.c. 28. Basionym: Bignonia indica L., Sp. Pl. (1753) 625. Type: Herb. Hermann No. 236;
vol. 5: Icon. 28, 29, 72 (BM). Synonyms: Bignonia pentandra Lour., Fl. Coch. 2 (1970) 379;
Bignonia tripinnata Noronha, Verh. Bat. Gen. 5 (1790) 8; Calosanthes indica Blume, Bijdr. (1826)
760; Hippoxylum indicum Rafin., Tellur. (1838) 78.
Medium-sized tree, partly deciduous, 2040 m tall, 1040 cm diameter. Bark smooth,
whitish; inner bark pale yellowish. Sapwood pale whitish, soft. Leaves 34-times pinnate,
up to 2 m long, tufted at the branch ends; petioles long, rachis swollen at points of
articulation; leaflets ovate to oblong, 69.5 x 2.55 cm, terminal one always bigger; base
cuneate or mostly oblique, apex acute to acuminate; lateral veins 45 pairs; intercostal veins
reticulate; glands scattered on the flower surface. Inflorescence 25100 cm long. Flowers
nocturnal, c. 9 cm long; calyx coriaceous, 2.53.5 cm long, 1.52 cm across, containing
38
39
40
water in bud, campanulate, dirty brown-violet, becoming almost woody in the fruit; corolla
c. 10 cm long, reddish outside, yellowish to pinkish inside, fleshy, funnel-shaped, the 5
unequal lobes with wrinkled margin; stamens all fertile, hairy at the base, included. Fruit
25100 x 510 cm, hanging conspicuously resembling the big flat blade of a slash-knife.
Seeds numerous, 48 x 2.54 cm including the membranous and transparent wing.
Vernacular names. Sabahparang pamol (colloquial Malay). Sarawakbinkuli (Iban),
gimurai, murai (Padawan Bidayuh), parang nyabor (Iban).
Distribution. As for genus.
Ecology. Secondary forest, mostly below 1000 m.
4. RADERMACHERA Zoll. & Moritzi

(J.C.M. Radermache, 17411783, patron of science in the Netherlands Indies)
in Zollinger, Syst. Verz. Pfl. Ind. Arch. 3 (1855) 53; Merrill l.c. (1921) 525; van Steenis l.c. (1927)
953, l.c. (1928) 238, l.c. (1977) 149, Blumea 23 (1976) 121; Masamune l.c. 652; Kochummen l.c. 42;
Anderson l.c. 152; Cockburn l.c. 20; Corner l.c. 180; Whitmore, Tantra & Sutisna l.c. 29.
Trees or shrubs. Twigs lenticellate. Leaves 13-times pinnate, opposite; leaflets often with
gland-fields on the lower surface. Inflorescence a raceme or thyrse, terminal on branches or
borne on the trunk and branches (plant cauliflorous or ramiflorous). Flowers white, pink or
greenish yellow; calyx 25-lobed, often glandular; corolla 5-lobed, slightly two-lipped;
stamens 4 or 5; ovary elongate, glabrous or covered with tiny scales or tubercles. Fruit a
straight or twisted capsule with a corky central placenta. Seeds many, tiny and narrow, the
ends winged.
Distribution. 15 species; tropical Asia, from India and south China to Malesia; 3 species in
Sabah and Sarawak.
Key to Radermachera species

1.
Leaves typically once pinnate, sometimes 12-times pinnate. Inflorescences always

borne on leafy shoots................................................................................1. R. pinnata
Leaves 23-times pinnate. Inflorescences borne directly on main branches or on trunk
and branches (plant cauliflorous or ramiflorous)..........................................................2
2.
Leaves 3-times pinnate, the secondary rachises never winged; leaflet tips caudate
.............................................................................................................2. R. ramiflora
Leaves 23-times pinnate, the secondary rachises with 12-mm-wide wings; leaflet tips
acuminate to cuspidate............................................................3. Radermachera sp. A.
41
1. Radermachera pinnata (Blanco) Seemen
Fig. 4.
(Latin, pinnatus = feathery; the pinnate leaves)

J. Bot. 8 (1870) 147; Merrill, Sp. Blanc. (1918) 350; van Steenis l.c. (1927) 973, l.c. (1928) 248, l.c.
(1976) 129, l.c. (1977) 153; Kochummen l.c. 42; Corner l.c. 181; Whitmore, Tantra & Sutisna l.c. 29.
Basionym: Millingtonia pinnata Blanco, Fl. Filip. (1827) 501. Type: Merrill Sp. Blanc. 834,
Philippines, Luzon, Rizal Province (neotype A; isoneotypes BO, K, L). Synonym: R. whitfordii Merr.,
Philip. J. Sc. 75 (1912) 352.
subsp. acuminata (Steenis) Steenis

l.c (1976) 129, l.c. (1977) 154; Whitmore, Tantra & Sutisna l.c. 29. Basionym: R. lobbii (Teijsm. &
Binn.) Miq. subsp. acuminata Steenis l.c. (1928) 247. Type: Endert E 1309, S Sumatra, Lampong,
Tandjong Karang (holotype BO; isotype L). Synonyms: R. lobbii Teijsm. & Binn., Nat. Tijd. Ned.
Ind. 25 (1863) 413; R. corymbosa Steenis l.c. (1928) 249.
Tree 840 m tall, 30 cm diameter; buttresses to 2 m high. Bark scaly to flaky, light brown
to grey; inner bark pale yellow. Sapwood white. Leaves 12-times pinnate, petioles 810
cm long; leaflets obovate to obovate-elliptic, 79 x 2.54 cm, coriaceous to chartaceous;
base cuneate, apex acute to rounded; lateral veins 79 pairs. Inflorescence a terminal
thyrse. Flowers mildly fragrant; calyx purple, c. 1.5 cm long, c. 11.5 cm across, with a few
scattered glands outside, lobes 3; corolla about 1.5 cm long, c. 11.2 cm across, basal part
narrow tubular and white, upper inflated portion pale lilac, throat and medium inner surface
of the lobes orange, lobes 5 with pale, minute hairs on the margin; stamens 4, didynamous;
ovary glabrous, style glabrous, 1.22 cm long. Fruit a capsule, hanging in clusters at the
end of branches, 1627.5 x 0.51 cm; persistent calyx c. 1.8 cm long. Seeds c. 2.5 x 0.5 cm,
winged.
Vernacular names. Sarawak/Kalimantanbinutan, kujuk langit (Kapuas Dayak).
Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, Borneo, and the Philippines.
In Sabah and Sarawak, generally uncommon but localised populations may be encountered
in ultramafic and limestone areas in the northern, eastern and southern parts of Sabah, and
around the Niah Caves and Kuching areas in Sarawak.
Ecology. Primary and secondary forest, to 800 m, associated with ultramafic soils (Sabah)
or limestones (Sarawak). Flowering in May, JulyMarch; fruiting in MayNovember.
Taxonomy. This species consists of two subspecies, according to van Steenis l.c. (1977)
153, l.c. (1976) 129. The other subspecies, R. pinnata subsp. pinnata, is restricted to
Celebes, the Moluccas and the Philippines. The main difference is that in subsp. pinnata the
intercostal veins on the lower leaflet surface form a dense and prominent network, whereas
in subsp. acuminata the veins are not conspicuous or prominent. Subsp. acuminata is
slightly variable in leaflet shape and texture.
2. Radermachera ramiflora Steenis

(Latin, ramiflorus = flowering on the branches)
J. Bot. 74 (1934) 5, l.c. (1976) 130, l.c. (1977) 152; Masamune l.c. 653; Whitmore, Tantra & Sutisna
l.c. 29. Type: Clemens 28672, British North Borneo, Mount Kinabalu (holotype BO; isotypes K, L).
Medium-sized tree, 3045 m tall, 3050 cm diameter; trunk fluted at base. Bark greyish,
fissured; inner bark whitish, turning yellowish on exposure. Sapwood yellowish white.
Twigs with corky bark. Leaves 3-times pinnate, crowded at the branch-tips; leaflets 4-6
pairs, elliptic-lanceolate, coriaceous, 3.58.5 x 1.53.5 cm; apex caudate; lateral veins 5-7
42
43
pairs and raised below, lower surface with scattered small glands near the base. Inflorescence a
raceme, borne on the trunk and branches (plants cauliflorous or ramiflorous), 1114 cm
long. Flowers yellow, erect on curved pedicels of c. 20 cm long; calyx dull reddish green to
dark brown when dry, in the bud pear-shaped with many scattered glands outside especially
near the base, 1821 x 68 mm, lobes 3; corolla deep ochre-yellow, the limb red, narrowly
salver-shaped with scattered 24 glands outside, slightly curved, the narrow basal part 35
cm long, lobes 5; stamens 4, exserted, didynamous, hairy at the base; ovary ribbed, style to
33.5 cm long. Fruit straight or twisted, 3570 cm long. Seeds 45 x 2.4 mm, the wing 67
mm.
Vernacular name. Sabahtuik-tuik hutan (Malay).
Distribution. Endemic to Sabah, known from Mt. Kinabalu, Ranau, Kota Marudu, and Mt.
Rara.
Ecology. Rainforest, also in disturbed forest on hill sides, to 1500 m, associated with
ultramafic soils. Flowering in January to March and around August; fruiting recorded in
April and December.
van Steenis l.c. (1977) noted that the calyx is eglandular, but the specimen Chew & Corner
RSNB 4290, examined by him, has a few scattered glands outside the calyx.
3. Radermachera sp. A.
van Steenis l.c. (1977) 152 included the only collection of this undescribed species
(Nooteboom & Aban 1603, Sabah, Kinabalu, at Kampong Kiau) in R. ramiflora, a conclusion
we disagree with. The species is poorly known; the young flower buds are not sufficient for
describing it properly, but the winged secondary rachis and cuspidate leaflet tips clearly
distinguish it from R. ramiflora.
44
BURSERACEAE
K.M. Kochummen
Kepong, Malaysia
A.W. Bennett in Hooker f., Fl. Brit. Ind. 1 (1875) 527; King, J. As. Soc. Beng. 62, 2 (1893) 235;
Merrill, EB (1921) 316, PEB (1929) 116; Ridley, FMP 1 (1922) 368; H.J. Lam, Bull. Jard. Bot. Btzg.
3, 12 (1932) 281; Masamune, EPB (1942) 362; Browne, FTSB (1955) 70; Leenhouts, FM 1, 5 (1956)
209, FM 1, 6 (1972) 917, FM 1, 7 (1976) 820; Backer & Bakhuizen f., FJ 2 (1965) 112; Smythies,
CST (1965) 30; Burgess, TBS (1966) 60; Kochummen, TFM 1 (1972) 121; Cockburn, TS 1 (1976) 34;
Anderson, CLTS (1980) 155; Wong, DMT (1982) 30; Corner, WSTM 2 (1988) 199; Whitmore, Tantra
& Sutisna, CLK 1 (1989) 34; Ng, Mal. For. Rec. 34, 1 (1991) 35.
Medium-sized to large buttressed trees, rarely shrubs; dioecious or occasionally monoecious;

crushed living parts with strong resinous smell. Bark pale grey to brown, smooth or scaly,
often lenticellate, surface sometimes with black or white resinous gum; inner bark pinkish,
or creamy, laminated or mottled, with droplets of clear or white resinous gum, with strong
resinous smell in most species (especially those of Canarium and Triomma). Sapwood often
whitish, shiny, sometimes pinkish or yellowish brown, vessels usually visible to the unaided
eye. Pith of twigs, petioles and petiolules often with vascular strands and resin ducts.
Leaves spiral, usually crowded at the tips of twigs, pinnately compound with opposite
leaflets and a terminal leaflet, rarely trifoliolate; petiolules of leaflets with distinct swellings
at both ends in most species (leaflets subsessile in Garuga); margin entire except in some
Canarium and Garuga species. Stipules present in all species of Garuga and in most species
of Canarium. Inflorescences usually axillary or terminal panicles, sometimes spikes,
racemes or thyrses. Flowers unisexual (except in Garuga), male and female on different
trees, 3- or 5-merous, usually greenish to cream, the remains of the other sex persisting;
sepals valvate, mostly united; petals valvate, free; stamens usually twice as many as petals,
filaments free or united, sometimes fused to the disc, anthers dehiscing inwards; disc
intrastaminal (except in Triomma), nectariferous, brightly coloured; ovary superior, with
35 cells, each cell with 2 ovules; style simple, stigma globular, often slightly lobed. Fruit a
drupe with a fleshy or leathery resinous rind and a more-or-less woody stone (pyrene) or in
Triomma a woody capsule; stone 3-celled or less. Seed one; cotyledons fleshy, variously
folded, rolled and convoluted; endosperm absent; germination epigeal or hypogeal, mostly
rapid; cotyledons divided into 3, 5 or more lobes (entire in Scutinanthe). Seedling with first
2 leaves opposite or alternate, subsequent leaves alternate, spiral, simple for several nodes,
then pinnate (in some Santiria the leaves are pinnate from start); sapling leaflets thinner,
with long drawn-out tips, longer petioles and more distantly spaced when compared to the
adult leaves.
Distribution. 16 genera and about 550 species; tropics and subtropics. In Sabah and Sarawak,
8 genera and 59 species are known.
45
Ecology. Burseraceae are common constituents of the main storey of mixed dipterocarp and
kerangas forests; they also occur in submontane forests to 1800 m. The drupaceous fruits
are dispersed by animals. In Triomma the winged seeds are dispersed by wind.
Timber. The standard Malaysian name for the timber of Burseraceae is kedondong (Malay).
It is a very homogeneous group except for slight colour differences, and a light hardwood,
moderately hard and moderately heavy; in strength Class `C, strong; working quality
variable, some easy, others difficult and blunting tools excessively. Not durable in tropical
conditions. Moderately susceptible to powder post-beetles. Difficult to treat with preservatives.
Seasoning without serious degrade, with low shrinkage. Sapwood pale, not sharply defined;
heartwood yellow-brown, pink-brown or red-brown, only slightly darkening on exposure,
surface mostly glossy, without figure; grain interlocked. Texture moderately fine and even.
Most species of Dacryodes, Santiria and Scutinanthe have siliceous timbers while Canarium
(except C. apertum) and Triomma have no silica in the timber.
Uses. Timber suitable for general building construction and carpentry work, plywood and
weather boarding. The family abounds in fragrant balsams and resins (e.g., balsam from
Commiphora opobalsamum; frankincense from Boswellia species especially B. sacra; myrrh
from Commiphora myrra). The resins are used in traditional medicine in Peninsular
Malaysia. The resin from Canarium luzonicum, the Manila-elemi is used in pharmacy in
ointments and plasters. The oil from the kernel is used locally. Commercial quantities of the
oil are now being produced in the Solomon Islands from Canarium species for industrial use
in the manufacture of skin- and hair-care products under the trade name "Solomon Nut Oil".
The seeds of Canarium album, Canarium indicum (ngali-nut; Solomon Islands), C. ovatum
and C. vulgare (kenari, Indonesian) are eaten. Certain species, e.g., C. album, C. ovatum
and C. vulgare, are planted as avenue trees, or for wind-breaks and also as shade-trees for
nutmeg plantations.
Taxonomy. Although the Burseraceae is taxonomically closely related to the Meliaceae and
Simaroubaceae it is liable to be confused with the pinnate-leaved species of the Anacardiaceae.
The sole distinguishing character is the number of ovules per cell which is two in every cell
in the Burseraceae and one in the Anacardiaceae. Vegetatively, the swollen petiolule is a
good diagnostic feature for most members of the Burseraceae. The family is divided into 3
tribes. Except for Triomma which belongs to tribe Bursereae with capsular fruit, all the
genera in Borneo belong to the tribe Canarieae with a drupaceous fruit. Triomma retains a
capsular fruit which must be the primitive fruit of the family and which is perhaps to be
regarded as an ancient relic character.
Key to genera
1.
Flowers 45-merous. Fruit with 1 or more pyrenes..................................................2

Flowers 3-merous. Fruit with 1 pyrene.........................................................................5
2.
Flowers bisexual. Stipellae present. Leaflet margin toothed...........................3. Garuga

Flowers unisexual. Stipellae absent. Leaflet margin entire or occasionally toothed.......3
3.
Stamens 5; disc extrastaminal. Fruit a 3-winged capsule. Leaflets withering yellow,

with pink veins....................................................................................................8. Triomma
46
BURSERACEAE (KOCHUMMEN)
Stamens 8 or 10; disc intrastaminal. Fruit a globular or ellipsoid drupe. Leaflets not
so.................................................................................................................................4
4.
Receptacle of flower cup-shaped; ovary 3-celled.....................................7. Scutinanthe

Receptacle flat; ovary 45-celled..................................................................5. Protium
5.
Fruit thick-walled, bony, seated on enlarged calyx. Leaves often stipulate, leaflet
margin often toothed.................................................................................1. Canarium
Fruit thin-walled, calyx not enlarged in fruit. Leaves without stipules, leaflet margin
entire...........................................................................................................................6
6.
Remains of stigma on the fruit always distinctly off-centre. Fruit surface smooth when
dry................................................................................................................6. Santiria
Remains of stigma on the fruit not off-centre. Fruit smooth or coarsely wrinkled when
dry............................................................................................................................................7
7.
Dry fruits smooth. Cotyledons entire. Inflorescences axillary, to 10 cm long...4.

Haplolobus
Dry fruits coarsely wrinkled. Cotyledons divided. Inflorescences axillary and terminal,
much longer.......................................................................................................2. Dacryodes
1. CANARIUM Stickman
(from the Moluccan name kenari)
kedondong (Malay)
Herb. Amb. (1754) 10 (erroneously Cenarium); King l.c. 236; Ridley l.c. (1922) 369; H.J. Lam, Bull.
Jard. Bot. Btzg. 3, 12 (1932) 422; Burkill, EPMP 1 (1935) 424; Browne l.c. 70; Leenhouts l.c. (1956)
249, l.c. (1972) 92, Blumea 9 (1959) 275; Backer & Bakhuizen f. l.c. 114; Kochummen Bhimea 8
(1955) l.c. (1972) 126, Sandakania 5 (1994) 73; Cockburn l.c 37; Anderson l.c. 155; Corner l.c.
200; Wong l.c. 30; Whitmore, Tantra & Sutisna l.c. 34; Ng l.c. 37.
Medium-sized to large buttressed trees, rarely shrubs. Bark grey-fawn or light yellowbrown, smooth, scaly or dippled, with many small lenticels; inner bark pinkish or reddish,
laminated, soft, aromatic, with clear sticky exudate, rarely with non-sticky exudate, which
becomes dark brown on exposure. Sapwood whitish, darkening inwards. Twigs usually
round, the pith nearly always with vascular strands. Leaves often with stipules of various
shapes and sizes, at base of rachis or on it, soon falling off; leaflets generally with pointed
tips, the margin entire, dentate, or serrate, basal pair of leaflets usually smaller; petiole
rounded, flattened or channelled especially towards base, often swollen at base. Inflorescences
axillary or terminal panicles, spikes, racemes, or thyrses, female ones often reduced and
smaller. Flowers unisexual, male and female on separate trees, 3-merous; receptacle flat or
concave; calyx cup-shaped, lobes deltoid, outside glabrous or hairy, inside always densely
silky-hairy; petals 3, creamy, free, usually overlapping in bud, nearly always ovoid-oblong,
the tip inflexed, fleshy and thick with thin margins, outside hairy in the centre, inside usually
glabrous; stamens 6 in one whorl, rarely only 3, free to entirely connate, sometimes adnate
to the disc, in female flowers sterile and often less-developed, filaments flattened, anthers
47
opening by a longitudinal slit; disc inside to the stamens (intrastaminal), 6-lobed, strongly
developed in male flowers; ovary in female flowers stalked if receptacle is concave, ovoid
to ellipsoid, style cylindrical, stigma globular, slightly 3-lobed; in the male much reduced,
often fused to disc. Fruit a drupe seated on persistent enlarged calyx, with apical stigma,
blue black when ripe (rarely ivory white or red), hairy especially near base and apex, or
glabrous; pericarp fleshy or fibrous, wrinkled when dry; stone hard, woody, round or
triangular in cross-section, with 3 cells, often reduced to 2 or 1, containing woody intrusions
of the placenta, penetrating between cells as wings, often visible as surface ribs. Seed 1 per
cell; testa brown; without endosperm; cotyledons oily, 3-lobed in most species but 5-lobed
in C. megalanthum.
Distribution. c. 100 species; tropical W and E Africa, Madagascar, Mauritius, Sri Lanka,
SE Asia from S Deccan to S China and Hainan, Malesia, NE Australia and Melanesia; 23
species in Sabah and Sarawak.
Ecology. Common in the lowland mixed dipterocarp forests, rare in submontane forests to
1800 m.
Uses. Canarium species produce general utility timber. Burkill (l.c.) documents some of the
uses. C. pimela is cultivated in SE China as an ornamental and as a fruit tree. The fruits are
highly esteemed by the Chinese. C. indicum, C. ovatum and C. vulgare are planted for their
nuts. They are an important constituent of the diet in the Solomon Islands. In Malaysia and
China they are a valued titbit. In Sarawak C. odontophyllum is widely cultivated for the
edible fruits. C. pseudodecumanum has edible fruits; the oil pressed out of the seeds is
locally used. The resin is used in pharmacy in ointments and plasters, mainly from C.
luzonicum, the "Manila-elemi". It is also a constituent of cellulose lacquers. The shell of the
nut is used as fuel substitute in the Solomon Islands. Very little is known about the
economic value of the other species.
Taxonomy. The genus can be subdivided into 3 subgenera: subgenus Canarium consisting
of sections Canarium and Pimela is centred in Malesia, subgenus Africanarium in W Africa,
and subgenus Canariellum restricted to E Queensland and New Caledonia. In Sabah and
Sarwak, section Canarium can be recognised by its foliaceous or pectinate (comb-like)
stipules, its leaflets which dry brown, its stamens that are free or adnate to the disc, the
usually glabrous disc and pistil, and the larger fruit (57 cm long) with frequently tomentose
calyx. In contrast, section Pimela has lanceolate or narrow stipules, leaflets which dry
greyish green, stamens that are often partly or entirely connate, a disc and pistil that are
mostly pilose, and smaller fruit (often 2.53.5 cm long) with usually glabrous calyx.
Key to Canarium species

1.
Leaflets glabrous..........................................................................................................2
Leaflets hairy............................................................................................................16
2.
Leaflet margin toothed...............................................................................................3

Leaflet margin entire........................................................................................................6
48
3.
Leaflets distinctly whitish below...............................................................13. C. littorale

Leaflets not so, rarely faintly whitish below....................................................................4
4.
Stipules absent. Leaflet base rounded, margin faintly toothed...20. C.

pseudodecumanum
Stipules present. Leaflet base not rounded, margin distinctly toothed...........................5
5.
Stipules deeply irregularly lobed. Rachis yellowish..........................5. C. denticulatum

Stipules suborbicular, not lobed. Rachis brownish........................10. C. kinabaluensis
6.
Leaflets sessile, lateral veins perpendicular to midrib. Trees with stilt-roots

........................................................................................................4. C. decumanum
Leaflets stalked, lateral veins not so. Trees without stilt-roots.....................................7
7.
Plants without stipules.................................................................................................8

Plants with stipules....................................................................................................10
8.
Lateral veins 57 pairs. Leaflet apex with 12-cm-long tip...........22. C. pseudopimela

Lateral veins 714 pairs. Leaflet apex without such long tips......................................9
9.
Twigs with prominent leaf-scars. Petals clawed...............21. C. pseudopatentinervium

Twigs without prominent leaf-scars. Petals not clawed...........................19. C. pilosum
10. Stipules needle-like...................................................................................................11

Stipules broad and flat...............................................................................................13
11. Rachis black. Flowers reddish when fresh.........................................6. C. dichotomum
Rachis not black. Flowers not reddish when fresh......................................................12
12. Leaflet stalk about 5 mm long; lateral veins 1518 pairs................11. C. kostermansii
Leaflet stalk longer, to 20 mm long; lateral veins 611 pairs...................2. C. asperum
13. Leaves faintly glaucous below................................................................14. C. littorale
Leaves not glaucous below.........................................................................................14
14. Lateral veins and reticulations invisible......................................18. C. patentinervium
Lateral veins and reticulations visible........................................................................15
15. Lateral veins prominently raised above; leaflet base usually rounded.....7. C. divergens
Lateral veins not raised; leaflet base cuneate.........................................3. C. caudatum
16. Leaflet margin toothed...............................................................................................17
Leaflet margin entire.................................................................................................24
17. Leaflets sessile, margin minutely toothed.............................20. C. pseudodecumanum
Leaflets stalked, margin coarsely and distinctly toothed.............................................18
18. Stipules absent...........................................................................................................19
Stipules present........................................................................................................21
49
19. Apex of leaflet blunt, rounded or notched................................................1. C. apertum

Apex of leaflet pointed...............................................................................................20
20. Leaflet margin curled inwards.........................................................9. C. grandifolium

Leaflet margin not so...................................................................23. C. sarawakanum
21. Stipules linear. Twigs woolly..................................................................19. C. pilosum
Stipules broad and flat. Twigs not woolly...................................................................22
22. Stipules deeply incised................................................................17. C. odontophyllum
Stipules not deeply incised.........................................................................................23
23. Stipules kidney-shaped with wavy margin..............................................14. C. littorale
Stipules rounded to ovate, margin not wavy..................................13. C. latistipulatum
24. Stipules 34-lobed. Leaflet base rounded......................................15. C. megalanthum
Stipules linear, not lobed. Leaflet base not rounded....................................................25
25. Leaflet margin curled inwards....................................................8. C. fusco-calycinum
Leaflet margin not curled inwards..............................................................................26
26. Ultimate leafy twigs stout, more than 1 cm thick. Leaflets (except basal ones) subsessile, base rounded or subcordate..................................................... . 10. C. hirsutum
Ultimate leafy twigs slender, less than 1 cm thick. Leaflets distinctly stalked, base
cuneate..................................................................................................16. C. merrillii
1. Canarium apertum H.J. Lam

(Latin, apertus = open or unshielded; the basal gaps between petals)
Ann. Jard. Bot. Btzg. 42 (1932) 214, Bull. Jard. Bot. Btzg. 3, 12 (1932) 491; Masamune l.c. 362;
Leenhouts l.c. (1956) 275, l.c. (1959) 386; Burgess l.c. 60; Kochummen l.c. (1972) 128; Anderson l.c.
155; Whitmore, Tantra & Sutisna l.c. 34. Type: Beccari PB 1630, Sarawak, Matang (holotype FI;
isotypes K, L). Synonym: Santiria serrulata Engl. in DC., Mon. Phan. 4 (1883) 160; non C.
serrulatum Miq., Fl. Ind. Bat. 1, 2 (1859) 646.
Very large tree to 40 m tall, 80 cm diameter; buttresses to 2 m high. Bark grey-brown,

lenticellate, scaly; inner bark yellow-brown. Twigs angled, powdery reddish brown hairy
when young. Stipules absent. Leaves with 34 pairs of leaflets; rachis and petiole powdery
brown-hairy; blade oblong, elliptic or ovate, 4.510.5 x 35.5 cm, sparsely rough-hairy
below and on midrib above; base rounded to subcordate, margin finely toothed towards
apex, apex blunt or rounded; midrib flattened above; lateral veins 815 pairs, prominently
raised below, giving a bullate appearance to the blade, flat above; intercostal veins reticulate,
raised below, faint above; petiolules 23 mm long, powdery hairy. Flowers (female)
yellow-brown, in terminal broadly thyrsoid panicles; petals clawed. Fruits ovoid, pointed,
hairy, circular in cross-section, 45 x 22.5 cm.
Distribution. Sumatra, Peninsular Malaysia and Borneo. Uncommon in Sabah, recorded from
Beaufort, Keningau, Sandakan, Tawau and Tenom. In Sarawak, reported from Belaga, Lundu,
Miri and the Sabal Forest Reserve. Also occurs in Kalimantan.
Ecology. In lowand mixed dipterocarp forests to 500 m, on clay-rich and yellow sandy soils
and alluvium. Flowers collected in January and April and fruits in May and October.
50
2. Canarium asperum Benth.

(Latin, asper = rough, uneven; possibly the rough-hairy inflorescence)
in Hooker, Lond. J. Bot. 2 (1843) 215; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 461; Masamune
l.c. 362; Leenhouts l.c. (1956) 293, l.c. (1959) 439; Backer & Bakhuizen f. l.c. 115; Whitmore, Tantra
& Sutisna l.c. 34. Type: Hinds, s.n., New Guinea (K).
subsp. asperum var. asperum

Synonyms: Canariopsis aspera (Benth.) Miq., Fl. Ind. Bat. 1, 2 (1859) 653; Canarium
villosum Benth. & Hook. f. ex F.-Vill., Nov. App. (1880) 40; C. molle Engl. in DC. l.c. 109
(for a complete list of synonyms, cf. Leenhouts l.c. (1956).
Small to medium-sized tree to 30 m tall, 50 cm diameter. Bark pale brown, scaly with large
flakes; inner bark dull brown with white sticky exudate. Sapwood cream. Stipules needlelike, inserted near the base of petiole. Leaves with up to 6 pairs of leaflets; rachis glabrous,
rarely hairy; petiolules of lateral leaflets 0.52 cm long, swollen towards apex (only 2 mm
long on collections from Pulau Gaya), rarely hairy; blade elliptic to lanceolate, 7.516 x
46.5 cm; base rounded to broadly cuneate, slightly unequal, margin entire, apex pointed;
midrib raised above; lateral veins 611 pairs, raised below, faintly raised above; intercostal
veins scalariform-reticulate, raised below, faint above. Flowers in axillary spikes or
racemes. Fruits ovoid to subglobose, 914 x 411 mm.
Distribution. E Java, Borneo, Lesser Sunda Islands, Philippines, Celebes, Moluccas, New
Guinea and the Solomon Islands. In Sabah uncommon, recorded only from Pulau Selipol, P.
Gaya, P. Banggi, Lahad Datu and Kudat. Not yet recorded from Sarawak. Also known from
Kalimantan.
Ecology. Rocky coasts and inland lowland mixed dipterocarp forests.
Leenhouts l.c. (1956 & 1959) recognised two subspecies, viz. subsp. asperum and papuanum, of
which only subsp. asperum occurs in Sabah and Sarawak. Of subsp. asperum, he distinguished
two varieties, namely var. asperum and var. clementis with the latter endemic to the
Philippines.
3. Canarium caudatum King

(Latin, caudatus = tailed; the gradually tapering leaflet apex)
l.c. 240; Ridley l.c. (1922) 370, Kew Bull. (1930) 81; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932)
443; Masamune l.c. 362; Leenhouts l.c. (1956) 259, l.c. (1959) 343; Kochummen l.c. (1972) 129.
Type: King's collector 10227, Perak (holotype CAL; isotypes A, BM, G, L).
Medium-sized to large tree to 36 m tall, 40 cm diameter. Bark grey, cracking and scaly;
inner bark with strong resinous smell. Stipule usually falling off early, kidney-shaped.
Leaves with 24 pairs of leaflets; petiolules of lateral leaflets c. 1 cm long, swollen at both
ends; blade elliptic or ovate, 5.516.5 x 38 cm; base rounded or broadly cuneate, margin
entire, apex pointed, tip c. 1 cm long; midrib raised above; lateral veins 811 pairs, distinct
51
on both surfaces; intercostal veins reticulate, visible on both sides. Flowers in terminal
thyrsoid inflorescences; stamens 3 in male flowers and 6 in female flowers. Fruits spindleshaped, 5.58 x 23.5 cm, with saucer-shaped persistent calyx.
Key to forms
Stipules small, kidney-shaped, inserted partly on the twig....................................................
forma caudatum
Synonym: C. pauciflorum Ridl. l.c. (1930) 80.
Sumatra, Peninsular Malaysia and Borneo. Of scattered distribution in Sabah and Sarawak.
Stipules auricle-shaped, inserted on the petiole.....................................................................
forma auriculiferum Leenh.
Blumea 8 (1955) 181. Type: Haviland 2877, Sarawak (holotype SING; isotype BM, K, L, SAR).
Sumatra, Peninsular Malaysia, and Borneo. In Borneo uncommon, known by a few

collections from Sabah (SAN 33828), Sarawak (e.g., S. 5840, S. 12032 and S. 46003), and
Kalimantan.
Ecology. Lowland mixed dipterocarp and kerangas forests to 230 m.
4. Canarium decumanum Gaertn.

(Latin, decumanus = the largest or greatest; the size of the fruit and tree)
Fruct. 2 (1791) 99; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 435; Leenhouts l.c. (1956) 276, l.c.
(1959) 389; Burgess l.c. 60; Cockburn l.c. 39; Whitmore, Tantra & Sutisna l.c. 34. Type: Hort. Bot.
Bog. VI E 5 (= Pl. Bog. Exsicc. 116), Moluccas (neotype L; isoneotypes B, BO, BRSL, G, K, L, NY).
Very large tree to 54 m tall, 150 cm diameter; buttresses to 5 m tall, stilt-roots present. Bark
grey, smooth to scaly or dippled; inner bark orange-red, granular, with strong mango smell,
exudate brownish and sticky. Sapwood white. Twigs pale white with prominent leaf-scars.
Stipules inconspicuous. Leaves closely spirally arranged, with 45 pairs of almost sessile
leaflets; rachis powdery yellowish hairy; blade oblong, elliptic or ovate, 615.5 x 37 cm;
base rounded or subcordate, margin entire, apex pointed; midrib flattened above; lateral
veins 1223 pairs, almost perpendicular to midrib, forking and forming reticulations a few
millimeters away from margin, visible on both surfaces; intercostal veins scalariformreticulate, very faint; petiole flattened above with sharp edges. Flowers hairy, in axillary
thyrsoid inflorescences. Fruits ellipsoid, 78.5 x 4.56 cm, rough hairy.
Vernacular name. Sabahpamatudon (Malay, Dusun).
Distribution. Borneo, Moluccas, New Guinea. In Sabah uncommon, recorded from Lahad
Datu and Sandakan; not yet reported from Sarawak; also known in Kalimantan.
Ecology. Lowland forests.
52
5. Canarium denticulatum Blume

(Latin, denticulatus = with very small teeth; the leaflet margin)
Bijdr. (1826) 1162; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 521; Masamune l.c. 362; Leenhouts
l.c. (1956) 272, l.c. (1959) 367; Backer & Bakhuizen f. l.c. 115; Burgess l.c. 60; Kochummen l.c.
(1972) 129; Cockburn l.c. 41; Whitmore, Tantra & Sutisna l.c. 34. Type: Blume 743, Java (L,
lectotype).
Medium-sized tree to 30 m tall, 50 cm diameter; buttresses short. Bark grey-white, smooth;

middle bark green; inner bark pinkish. Sapwood white. Twig pale whitish. Stipules persistent,
yellowish, inserted on petiole, deeply irregularly lobed. Leaves with 26 pairs of leaflets;
rachis pale yellow; petiolules of lateral leaflets 815 mm long, yellowish (rachis hairy in
saplings); blade elliptic or oblong, 623.5 x 39 cm; base unequal, rounded or cuneate,
margin faintly toothed towards apex, apex pointed; midrib raised above; lateral veins 913
pairs, curving and joining near margin, visible on both surfaces; intercostal veins reticulate,
faintly visible on both surfaces; glabrous or reddish brown-hairy on the lower side. Flowers
white, hairy, in an axillary thyrsoid inflorescence. Fruits ellipsoid, 2.53 x 1.5 cm.
Key to subspecies
Leaflets reddish brown-hairy below, margin toothed.............................................................
subsp. kostermansii Leenh.
Blumea 8 (1955) 181. Type: Kostermans 5226, E Borneo, Sg. Menubar region (holotype L;
isotypes BM, BO, K).
Endemic to Borneo. Uncommon, in Sabah only known from several collections, all
from the east coast districts; also in Kalimantan.
Leaflets glabrous, margin entire or faintly toothed................................................................
subsp. denticulatum
Synonyms: C. fissistipulum Miq., Fl. Ind. Bat. Suppl. (1861) 521; C. kunstleri King l.c. 184; C.
laciniatum Elmer, Leafl. Philip. Bot. 3 (1911) 1084.
Andamans, Burma, Sumatra, Peninsular Malaysia, Borneo, and the Philippines. In Sabah
reported from Ranau, Kalabakan, Kinabatangan, Lahad Datu, Sandakan, Tawau. In
Sarawak known from Batu Laga, Bt. Mersing, Ulu Tinjar, Bt. Raya, Ulu Rajang, Segam
FR, Lambir NP and Marudi; also known in Kalimantan. Common in the lowlands, rarely
to 750 m, in mixed dipterocarp forest on friable fertile soils, particularly on basic volcanic
rocks.
6. Canarium dichotomum (Blume) Miq.

(Latin, dichotomus = having divisions always in pairs; branching of the inflorescence)
l.c. (1859) 648; Merrill l.c. (1929) 116; H.J. Lam, Bull Jard. Bot. Btzg. 3, 12 (1932) 447; Masamune
l.c. 363; Leenhouts l.c. (1956) 283, l.c. (1959) 423; Burgess l.c. 60; Anderson l.c. 155; Whitmore,
Tantra & Sutisna l.c 34. Basionym: Pimela dichotoma Blume, Mus. Bot. Lugd. Bat. 1 (1850) 222.
Type: Korthals 957, Sumatra (holotype L; isotype U). Synonym: C. endertii H.J. Lam, Ann. Jard.
Bot. Btzg. 42 (1932) 210.
Medium-sized tree to 27 m tall, 25 cm diameter. Bark brownish to reddish, smooth to scaly;

inner bark reddish brown. Sapwood pale white. Twigs 0.52(2.5) cm thick, grey or dark
brown to blackish, powdery brown-hairy when young. Stipules linear, hairy, on the base of
rachis, persistent. Leaves with 14 (rarely 68) pairs of leaflets; rachis black, powdery
53
brown-hairy; petiolules of lateral leaflets 0.52 cm long, hairy; blade elliptic to lanceolate or
oblong, 820 x 49.5 cm, sparsely hairy below or glabrous; base broadly cuneate, unequal,
margin entire, apex pointed, tip c. 1 cm long; midrib raised above; lateral veins 915 pairs,
raised below, distinct above; intercostal veins reticulate, faintly visible on both surfaces.
Flowers in terminal and axillary thyrsoid inflorescences, main branches in male
inflorescences repeatedly dichotomously branched, axis reddish. Fruits oblong, 34 x
1.21.5 cm, triangular; calyx funnel-shaped.
Distribution. Sumatra, Borneo. Of scattered distribution in Sabah and Sarawak. Also known in
Kalimantan.
Ecology. Lowland forest, rarely in submontane forest to 1200 m.
7. Canarium divergens Engl.

(Latin, divergium = going different ways; the laxly branched inflorescence)
in DC. l.c. 143; Merrill l.c. (1921) 316; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 503; Masamune
l.c. 363; Leenhouts l.c. (1956) 260, l.c. (1959) 346; Anderson l.c. 155. Type: Beccari PB 2237,
Sarawak, Matang (holotype P; isotypes FI, K).
Medium-sized tree to 30 m tall, 45 cm diameter; buttresses short. Bark grey-white,

lenticellate, scaly. Twigs brownish, glabrous. Stipules deciduous. Leaves with 24 pairs of
leaflets; petiolules of lateral leaflets 715 mm long; blade elliptic, ovate or obovate, 718 x
3.59.5 cm, glabrous; base cuneate to rounded, margin entire to faintly toothed, apex
pointed, tip c. 1 cm long; midrib raised above; lateral veins 713 pairs, raised below, faint
above, curving and joining near margin; intercostal veins scalariform-reticulate, raised
below, faint above. Inflorescences thyrsoid, terminal and axillary; male to 37 cm long,
hairy. Fruits yellowish green when fresh, ellipsoid, 68.5 x 2.53.5 cm; calyx funnelshaped with wavy margin.
Distribution. Endemic to Borneo. Uncommon in Sabah, known by a single collection SAN
50076 from Beaufort; more common in Sarawak, (Semengoh, G. Buri, Matang, Lambir
National Park, Bintulu and Ulu Anap); also recorded from Brunei.
Ecology. Mixed dipterocarp forests on deep sandy humult ultisols, to 450 m.
8. Canarium fusco-calycinum Stapf ex Ridl.

(Latin, fuscus = dark or dark brown, calycinus = calyx; the colour of the calyx)
l.c. (1930) 82; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 499; Masamune l.c. 363; Leenhouts l.c.
(1956) 283, l.c. (1959) 424; Anderson l.c. 155. Type: Haviland 1981, Sarawak, Kuching (holotype K;
isotypes BM, SAR).
Medium-sized tree to 20 m tall, 50 cm diameter. Twigs rusty brown-hairy. Stipules linear,

falling off early. Leaves with 2 pairs of leaflets; rachis hairy; petiolules of lateral leaflets
23 mm long, hairy; blade hairy below, ovate to oblong, 4.515 x 39 cm; base broadly
cuneate, margin entire, incurled, apex pointed, tip c. 1 cm long; midrib raised above, hairy;
lateral veins 1217 pairs, raised below, visible above, curving and joining near margin;
54
intercostal veins scalariform-reticulate, raised below, faint above. Flowers (male) in terminal
thyrsoid inflorescences, hairy; stamens united. Fruits ellipsoid, 33.5 x 1.21.5 cm, triangular in cross-section; calyx red-brown hairy, with distinct lobes.
Distribution. Endemic to Sarawak. Uncommon, reported from the 1st, 4th and 7th Div.
9. Canarium grandifolium (Ridl.) H.J. Lam

(Latin, grandis = large; folium = leaf)
Ann. Jard. Bot. Btzg. 42 (1932) 215, Bull. Jard. Bot. Btzg. 3, 12 (1932) 527; Leenhouts l.c. (1956)
275, l.c. (1959) 386; Kochummen l.c. (1972) 129. Basionym: Trigonochlamys grandifolia Ridl., J.
Str. Br. R. As. Soc. 54 (1916) 31, l.c. (1922) 381. Type: Cantley's collector, s.n., Singapore (lectotype
SING).
Medium-sized to large tree to 40 m tall, 60 cm diameter; buttresses tall. Bark greyish,

dippled and scaly. Twigs densely fulvous tomentose. Stipules absent. Leaves with 57 pairs
of leaflets, leaflet stalks glabrous to hairy; blade elliptic to obovate, 1017 x 58 cm,
densely hairy beneath and on midrib above; base cuneate to rounded, margin recurved,
minutely toothed to entire, apex rounded with short tip; midrib sunken above; lateral veins
914 pairs, prominently raised below; intercostal veins equally prominent, sunken above.
Inflorescences terminal, rarely lateral, densely red-tomentose; male thyrsoid, female racemose
to spicate. Flowers pubescent; petals clawed; stamens free. Fruits ellipsoid, c. 5 x 3.5 cm,
slightly hairy at apex.
Distribution. Peninsular Malaysia and Borneo. In Sabah and Sarawak very uncommon,
known only from two collections (SAN 44553 and S. 36641); also known from Brunei.
10. Canarium hirsutum Willd.

(Latin, hirsutus = covered with rough hairs; the fruit)
Sp. Pl. 4 (1805) 760; Ridley l.c. (1922) 374; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 466;
Masamune l.c. 363; Leenhouts l.c. (1956) 287, l.c. (1959) 424; Backer & Bakhuizen f. l.c. 115;
Burgess l.c. 60; Kochummen l.c. (1972) 130; Cockburn l.c. 42; Anderson l.c. 155; Whitmore, Tantra
& Sutisna l.c. 35. Type: bb. 33772, Moluccas (neotype L; isoneotypes A, BO, SING). Synonyms: C.
hispidum Blume, Cat. (1823) 109; C. subcordatum Ridl. l.c. (1922) 374.
Medium-sized tree to 25 m tall, 60 cm diameter. Twigs very stout, 1.53.5 cm thick, reddish
brown-hairy near tip, pith large with many small vascular strands. Stipules present (except
in two varieties), caducous, inserted on the petiole, subulate, 412 mm long. Leaves to 2 m
long, with 413 pairs of leaflets; blade subsessile except basal ones, glabrous or hairy, ovate
to lanceolate, 545 x 2.515 cm; base rounded to cordate, margin entire, apex pointed;
midrib raised above; lateral veins 1230 pairs, visible on both surfaces; intercostal veins
55
finely reticulate, visible; petiole to 2 cm thick near base with sharp edges. Inflorescences
axillary, male thyrsoid, female subracemose. Flowers c. 1 cm long, shortly stalked. Fruits
with irritant reddish brown hairs, oblong to ovoid, 2.83.5 x 22.5 cm.
Distribution. Throughout Malesia (except Lesser Sunda Islands), Carolines and Solomon
Islands. In Sabah rather common, but in Sarawak very uncommon and represented by a
single collection (S. 25192). Also known in Kalimantan.
Ecology. Mixed dipterocarp forest, rarely to 1800 m.
Two subspecies and a few varieties are recognised for this species. In Sabah and Sarawak
only subspecies hirsutum is present with its variety hirsutum and forma scabrum (Blume)
Leenh.
11. Canarium kinabaluensis Leenh.

(of Mt. Kinabalu)
l.c. (1955) 182, l.c. (1956) 260, l.c. (1959) 346; Burgess l.c. 60; Whitmore, Tantra & Sutisna l.c. 35.
Type: Ramos 1698, North Borneo, Sandakan (holotype L; isotypes A, BO, BM, K, P).
Medium-sized tree to 25 m tall, 50 cm diameter; buttresses steep, to 1 m high. Bark grey,

smooth; inner bark orange with strong resinous smell. Sapwood white. Twigs blackish.
Stipules suborbicular, inserted on the base of the petiole. Leaves with 24 pairs of leaflets;
rachis black, cracking; petiolule black on drying; blade drying to reddish brown, thickly
leathery, elliptic-ovate or lanceolate, 820 x 3.510 cm; base broadly cuneate, margin very
faintly toothed, apex pointed; midrib raised above, sharply keeled below; lateral veins 11
13 pairs, raised below; intercostal veins scalariform. Inflorescences and flowers unknown.
Fruits spindle-shaped, green drying brown, strongly wrinkled, 68 x 24 cm. Seeds one.
Distribution. Endemic to Borneo. Uncommon, collected from Papar, Sandakan and Mt.
Kinabalu in Sabah; also known from Kalimantan.
Ecology. Lowland to submontane forests at 3001500 m.
12. Canarium kostermansii Leenh.

(A.J.G.H. Kostermans, 19071994, botanist at the Forest Research Institute Bogor and the
Herbarium Bogoriense, Indonesia)
l.c. (1955) 191, l.c. (1956) 281, l.c. (1959) 398; Burgess l.c. 61; Whitmore, Tantra & Sutisna l.c. 35.
Type: Kostermans 5315, E Borneo (holotype L; isotypes BM, K).
Medium-sized to large tree reaching 35 m tall, 60 cm diameter. Bark brown, smooth. Twigs
slender, to 1 cm thick, pith with 2 concentric cylinders of vascular strands. Stipules subulate,
caducous. Leaves with 47 pairs of leaflets; leaflet stalk about 0.5 cm long; blade
lanceolate, 1022 x 37 cm; base unequal, broadly cuneate, margin entire, apex pointed;
midrib raised above; lateral veins 1518 pairs, faintly curving towards the margin, raised
above; intercostal veins reticulate, distinct on both surfaces. Inflorescences (female) axillary,
racemose. Fruits spindle-shaped, 3 x 11.5 cm, glabrous; calyx funnel-shaped.
56
Distribution. Endemic to Borneo. Uncommon in Sabah and Sarawak; also found in

Kalimantan.
Ecology. Lowlands and hill forests to 900 m.
13. Canarium latistipulatum Ridl.

(Latin, latus = broad, wide, stipula = stipule)
(1956) 258, l.c. (1959) 341; Whitmore, Tantra & Sutisna l.c. 35. Type: Haviland & Hose 2245,
Sarawak, Kuching (holotype K; isotype SAR).
Small to medium-sized tree to 27 m tall. Young twigs powdery brown-hairy. Stipules subpersistent, rounded to ovate, 12 x 11.7 cm, inserted on the base of petiole. Leaves with 3
pairs of leaflets; rachis powdery brown-hairy; petiolule of lateral leaflets 510 mm long,
hairy; blade sparsely hairy below, drying reddish brown, oblong or elliptic, 615 x 3.56
cm; base broadly cuneate, margin toothed, apex pointed, tip c. 1.5 cm long; midrib raised
above; lateral veins 710 pairs, raised below, curving and joining near margin; intercostal
veins scalariform-reticulate, distinct below, faint above. Flowers (male) hairy, in terminal
thyrsoid inflorescences. Fruits oblong, 6.27.2 x 2.23 cm, with tapered base; calyx funnelshaped, powdery brown-hairy.
Distribution. Endemic to Borneo. Common in Sabah but uncommon in Sarawak and known
only by 2 other collections (S. 3458 & S. 52998) beside the type.
14. Canarium littorale Blume

(Latin, littoralis = of the seashore; its main habitat)
l.c. (1826) 1164; Merrill l.c. (1929) 116; H.J. Lam, Bull. Jard. Bot. Btzg. 3,12 (1932) 498; Masamune
l.c. 364; Leenhouts l.c. (1956) 256, l.c. (1959) 337; Backer & Bakhuizen f. l.c. 115; Burgess l.c. 61;
Kochummen l.c. (1972) 130; Cockburn l.c. 42; Anderson l.c. 155; Whitmore, Tantra & Sutisna l.c. 35.
Type: Blume 1736, Java, Nusa Kambangan (holotype L; isotypes BO, MEL, U).
Medium-sized to large tree to 40 m tall, 60 cm diameter. Bark grey, smooth to scaly; inner
bark orange-red or brownish. Sapwood pale white. Twigs glabrous or hairy. Stipules
dropping off early or semi-persistent, kidney-shaped with wavy to deeply lobed margins.
Leaves with 26 pairs of leaflets; rachis glabrous or softly hairy; petiolules of lateral leaflets
12 cm long, glabrous or hairy, often swollen at both ends; blade ovate to oblong or
lanceolate, 325 x 29 cm, glabrous or sparsely to densely hairy below, sometimes white
waxy below; base cuneate to subcordate, margin entire to faintly or prominently toothed,
apex pointed; midrib raised above; lateral veins 920 pairs, sometimes curving and joining
near margin, raised below, faint or sunken above; intercostal veins scalariform-reticulate or
reticulate, distinct below, faint above, rarely sunken above. Flowers usually in terminal
57
thyrsoid (male) or subracemose (female) inflorescences. Fruits ellipsoid or ovoid, 4.57 x

1.53 cm, sparsely hairy or glabrous with rugose and wrinkled surface when dried.
An extremely variable species in vegetative and floral characters. Leenhouts (l.c. 1959)
recognises 5 forms (littorale, pruinosum, purpurascens, rufum, and tomentosum), of which
3 occur in Sabah and Sarawak.
Key to forms
1.
Leaflets whitish beneath................................................................................................

forma pruinosum (Engl.) Leenh.
l.c. (1956) 258. Basionym: C. pruinosum Engl. in DC. l.c. 106, Merrill l.c. (1921) 317,
Masamune l.c. 365. Type: Beccari PB 1970, Sarawak (holotype K; isotype FI).
Confined to Sabah and Sarawak, common in secondary forest, especially on

periodically flooded sandy alluvium.
Leaflets not whitish beneath.........................................................................................2
2.
Leaflets densely hairy below; margin prominently toothed; lateral veins sunken
above...........
forma rufum (A.W. Benn.) Leenh.
l.c. (1959) 339. Basionym: C. rufum A.W. Benn. in Hooker f. l.c. 533, Masamune l.c. 365.
Type: Maingay 1434, Malacca (holotype K; isotypes CAL, L).
Widely distributed in Sabah, less common in Sarawak. Also in Indo-China, Sumatra,

Peninsular Malaysia, and Java.
Leaflets glabrous below; margin faintly toothed; lateral veins not sunken above............
forma littorale
Synonyms: C. glaucum Blume l.c. (1850) 219; C. serricuspe Miq. l.c. (1859) 649; C.
serrulatum Miq. l.c. (1859) 646; C. acutum Engl. in DC. l.c. 113; C. giganteum Engl. in DC.
l.c. 106; C. flavum Ridl. l.c. (1930) 81.
Sumatra, Peninsular Malaysia, Java and Borneo. In Sabah and Sarawak common in
the lowland mixed dipterocarp to submontane forests to 1100 m. Also found in
Kalimantan.
15. Canarium megalanthum Merr.

(Greek, mega = large, anthos = flower)
Philip. J. Sc. 30 (1926) 81; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 446; Masamune l.c. 324;
Leenhouts l.c. (1956) 274, l.c. (1959) 370; Burgess l.c. 61; Kochummen l.c. (1972) 131; Anderson l.c.
155; Whitmore, Tantra & Sutisna l.c. 35. Type: Wood 1213, British North Borneo (holotype UC;
isotypes A, K, L).
Emergent tree to 35 m tall, 50 cm diameter; buttresses present. Bark grey, smooth to

dippled and scaly; inner bark reddish brown. Sapwood yellowish white. Stipules subpersistent, inserted on the petiole near base, 34-lobed. Leaves with 45 pairs of leaflets;
rachis powdery yellowish hairy; petiolules of lateral leaflets 11.5 cm long, powdery hairy;
blade powdery hairy, oblong to lanceolate, 817 x 47 cm; base rounded, margin entire,
apex pointed; midrib flattened above; lateral veins 1012 pairs, raised on both surfaces;
58
intercostal veins scalariform-reticulate, visible below. Flowers hairy, in terminal thyrsoid

inflorescences. Fruits ellipsoid, sparsely hairy, 57.5 x 2.84.5 cm.
Distribution. Sumatra, Peninsular Malaysia, Borneo; uncommon in Sabah and Sarawak.
Also known in Kalimantan.
Ecology. Ridges in mixed dipterocarp forest to 360 m. Flowering in May and fruiting in
May and September.
Uses. In Brunei this species is cultivated for its edible fruits which are among the largest in
the genus.
16. Canarium merrillii H.J. Lam

(E.D. Merrill, 18761954, American botanist)
in Merrill l.c. (1929) 117; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 489; Masamune l.c. 364;
Leenhouts l.c. (1956) 282, l.c. (1959) 402; Burgess l.c. 61; Anderson l.c. 156; Whitmore, Tantra &
Sutisna l.c. 35. Type: Elmer 20326, British North Borneo, Sandakan (holotype PNH; isotypes A, BM,
G, K, L, NY, P, SING, U, Z).
Small to medium-sized tree to 25 m tall, 30 cm diameter. Bark grey, scaly; inner bark
brownish. Sapwood white. Twigs pale grey, rough hairy. Stipules linear, to 5 mm long,
inserted at base of petiole, semi-persistent. Leaves with 45 pairs of leaflets; rachis hairy;
petiolules of lateral leaflets 25 mm long, hairy; blade rough hairy below (like sandpaper to
the touch), elliptic, oblong or obovate, 5.516 x 35.5 cm; base cuneate, unequal, margin
entire, apex pointed, tip c. 1 cm long; midrib faintly sunken above; lateral veins 614 pairs,
curving and joining near margin, raised below, visible above; intercostsal veins reticulate,
distinct below, faint above. Flowers yellow, in terminal or axillary thyrsoid inflorescences.
Fruits ellipsoid, pointed, 34.2 x 1.21.5 cm.
Distribution. Endemic to Borneo. Of scattered distribution in Sabah, Sarawak and
Kalimantan.
Ecology. Mixed dipterocarp to submontane forests to 1400 m.
17. Canarium odontophyllum Miq.
Fig. 1.
(Greek, odontos = toothed, phyllon = leaf; the toothed leaflets)

l.c. (1861) 521; Merrill l.c. (1929) 118; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 519; Masamune
l.c. 364; Leenhouts l.c. (1956) 271, l.c. (1959) 365; Burgess l.c. 60; Cockburn l.c. 43; Anderson l.c.
156; Whitmore, Tantra & Sutisna l.c. 35. Type: Teijsmann HB 692, Sumatra (holotype U; isotype L).
Synonyms: C. beccarii Engl. in DC. l.c. 107; C. palawanense Elmer, Leafl. Philip. Bot. 5 (1913)
1754; C. multifidum H.J. Lam, Ann. Jard. Bot. Btzg. 42 (1932) 215.
59
60
Tree to 30 m tall, 30 cm diameter. Bark grey-brown; inner bark brownish. Sapwood pale.
Twigs stout, whitish, 1.52 cm thick, rusty hairy. Stipules persistent, irregularly toothed or
lobed, c. 6 x 3 cm. Leaves with 38 pairs of leaflets; rachis yellowish hairy; petiolules of
lateral leaflets 310 mm long, hairy; blade densely velvety hairy below, oblong to
lanceolate, 9.528 x 411 cm; base broadly cuneate to rounded, margin toothed, apex
pointed; midrib raised above, hairy; lateral veins 1223 pairs, raised below, faint to sunken
above; intercostal veins scalariform-reticulate, raised below, faint to sunken above. Flowers
hairy, in axillary or terminal thyrsoid inflorescences. Fruits ovoid to ellipsoid, 34 x 2.53
cm.
Distribution. Sumatra, Borneo, and the Philippines. Widely distributed in Sabah, uncommon
in Sarawak. Also known in Kalimantan.
Ecology. Mixed dipterocarp forests on fertile clay soils to 500 m. Flowering in May, June
and November and fruiting in March and August.
Uses. Widely cultivated in Sarawak and Brunei for its fruits.
18. Canarium patentinervium Miq.

(Latin, patens = spreading, nervus = nerves; the leaflet venation)
(1956) 258, l.c. (1959) 342; Burgess l.c. 61; Kochummen l.c. (1972) 134; Whitmore, Tantra & Sutisna
l.c. 36. Type: Teijsmann HB 3736, Sumatra, Palembang (holotype U; isotype L). Synonyms: C.
nitidum A.W. Benn. in Hooker f. l.c. 533; C. parvifolium A.W. Benn. in Hooker f. l.c. 536.
Tree to 24 m tall, 25 cm diameter. Bark grey, smooth. Twigs brownish, lenticellate. Stipules
falling off early, kidney-shaped. Leaves with 24 pairs of leaflets; petiolules of lateral
leaflets 11.5 cm long, cracking; blade thickly leathery, glabrous, drying to dark brown,
ovate to oblong, 712.5 x 2.56 cm; base cuneate, margin entire, apex pointed; midrib
flattened above, rarely sunken; lateral veins 710 pairs, raised below, faint above, curving
and joining near margin; intercostal veins reticulate, faintly visible on both surfaces,
sometimes inconspicuous. Flowers hairy, usually in terminal thyrsoid inflorescences. Fruits
ellipsoid, 66.5 x 2.23 cm, apex pointed.
Distribution. Sumatra, Peninsular Malaysia, and Borneo. Widespread, collected from Tenom,
Tawau and Tuaran in Sabah, and from Lundu, Semengoh to Bukit Lambir in Sarawak. Also
found in Kalimantan.
Ecology. Mixed dipterocarp forest on yellow sandy and leached clay soils, and kerangas
forest, to 450 m.
19. Canarium pilosum A.W. Benn.

(Latin, pilosus = having soft and distinct hairs; the twigs and leaves)
in Hooker f. l.c. 533; King l.c. 243; Ridley l.c. (1922) 372; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12
(1932) 472; Masamune l.c. 365; Leenhouts l.c. (1956) 281, l.c. (1959) 398; Burgess l.c. 61;
Kochummen l.c. (1972) 134; Anderson l.c 156; Whitmore, Tantra & Sutisna l.c. 36. Type: Maingay
3103, Malacca (holotype K; isotype CAL).
Small to medium-sized tree to 27 m tall, 25 cm diameter. Bark pale brown, smooth; inner
bark pink. Sapwood white. Twigs densely woolly hairy. Stipules persistent, linear, inserted
at the base of petiole or on the petiole, rarely absent. Leaves with 23 pairs of leaflets;
rachis densely hairy, rarely glabrous; petiolules of lateral leaflets 23 mm long, densely
hairy, rarely glabrous; blade densely hairy below, sparsely hairy above, rarely completely
61
glabrous, elliptic, oblong or obovate, 719 x 3.59 cm; base cuneate or rounded, margin
faintly to distinctly toothed, rarely entire, apex pointed, tip c. 1 cm long; midrib raised
above; lateral veins 714 pairs, raised on both surfaces, curving and joining near margin;
intercostal veins scalariform-reticulate, raised below, faint above. Flowers hairy, in axillary
or terminal panicles. Fruits oblong, 23 x 11.5 cm.
Key to subspecies
Stipules present. Leaflets hairy below, margin toothed..........................................................
subsp. pilosum
Synonyms: C. grandifolium A.W. Benn. in Hooker f. l.c. 533; C. hirtellum A.W. Benn. in Hooker
f. l.c. 533; C. pilosum var. hirtellum (A.W. Benn.) Ridl. l.c. (1922) 372; C. motleyanum Engl. in
DC. l.c. 133.
Sumatra, Peninsular Malaysia and Borneo. In Sabah and Sarawak uncommon, recorded
from Beaufort, Beluran, Papar, and Sandakan (Sabah), and from Kapit, Marudi, Miri
(Sarawak). Also known in Kalimantan. Lowland forest.
Stipules absent. Leaflets glabrous below, margin entire........................................................
subsp. borneensis Leenh.
l.c. (1955) 193. Type: Clemens 40163, British North Borneo, Mt. Kinabalu (holotype L; isotypes
A, BM, BO, G, NY). Synonym: Dacryodes scandens Husson, Blumea 7 (1952) 164.
Uncommon, known from Mt. Kinabalu, Lahad Datu and Sandakan in Sabah, and G.
Lambir in Sarawak. Also found in Kalimantan.
20. Canarium pseudodecumanum Hochr.

(Latin, pseudo = false; resembling C. decumanum)
Pl. Bog. Exs. (1904) 61; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 438; Leenhouts l.c. (1956) 275,
l.c. (1959) 388; Burgess l.c. 61; Kochummen l.c. (1972) 135; Whitmore, Tantra & Sutisna l.c. 36.
Type: Hort. Bot. Bog. VI E 6a, Sumatra, Lampong (holotype L; isotypes B, BO, BRSL, G, K, L, WY,
P). Synonym: Canarium decumanum (non Gaertn.) Engl. in Merrill l.c. (1929) 116.
Tree to 40 m tall and 120 cm diameter; buttresses to 6 m high. Bark grey-white, smooth to
scaly; inner bark pale. Sapwood yellowish white. Twigs stout, 11.5 cm thick, with large
leaf-scars. Terminal bud 45 cm long, densely brown tomentose. Stipules absent. Leaves
with 46 pairs of leaflets; petiole stout, light brown pubescent, swollen at base and flattened
above; blade almost sessile, lanceolate or oblong, 1723 x 711 cm, glabrous above except
the midrib and lateral veins, densely minutely woolly tomentose below; base rounded or
subcordate, margin minutely toothed, apex pointed; midrib flattened to raised above; lateral
veins 1723 pairs, distinctly arching near margin, prominent below; intercostal veins
prominent giving pitted appearance to the lower surface, veins and reticulations raised
above. Inflorescences (male) axillary thyrses. Flowers pubescent. Fruits ellipsoid, 78 x
46 cm, densely tomentose when young.
Distribution. Sumatra, Peninsular Malaysia, and Borneo. In Sabah recorded from the Ranau,
Sandakan and Tawau districts; not recorded from Sarawak. Also known in Kalimantan.
62
Distribution. Sumatra, Peninsular Malaysia, and Borneo. In Sabah recorded from the Ranau,
Sandakan and Tawau districts; not recorded from Sarawak. Also known in Kalimantan.
Ecology. Lowland forest to 280 m.
Uses. The resin is used for caulking boats. The fruit is edible. An edible oil is extracted from
the seeds.
Closely related to C. decumanum but differing in the absence of stipules and by the toothed
and densely woolly tomentose lower surface of the leaflets.
21. Canarium pseudopatentinervium H.J. Lam

(resembling C. patentinervium)
Leenhouts l.c. (1956) 274, l.c. (1959) 385; Burgess l.c. 61; Anderson l.c. 155; Whitmore, Tantra &
Sutisna l.c. 36. Type: bb. 153, T. 3P. 336, Sumatra, Palembang, Lamatang Ilir near G. Megang
(lectotype BO).
Medium-sized to large tree to 45 m tall, 100 cm diameter; buttresses to 5 m tall. Bark pale
yellow-brown, irregularly scaly to dippled. Twigs whitish, with prominent leaf-scars, pith
with many peripherally arranged vascular strands and often with a central cavity. Stipules
absent. Leaves with 14 pairs of leaflets; blade drying greenish, ovate to elliptic, 512 x
27 cm; base asymmetric, rounded to broadly cuneate, margin entire, apex blunt to pointed;
midrib flattened above; lateral veins 714 pairs, curving near margin; intercostal veins
reticulate, prominent on both surfaces; petiolules of lateral leaflets 59 mm long, not
swollen at both ends. Inflorescences (female) in terminal panicles. Flowers (female) subsessile, densely tomentose, petals distinctly clawed, apex inflexed, stamens free. Fruits
ellipsoid, 57 x 23 cm; calyx triangular, hairy inside.
Distribution. Sumatra, Borneo. In Sarawak uncommon, known only by a single collection
S. 18215 from Belaga; not yet found in Sabah. Also occurs in Kalimantan.
Ecology. Lowland forest at c. 100 m.
22. Canarium pseudopimela Kochummen

(resembling C. pimela)
l.c. (1994) 73. Type: Ding Hou 489, Sarawak, G. Raya (holotype SAR; isotype L).
Small tree to 12 m tall, 15 cm diameter. Twigs brown, c. 3 mm thick, rounded. Stipules

absent. Leaves with 23 pairs of leaflets; blade thinly leathery, elliptic to narrowly obovate,
613 x 2.55 cm, slightly falcate; base cuneate, margin entire, apex cuspidate, tip 1.52 cm
long; midrib raised above; lateral veins 57 pairs, arching and joining near margin, visible
below, faint above; intercostal veins finely reticulate, distinctly visible below, faint above;
petiolules of lateral leaflets 45 mm long, petiolules of terminal leaflet 23.5 cm long.
Inflorescences (female) in racemes, terminal, glabrous, c. 23 cm long, with 1314 cm long
branches in few-flowered racemes, buds obovate. Flowers (female) c. 9 mm long, stalk c. 5
mm long; calyx funnel-shaped, with obtuse lobes, fleshy, with few irregular lines towards
base; petals fleshy, irregularly wrinkled on drying; rudimentary stamens 6, connected into a
cup-shaped staminal ring; disc undulate, c. 1 mm high; ovary hairy, faintly ridged, stigma
63
capitate. Infructescences c. 13 cm long. Fruits ellipsoid, 3 x 11.5 cm, triangular, apex

narrowed to sharp point; calyx almost flat; stalk c. 1.5 cm long.
Distribution. Endemic to Borneo. Uncommon, known by two collections only from G.
Lambir (S. 16613) and G. Raya (Ding Hou 489) in Sarawak.
Very close to C. pimela but differing in the terminal inflorescences, hairy ovary and in the
long-pointed leaflets.
23. Canarium sarawakanum Kochummen

l.c. (1994) 75. Type: Au S. 24802, Sarawak, Kapit (holotype SAR; isotypes A, BO, K, KEP, L, SAN,
SING).
Small tree to 8 m tall, 10 cm diameter. Twigs reddish brown hairy when young. Stipules
absent. Leaves with 12 pairs of leaflets; rachis finely reddish brown hairy; blade thinly
leathery, drying to reddish brown, glabrous above, sparsely reddish brown hairy below,
elliptic to narrowly obovate, 820 x 3.58.5 cm; base cuneate, margin sub-entire to faintly
toothed, apex acuminate, tip 12 cm long; midrib raised above; lateral veins 68 pairs,
distinct below, faintly raised above, curving and joining near margin; intercostal veins
reticulate, visible below, faint above; petiolules of lateral leaflets swollen near the apex,
11.5 cm long, that of terminal leaflet to 5 cm long. Inflorescences (female) axillary or
terminal, thyrsoid, to 22 cm long, with few side branches; rachis brownish hairy. Flowers
(male) with the calyx hairy outside, margins wavy to lobed; petals oblong, c. 4.5 mm long,
hairy outside; stamens 3, joined at the base, filaments to 2.5 mm long; rudimentary ovary
and style present. Fruits (immature) ellipsoid, 4.55.5 x 1.72 cm.
Distribution. Endemic to Borneo. Uncommon, known from a few collections from Kapit,
Lubok Antu, G. Penrissen and Simanggang in Sarawak.
Ecology. Lowland to low submontane forests to 900 m.
Close to C. latistipulatum but that species has semi-persistent stipules.
64
2. DACRYODES Vahl
(Greek, dakruon = a tear; the resin droplets on the bark surface)
kedondong (Sabah and Sarawak Malay), kimayau (Bidayuh in Sarawak)
kembayau (Sabah Dusun), seladah (Sarawak Iban)
Skrift. Dansk. Nat. Hist. Selsk. 4 (1810) 116; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 334;
Kalkman, Blumea 7 (1954) 500; Leenhouts l.c. (1956) 219, l.c. (1972) 917, l.c. (1976) 820; Backer
& Bakhuizen f. l.c. 114; Kochummen l.c. (1972) 136; Cockburn l.c. 43; Anderson l.c. 156; Wong l.c.
42; Whitmore, Tantra & Sutisna l.c. 36; Ng l.c. 37. Synonyms: Pachylobus G. Don, Gen. Syst. 2
(1832) 89; Canarium section Tenuipyrena Engl. in DC. l.c. 104; Curtisiana Ridl., J. Str. Br. R. As.
Soc. 82 (1920) 180; Hemisantiria H.J. Lam in Merrill l.c. (1929) 118.
Small to medium-sized, dioecious trees; buttresses small, short; sometimes with stilt-roots.
Bark smooth to scaly; inner bark orange, yellowish white or pink, exudate often clear,
rarely white. Sapwood pale. Stipules absent (in D. laxa the first pair of leaflets are smaller
and stipule-like). Leaflets entire, the base often strongly unequal; stalks (petiolules) strongly
swollen at both ends. Inflorescence an axillary or terminal panicle. Flowers unisexual, 3merous; sepals free or united; petals usually with thickened inflexed apex; stamens 6,
filaments free, bases united with the disc; disc intrastaminal, glabrous; ovary 3(2)-celled,
usually moderately reduced in male flowers; stigma sessile. Fruit a drupe, oblong or
ellipsoid, 1-seeded; stigma usually apical; pericarp fleshy and thick, coarsely wrinkled when
dry, glabrous; stone containing one fertile and two reduced cells; calyx persistent or
caducous. Seeds round in cross-section; cotyledons 911-lobed in D. rostrata, and 5-lobed
in D. costata, D. laxa and D. rugosa.
Distribution. About 40 species; tropical America, Africa and Asia. In Malesia mainly
centred in Sumatra, Peninsular Malaysia and Borneo; 11 species in Sabah and Sarawak.
Ecology. Lowland (including swamp) to submontane forests to 1500 m.
Uses. The timbers of Dacryodes are essentially similar to that of Canarium in anatomical
structure but they are on the whole heavier and the rays contain silica which makes them
difficult to saw. Fruits of Dacryodes rostrata f. cuspidata, known in Sarawak as keramoh
(Malay) and kembayau (Murut and Iban), are eaten.
Key to Dacryodes species

1.
Rachis and young twigs hairy......................................................................................2

Rachis and young twigs glabrous.................................................................................7
2.
Rachis and young twigs covered with long stiff hairs......................................5. D. laxa
Rachis and young twigs powdery hairy........................................................................3
3.
Ultimate leafy twigs c. 1 cm thick. Terminal bud c. 1.5 cm long.................2. D. elmeri

Ultimate leafy twigs to 0.5 cm thick. Terminal bud to 1 cm long..................................4
65
4.
Intercostal veins prominently raised below. Hairs not yellowish...................................5

Intercostal veins not raised, but visible below. Hairs yellowish...........10. D. rubiginosa
5.
Intercostal veins scalariform. Leaflets densely hairy below.......................8. D. nervosa

Intercostal veins reticulate. Leaflets sparsely hairy below.............................................6
6.
Petiolules strongly swollen at both ends; rachis glabrous..........................11. D. rugosa

Petiolules not strongly swollen at both ends; rachis hairy...........................1. D. costata
7.
Adaxial side of petiole flattened with sharp edges, petiolules of lateral leaflets 2.5-3.5
cm long....................................................................................................3. D. expansa
Petiole and petiolules not so.........................................................................................8
8.
Midrib below sharply keeled; petiole strongly grooved or flattened above.......................

..............................................................................................................6. D. longifolia
Midrib below not keeled; petiole rounded or flattened above........................................9
9.
Reticulations fine and more distinct on the upper than on the lower leaflet surface. Inner
bark reddish; cut bark and twigs with white sap..............................9. D. rostrata
Reticulations more distinct on the lower surface. Inner bark yellowish; cut bark and
twigs without white sap..............................................................................................10
10. Leaflets usually of 2 pairs only. Trees with stilt-roots........................7. D. macrocarpa

Leaflets more than 2 pairs. Trees without stilt-roots...................................................11
11. Lateral veins sunken above; intercostal veins raised below. Petiolule strongly swollen at
both ends..............................................................................................11. D. rugosa
Lateral veins not sunken above; intercostal veins faintly visible below. Petiolule not
prominently swollen..............................................................................4. D. incurvata
1. Dacryodes costata (A.W. Benn.) H.J. Lam

(Latin, costatus = ribbed; probably the prominent veins on the lower leaflet surface)
Kalkman l.c. (1954) 508; Leenhouts l.c. (1956) 222; Burgess l.c. 61; Kochummen l.c. (1972) 140;
Anderson l.c. 156; Whitmore, Tantra & Sutisna l.c. 36. Basionym: Santiria costata A.W. Benn. in
Hooker f. l.c. 537. Type: Maingay 313/3295, Malacca (holotype K; isotype CAL). Synonym:
Canarium costatum (A.W. Benn.) Ridl. l.c. (1930) 82.
Medium-sized to large tree to 45 m tall, 45 cm diameter; buttresses short. Bark grey-brown,

smooth to flaky; inner bark yellow-brown, with droplets of white sap. Sapwood whitish.
Ultimate leafy twigs to 0.5 cm thick, rusty-brown velvety hairy when young. Leaves with
13 pairs of leaflets; rachis powdery rusty hairy; petiole hairy, flattened above; petiolules of
lateral leaflets 515 mm long, powdery hairy to glabrous, slightly swollen at both ends;
blade sparsely hairy on midrib and veins below, elliptic to oblong, 6 15 x 2.56 cm; base
cuneate, often unequal, apex pointed, tip 12 cm long; midrib raised above; lateral veins
1014 pairs, prominently raised below, faint above, looping and joining near margin;
intercostal veins reticulate, distinct below, faint above. Inflorescences terminal or from
upper leaf axils, axes densely hairy. Flowers pubescent; petals whitish, glabrous; stamen
filaments free from the disc; disc annular to 6-lobed. Fruits ellipsoid or ovoid, 1.72.2 x
11.2 cm; stalk c. 1 cm long.
66
Distribution. Sumatra, Peninsular Malaysia, Borneo, and the Philippines. In Sabah

reported from Ranau, Sandakan and Tawau; in Sarawak, known only from the Lambir
National Park and Semengoh FR. Also known in Kalimantan.
Ecology. Widely distributed from lowland to hill forests to 540 m, mainly on ridges and
hillsides. Flowering in May and September, and fruiting in July, October and November.
This species is very similar to D. rugosa from which it can be distinguished by the hairy
petioles, and the less swollen leaflet stalks. The flowers of both species are quite distinct, in
D. costata the disc is annular to 6-lobed and the stamen filaments free from disc, while in D.
rugosa the disc is cup-shaped and stamen filaments adnate to disc.
2. Dacryodes elmeri H.J. Lam

(A.D.E. Elmer, 18701942; plant collector with the Bureau of Science, Manila, the
Philippines)
Ann. Jard. Bot. Btzg. 42 (1932) 203, Bull. Jard. Bot. Btzg. 3, 12 (1932) 344; Kalkman l.c.
(1954) 521; Leenhouts l.c. (1956) 225. Synonym: Hemisantria ?n. sp. H.J. Lam in Merrill
l.c. (1929) 199. Type: Elmer 21573, British North Borneo, Tawau (holotype L; isotypes A,
BKH, BM).
Tree, to 75 cm in diameter. Ultimate leafy twigs c. 1 cm thick, densely minutely villous; pith
with many scattered vascular strands. Terminal bud c. 1.5 cm long, densely pubescent.
Leaves with 34 pairs of leaflets; rachis strongly flattened at base, hairy at first becoming
glabrous; blade oblong, 1222 x 58.5 cm, chartaceous; base oblique-cuneate, on one half
rounded, apex pointed; lateral veins 1318 pairs, prominent beneath. Inflorescence and
flower unknown. Fruits ovoid, 44.7 x 2.25 cm, with rounded apex and base.
Distribution. Endemic to Borneo. Very uncommon, known from the type collection (Sabah)
and from Brunei.
Ecology. Lowland forest.
3. Dacryodes expansa (Ridl.) H.J. Lam

(Latin, expansus = spread out, diffuse; the thin, spread out petals)
Kalkman l.c. (1954) 510; Leenhouts l.c. (1956) 228, l.c. (1972) 919, l.c. (1976) 821. Basionym:
Canarium expansum Ridl. l.c. (1930) 83. Type: Haviland 2271, Sarawak, Kuching (K).
Small tree. Buds reddish brown, hairy. Leaves with 4 pairs of leaflets; adaxial side of
petiole flattened with sharp edges; petiolules of lateral leaflets 2.53.5 cm long, strongly
swollen at both ends; blade oblong to oblong-lanceolate, 1723 x 67.5 cm, brownish when
dried; base cuneate, apex pointed; midrib raised above; lateral veins 1012 pairs, prominent
below; intercostal veins faint. Inflorescences (male) probably lateral on axillary shoots, to
24 cm long, glabrous; pedicels 37 mm long, articulated. Flowers (male) glabrous; calyxlobes deltoid; petals very thin, spreading; stamens free from disc; disc annular, thick; ovary
in male flowers reduced. Infructescence and fruit unknown.
67
Distribution. Endemic to Borneo. Uncommon, known from the type collection only and
from Brunei.
4. Dacryodes incurvata (Engl.) H.J. Lam

(Latin, incurvatus = bending inward; the leaflet margin)
Ann. Jard. Bot. Btzg. 42 (1932) 204, Bull. Jard. Bot. Btzg. 3, 12 (1932) 362; Kalkman l.c. (1954) 506;
Leenhouts l.c. (1956) 224; Burgess l.c. 61; Kochummen l.c. (1972) 140; Anderson l.c. 156; Whitmore,
Tantra & Sutisna l.c. 36. Basionym: Canarium incurvatum Engl. in DC. l.c. 138. Type: Beccari PB
2937, Sarawak (holotype P; isotype FI). Synonyms: Santiria nitida Merr., Publ. Govt. Lab. Philip. 35
(1906) 29; Canarium nitens Merr., Philip. J. Sc. 10 (1915) Bot. 24; Hemisantiria nitida H.J. Lam in
Merrill l.c. (1929) 118; Canarium angulatum Ridl., Kew Bull. (1931) 493; Dacryodes angulata (Ridl.)
H.J. Lam, Ann. Jard. Bot. Btzg. 42 (1932) 204.
Medium-sized to large tree to 35 m tall, 30 cm diameter; buttresses steep to 2 m high. Bark

grey-brown, smooth, with horizontal rings; inner bark pink with droplets of white sap.
Sapwood pale white. Twigs dark brown. Leaves with 14 pairs of leaflets; petiole strongly
flattened at base; petiolules of lateral leaflets 12 cm long, swollen at both ends; blade
thinly to thickly leathery, shiny above, elliptic, oblong or lanceolate, 616 x 29 cm; base
rounded, subcordate or cuneate, apex pointed; midrib raised or flattened above; lateral veins
1016 pairs, raised on both surfaces; intercostal veins reticulate, faintly visible on both
surfaces. Flowers hairy, usually in terminal panicles. Fruits ovoid or ellipsoid, 2.53 x
1.72 cm, ripening to yellow and then purplish.
Distribution. Sumatra, Peninsular Malaysia, Borneo, and the Philippines. Common in Sabah,
recorded from Beaufort, Sipitang and Tawau. In Sarawak recorded from Bintulu, Lambir
National Park, Serian and Kuching. Also found in Brunei and Kalimantan.
Ecology. Mixed dipterocarp forests to 860 m; uncommon in mixed peat swamp forest.
Flowering in April, June and September, and fruiting in March and JuneNovember.
5. Dacryodes laxa (A.W. Benn.) H.J. Lam

(Latin, laxus = loose or distant; the inflorescence)
Leenhouts l.c. (1956) 224; Kochummen l.c. (1972) 141; Cockburn l.c. 45; Anderson l.c. 156;
Whitmore, Tantra & Sutisna l.c. 37. Basionym: Canarium laxum A.W. Benn. in Hooker f. l.c. 535.
Type: Maingay 366, Malacca (holotype K; isotypes CAL, L). Synonyms: Canarium fragile Engl. in
DC. l.c. 138; Santiria laxa (A.W. Benn.) King, l.c. 254.
Small to medium-sized tree to 30 m tall, 30 cm diameter. Bark grey, smooth to scaly; inner
bark pale yellow. Sapwood pale white. Twigs with long rough hairs when young. Leaves
with up to 5 pairs of leaflets; rachis rough-hairy; petiolules of lateral leaflets 310 mm long,
hairy; blade drying greenish, oblong, lanceolate or oblanceolate, 8.534 x 4.59 cm; base
cuneate, apex pointed, tip to 1.5 cm long; midrib raised above; lateral veins 930 pairs,
raised on both surfaces, curving and joining near margin; intercostal veins reticulate, raised
below, sunken above; petiole strongly swollen at base. Flowers in branched, long (c. 80 cm)
terminal panicles; petals glabrous. Fruits pink ripening to blue, oblong or ovoid, 2.54.5 x
12 cm, apex pointed; stalk 22.5 cm long, slender.
68
Distribution. Sumatra, Peninsular Malaysia, and Borneo. Widely distributed in Sabah. In

Sarawak reported only from Bako National Park, Semengoh Arboretum and Lundu. Also
known in Brunei.
Ecology. Common in mixed dipterocarp forests on yellow sandy clay soils, to 700 m.
Flowering from April to June, and fruiting from June to November.
6. Dacryodes longifolia (King) H.J. Lam

(Latin, longus = long, folium = leaf)
Leenhouts l.c. (1956) 228, l.c. (1972) 919; Kochummen l.c. (1972) 141. Basionym: Santiria
longifolia King l.c. 258. Type: King's collectors 6838, Perak (holotype K; isotype L). Synonyms:
Curtisiana penangensis Ridl. l.c. (1920) 180; Dacryodes longifolia var. penangensis (Ridl.) H.J. Lam,
Ann. Jard. Bot. Btzg. 42 (1932) 202.
Medium-sized to large tree to 40 m tall. Bark dark grey, smooth; inner bark orange-yellow.
Twigs brown, with scattered white lenticels. Leaves with 26 pairs of leaflets; petiole
strongly grooved or flattened above; petiolules of lateral leaflets 12 cm long, strongly
swollen at both ends; blade oblong to lanceolate, 6.518 x 2.54.5 cm; base usually
unequal, broadly cuneate to rounded, apex pointed; midrib raised above, sharply keeled
below; lateral veins 712 pairs, distinct below, faint above; intercostal veins reticulate,
faintly visible on both surfaces. Flowers (male) in axillary panicles, glabrous. Fruits ellipsoid,
2.53.5 x 1.52.5 cm, much wrinkled on drying; stalk c. 7 mm long.
Distribution. Sumatra, Peninsular Malaysia, Borneo and the Philippines. Common and
widely distributed in Sabah; less common in Sarawak.
Ecology. Lowland to submontane forests to 1500 m.
7. Dacryodes macrocarpa (King) H.J. Lam

(Greek, makros = large, karpos = fruit)
Kalkman l.c. (1954) 514; Leenhouts l.c. (1956) 228, l.c. (1972) 919, l.c. (1976) 820; Burgess l.c. 61;
69
70
Kochummen l.c. (1972) 142; Anderson l.c. 156; Whitmore, Tantra & Sutisna l.c. 37. Basionym:
Santiria macrocarpa King l.c. 256. Type: King's collectors 7298, Perak (holotype K; isotypes CAL,
SING).
Medium-sized to tall tree; stilt-roots and buttresses present. Bark reddish brown, cracking.
Twigs dark brown to blackish when dry. Leaves with 12 pairs of leaflets; petiolules of
lateral leaflets 0.72 cm long, swollen at both ends; blade elliptic, obovate or ovate, 612 x
36 cm; base often unequal, rounded to broadly cuneate, apex blunt, rounded or shortly
pointed; midrib flattened above; lateral veins 69 pairs, visible on both surfaces; intercostal
veins scalariform-reticulate, faintly visible on both surfaces. Flowers glabrous, in axillary
panicles. Fruits ovoid or ellipsoid, 2.54 x 2.5 cm.
Key to varieties
1.
Leaflets widest around the middle, equal-sided at base; lateral veins at right angle to
midrib...........................................................................................................................
var. patentinervia Leenh.
l.c (1976) 821. Type: Sinclair & Kadim 10492, Brunei, Bt. Labi FR (holotype L; isotypes K,
SAR, SING).
Endemic to Borneo. In Sabah uncommon, known only from Bt. Hampuan, Ranau
district (SAN 25331). In Sarawak scattered throught 3rd, 4th and 5th Div. in the
lowland to submontane forest to 1500m. Also in Brunei. Vernacular names:
Sarawakseladah (Iban). Bruneisabal (Iban), sibut (Tutong, Dusun).
Leaflets widest in the lower half, oblique at base; lateral veins at acute angle with
midrib..........................................................................................................................2
2.
Twigs blackish when dry. Midrib and veins not prominent beneath...............................
var. macrocarpa
Synonym: D. macrocarpa var. genuina H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 343.
Sumatra, Peninsular Malaysia, and Borneo. Uncommon in Sabah, more common in

Sarawak. Also in Brunei and Kalimantan. Vernacular names: Sabahasam-asam
(Dusun). Sarawakicerawas burung, keruas (Malay). Bruneikedondong, pasohpasoh (Malay).
Twigs and rachis scaly to powdery hairy and brown when dry. Midrib and veins
prominent below............................................................................................................
var. kostermansii (Kalkman) Kalkman
in Leenhouts l.c. (1956) 228. Synonym: D. kostermansii Kalkman l.c. (1954) 515. Type:
Kostermans 6725, SE Borneo, Loa Djanan, W Samarinda (holotype L; isotypes BO).
Endemic to Borneo. In Sabah known by a single collection (SAN 25572) from the
Silam FR, Lahad Datu district; not recorded in Sarawak. Also found in Kalimantan.
Ecology. Coastal and mixed swamp forests. Flowering in September and fruiting in
MarchApril and November.
71
8. Dacryodes nervosa (H.J. Lam) Leenh.

(Latin, nervosus = prominently nerved; the leaves)
Blumea 12 (1964) 19, l.c. (1972) 919. Basionym: Santiria nervosa H.J. Lam, Ann Jard. Bot. Btzg. 42
(1932) 206, Bull. Jard. Bot. Btzg. 12, 3 (1932) 387. Type: Grashoff 960, Sumatra (L).
Medium-sized tree to 30 m tall, 35 cm diameter; buttresses to 1.5 m tall. Bark smooth to

scaly, dark brown. Ultimate leafy twigs to 0.5 cm thick. Terminal bud 0.51 cm long.
Leaves with 14 pairs of leaflets; blade oblong to ovate, 5.517 x 37.5 cm, upper surface
greenish when dried, lower surface pubescent to glabrous; base broadly cuneate, asymmetric, apex pointed; midrib raised above; lateral veins 1015 pairs, prominent below;
intercostal veins scalariform, raised below; petiolule 811 mm long, hairy. Inflorescences
axillary. Flowers sessile or shortly stalked. Fruits pink when fresh, ellipsoid, 1.61.7 x 1.1
cm.
Distribution. Sumatra, Peninsular Malaysia, and Borneo. Uncommon, in Sarawak so far
recorded only from Bako National Park, Bintulu and Sadong (S. 13389, S.16228, S.18192,
S. 37488); not yet found in Sabah. Also occurs in Kalimantan.
9. Dacryodes rostrata (Blume) H.J. Lam
Fig. 2.
(Latin, rostratus = with a beak or narrowed tip; the leaves)

Kalkman l.c. (1954) 519; Leenhouts l.c. (1956) 225; Burgess l.c. 161; Kochummen l.c. (1972) 143;
Cockburn l.c. 45; Anderson l.c. 156; Whitmore, Tantra & Sutisna l.c. 37. Basionym: Santiria rostrata
Blume l.c. (1850) 213. Type: Korthals, s.n. (= Leiden no. 898.321228), Borneo (L). Synonym:
Hemisantiria rostrata (Blume) H.J. Lam in Merrill l.c. (1929) 119.
Small to very large tree to 45 m tall, 200 cm diameter. Bark dark grey, smooth to scaly or
dippled; inner bark reddish or pinkish with droplets of white sap. Sapwood whitish. Twigs
thin to very thick (0.42.5 cm), dark brown, lenticellate. Leaves with 28 pairs of leaflets;
petiolules of lateral leaflets 12 cm long, swollen at both ends; blade thinly to thickly
leathery, glabrous or (sometimes) with tiny inconspicuous hairs below, ovate to oblong, 8
30 x 3.511 cm; base strongly unequal, apex pointed, tip c. 2 cm long; midrib raised above;
lateral veins 515 pairs, raised on both surfaces; intercostal veins reticulate, faint to distinct
on both surfaces or sometimes more distinct above than below; petioles strongly flattened at
base in those with thick twigs. Flowers densely hairy, in axillary panicles 535 cm long.
Fruits ovoid to oblong, 23.5 x 12 cm, often strongly wrinkled on drying, ripening to blue,
with white sap.
Key to forms
Ultimate leafy twigs 1.52.5 cm thick. Leaflets at least 10 cm long, drying greenish grey;
reticulations more distinct on the upper than on lower surface..............................................
72
forma cuspidata (Blume) H.J. Lam

Bull. Jard. Bot. Btzg. 3, 12 (1932) 351. Basionym: Dracontomelon cuspidatum Blume l.c. (1850).
Type: Korthals, s.n. (= Leiden no. 897. 363270), Borneo (L). Synonym: Canarium minahassae
Koord. l.c. 96; Santria samarensis Merr. l.c. (1915) 315; C. crassifolium Merr. l.c. (1915) 274; C.
cuspidatum (Blume) Merr. l.c. (1921) 316; C. reticulatum Ridl. l.c. (1930) 83.
Indo-China, Sumatra, Borneo, Philippines, and N Celebes. In Sabah, known from a few
collections from Labuan FR (FDBNB 44236), Bt. Batangan, Sipitang district (SAN
16612), and Nabutan, Ranau district (SAN 100230). In Sarawak recorded from Bt.
Rawan (S. 45575) and Semengoh Arboretum (S. 32406) in 1st Div., Ulu Balleh, Kapit,
3rd Div. (S. 29100); and Ulu Dapoi, Marudi (S. 23056 & S. 23474), Kebulu Protected
Forest (S. 49126), and Dulit Range (S. 46672) in the 4th Div. Vernacular names:
Sabahkedondong, salong banggi (Malay). Sarawakseladah (Iban).
Ultimate leafy twigs to 0.5 cm thick. Leaflets smaller, drying reddish brown; reticulations
not so...................................................................................................................................
forma rostrata
Synonyms: D. rostrata f. genuina H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 351; Santiria
montana Blume l.c. (1850) 212; C. montanum (Blume) Korth. ex Miq. l.c. (1859) 649: C.
kedondon A.W. Benn. in Hooker f. l.c. 535.
Sumatra, Peninsular Malaysia, and Borneo. Common and widely distributed in Sabah
and Sarawak. Vernacular names: Sabahkedondong, kambayau (Dusun).
Ecology. Very common in mixed dipterocarp forest to 800 m. Flowering in February
September, and fruiting in April and JuneDecember.
10. Dacryodes rubiginosa (A.W. Benn.) H.J. Lam

(Latin, rubiginosus = of rusty colour; the tomentum)
Kalkman l.c. (1954) 521; Leenhouts l.c. (1956) 225; Kochummen l.c. (1972) 143; Whitmore, Tantra &
Sutisna l.c. 37. Basionym: Canarium rubiginosum A.W. Benn. in Hooker f. l.c. 535, non C.
rubiginosum Miq. l.c. (1859) 651 (= Santiria rubiginosa Blume). Type: Maingay 309, Malacca
(holotype K; isotypes CAL, L).
Medium-sized tree to 20 m tall, 20 cm diameter. Ultimate leafy twigs to 0.5 cm thick,

yellowish brown velvety hairy when young. Leaves with 23 pairs of leaflets; rachis
appressed hairy; petiolules of lateral leaflets 0.51 cm long, swollen at both ends; blade
sparsely hairy below, elliptic or oblong, 9.519.5 x 2.56.5 cm; base cuneate, unequal, apex
pointed, tip c. 1.5 cm long; midrib raised above; lateral veins 812 pairs, prominently raised
below, faintly raised above; intercostal veins scalariform-reticulate, visible below. Flowers
(male) tomentose, in terminal inflorescences, axes densely yellowish brown hairy. Fruits
ellipsoid, 22.5 x 1 cm.
Distribution. Peninsular Malaysia and Borneo. In Sabah scattered. In Sarawak uncommon,
known only from two collections (S. 43194 and S. 43377) from Samunsam Wildlife Sanctuary
and Ulu Sg. Semawat respectively. Also found in Kalimantan.
73
11. Dacryodes rugosa (Blume) H.J. Lam

(Latin, rugosus = wrinkled; the bullate leaflets)
Kalkman l.c. (1954) 505; Leenhouts l.c. (1956) 221; Backer & Bakhuizen f. l.c. (1972) 114; Burgess
l.c. 62; Kochummen l.c. 144; Cockburn l.c. 47; Anderson l.c. 157; Whitmore, Tantra & Sutisna l.c. 37.
Basionym: Santiria rugosa Blume l.c. 212. Type: Korthals, s.n. (= Leiden no. 898.321232), Java
(L). Synonyms: Canarium rugosum (Blume) Miq. l.c. (1859) 649 (including var. sumatranum);
Hemisantiria rugosa (Blume) H.J. Lam in Merrill l.c. (1929) 119.
Small to medium-sized tree, rarely to 30 m tall and 40 cm diameter. Bark grey-white,

smooth to scaly; inner bark yellowish. Sapwood pale. Ultimate leafy twigs 0.30.5 cm
thick. Leaves with 14 pairs of leaflets; rachis flattened above; petiole glabrous; petiolules
of leaflets strongly swollen at both ends; blade glabrous or appressed pubescent below or
with erect hairs on midrib and veins below (var. virgata), elliptic, ovate to oblonglanceolate, 622 x 2.511 cm, sometimes bullate; base cuneate, apex pointed with long tip;
midrib raised above; lateral veins 712 pairs, raised below, flattened above (sunken in var.
rugosa), often curving and joining near margin; intercostal veins reticulate, raised below,
faint or sunken above. Flowers in axillary panicles, glabrous (var. virgata) or hairy; stamen
filaments adnate to the disc; disc cup-shaped. Fruits ovoid, slightly oblique, 1.52.5 x 1
1.5 cm.
Key to varieties
Lateral veins sunken on upper leaflet surface. Leaflets glabrous or appressed-hairy on lower
surface. Flowers densely hairy..............................................................................................
var. rugosa
Synonyms: Dacryodes rugosa var. genuina H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 347;
Santiria fasciculata A.W. Benn. in Hooker f. l.c. 539.
Sumatra, Peninsular Malaysia, Java, and Borneo. Widespread in Sabah and Sarawak in
mixed dipterocarp forest on yellow sandy clay soils. Also in Kalimantan.
Lateral veins not sunken on upper leaflet surface. Leaflets with scattered erect hairs on
midrib and veins on lower side. Flowers glabrous.................................................................
var. virgata (Blume) H.J. Lam
Bull. Jard. Bot. Btzg. 3, 12 (1932) 348. Basionym: Santiria virgata Blume l.c. (1850) 213.
Synonym: Canarium virgatum (Blume) Miq. l.c. (1959) 650. Type: Korthals, s.n., W Borneo, G.
Pamaton (holotype L; isotype BO).
Widely distributed in Sabah and Sarawak; also in Kalimantan.

Ecology. Lowland to hill mixed dipterocarp forests to 900 m. Flowering in AprilNovember,
and fruiting in MarchDecember.
3. GARUGA Roxb.
(an Indo-Malayan plant name, origin uncertain)
Pl. Corom. 3 (1811) 5; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 325; Kalkman l.c. (1954) 459;
Leenhouts l.c. (1956) 215; Kochummen l.c. (1972) 144; Whitmore, Tantra & Sutisna l.c. 37.
Deciduous trees. Pith of branches and petioles without vascular strands. Leaves with
stipules, stipellae (stipule-like outgrowths at the base of leaflets) often present; margin of
leaflets toothed. Inflorescence an axillary panicle. Flowers bisexual, 5-merous; receptacle
74
concave, globose or cup-like; sepals free; stamens 10; disc adnate to the receptacle, glabrous
with 10 lobes between the stamens; ovary 5-celled. Fruit drupaceous; pericarp fleshy;
pyrenes 15, furrowed, bony. Cotyledons twisted and folded.
Distribution. 4 species; continental SE Asia, with a variety of one species in Malesia,
Melanesia, N Australia, and the Solomon Islands.
Garuga floribunda Decne. var. floribunda
Fig. 3.
(Latin, floribundus = with abundant flowers)

Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 477; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 326;
Kalkman l.c. (1954) 463; Leenhouts l.c. (1956) 215; Kochummen l.c. (1972) 144, Whitmore, Tantra &
Sutisna l.c. 37. Type: sine coll., s.n., Timor (holotype G; isotype NY). Synonyms: Guaiacum abilo
Blanco, Fl. Filip. (1837) 364; Garuga abilo (Blanco) Merr., Publ. Govt. Lab. Philip. 35 (1905) 73;
Garuga littoralis Merr. l.c. (1915) 27; Garuga clarkei Merr. l.c. (1915) 29.
Small to medium-sized tree to 12 m tall, 10 cm diameter. Bark grey, smooth. Twigs covered
with many leaf-scars, minutely tomentose at tips. Stipules inserted at base of petioles,
caducous. Leaves crowded at ends of twigs, leaflets 410(15) pairs; blade subsessile,
elliptic to oblong or lanceolate, 519 x 25.5 cm; base oblique, cordate or rounded, margin
toothed, apex pointed; midrib raised above; lateral veins 1020 pairs, distinct on both
surfaces; intercostal veins reticulate, visible on both surfaces. Flowers with cup-shaped
receptacle; petals tomentose inside. Fruits obliquely pear-shaped, 59 x 512 mm.
Distribution. Peninsular Malaysia, Java, Borneo, Philippines, Celebes, Lesser Sunda Islands,
Moluccas, New Guinea, Melanesia, and W Australia. In Borneo uncommon, known only from
some islands (Gaya and Sipadan) in Sabah.
Ecology. Coastal forests at low altitude.
A second variety, var. gamblei, is known from E India, Sikhim, Bangladesh, W China and
Hainan.
4. HAPLOLOBUS H.J. Lam

(Greek, haplos = single, lobus = lobe; the single-lobed seed)
Ann. Jard. Bot. Btzg. 42 (1932) 25, Bull. Jard. Bot. Btzg. 3, 12 (1932) 404, Blumea 9 (1958) 237;
Husson & H.J. Lam, Blumea 7 (1953) 413; Leenhouts l.c. (1956) 239, Blumea 20 (1972) 283 & 311;
Whitmore, Tantra & Sutisna l.c. 37; Kochummen, Sandakania 5 (1994) 75.
Small to medium-sized, dioecious trees. Bark greyish, scaly. Pith of branches and petioles
mostly without vascular strands. Leaves without stipules, imparipinnate; leaflets entire.
Inflorescences paniculate, axillary or borne on leafless twigs or stems, sometimes with
vegetative terminal bud. Flowers 3-merous, functionally unisexual, organs of the other sex
always present and sometimes only little reduced in size; sepals united; petals free with
75
inflexed tip; stamens 6 (occasionally only 3), inserted outside the annular disc or filaments
slightly connate to it at base; ovary 3-celled, each cell with 2 collateral, pendulous, axile
ovules; stigma 3-lobed to subglobular. Fruits ovoid to subglobular, calyx persistent,
spreading, stigma terminal; pericarp dry, thin, smooth. Seed mostly 1 in each pyrene;
cotyledons plano-convex, thick, entire; germination hypogeal.
Distribution. 19 species; from Borneo and Celebes through the Moluccas and New Guinea
to Western Polynesia. 6 species in Sabah and Sarawak.
Ecology. From sea-level to 1800 m.
Taxonomy. Species of this genus are difficult to distinguish from some Santiria and
Dacryodes species in the sterile state or even with flowers. Generally in Haplolobus the
twigs are lenticellate, occasionally some species have galls or their holes on the leaf-blade,
and often the inflorescences are short and usually terminate in a vegetative bud.
Key to Haplolobus species

1.
Leaflets hairy below.....................................................................................................2

Leaflets glabrous below................................................................................................3
2.
Leaflets densely hairy below; lateral veins 2530 pairs; petiole thin, lower part
distinctly grooved....................................................................................1. H. beccarii
Leaflets sparsely hairy below; lateral veins 79 pairs; petiole thicker, not grooved.........
.........................................................................................................2. H. bintuluensis
3.
Petiole channelled above, with incurved edges, petiolules with horizontal cracks,
swollen at both ends............................................................................5. H. leenhoutsii
Not this combination of characters...............................................................................4
4.
Leaflets drying greenish brown above, reddish brown below. Young twigs strongly
angled..................................................................................................4. H. kapitensis
Leaflets drying the same colour on both surfaces. Young twigs not angled...................5
5.
Leaflets drying dark brown, strongly asymmetric; midrib not in the centre, sharply
keeled below....................................................................................3. H. inaequifolius
Leaflets drying yellowish; midrib not so..........................................6. H. sarawakanus
1. Haplolobus beccarii Husson

(Odoardo Beccari, Italian explorer and botanist, 18431920)
in Husson & H.J. Lam l.c. 431; Leenhouts l.c. (1956) 243, l.c. (1972) 292; H.J. Lam l.c. (1958) 248;
Whitmore, Tantra Sutisna l.c. 37. Type: Beccari PB 1803, Sarawak (FI).
76
77
78
Tree. Twigs stout to 2.5 cm diameter, lenticellate. Leaves with 6 pairs of leaflets; pith of
petioles with 1520 vascular strands; rachis to 23 cm long; blade lanceolate-oblong, 2136
x 7.510 cm, densely hairy on the under surface; base cuneate, apex pointed; lateral veins
2530 pairs, arching near margin; petiole flattened on the upper side, and distinctly striate
on the lower side. Inflorescences and flowers unknown. Infructescences borne on leafless
twigs or stems. Fruits oblong, with pointed tip, 1315 x 12 mm.
Distribution. Endemic to Sarawak. Uncommon, known only from the type specimen from
Mt. Matang.
2. Haplolobus bintuluensis Kochummen

(of Bintulu in Sarawak)
l.c. (1994) 75. Type: Sibat S. 24562, Sarawak (holotype SAR; isotypes A, K, L, SAN, SING).
Treelet to 3 m tall, 5 cm diameter. Twigs c. 7 mm thick, greyish, with white lenticels.

Terminal bud oblong, c. 10 x 3 mm, rusty hairy. Leaves: rachis c. 23 cm long, slightly
swollen at base, dark brown-hairy with 23 pairs of leaflets; petiolules of lateral leaflets 8
15 mm long, dark brown hairy, swollen at both ends, petiolules of terminal leaflets 3.54.5
cm long; blade thinly leathery, glabrous above except midrib, dark brown hairy on the
midrib and veins below, 10.517 x 5.57 cm; base cuneate, apex long-pointed, tip narrow,
12 cm long; midrib flattened above; lateral veins 79 pairs, distinct below, faint above,
curving and joining near margin; intercostal veins reticulate, faintly visible below.
Inflorescences and flowers unknown. Infructescence c. 2 cm long. Fruits oblong, reddish
when fresh, yellowish brown on drying, 1620 x 1112 mm, with pointed tip.
Distribution. Endemic to Sarawak. Uncommon, known from the type collection only, from
the Nyabau Catchment area, Bintulu.
Ecology. Mixed dipterocarp forest on yellow-red sandy humult ultisol, at about 100 m.
3. Haplolobus inaequifolius Kochummen

(Latin, inaequalis = unequal, folium = leaf; the asymmetric leaflets)
l.c. (1994) 78. Type: Yii S. 40756, Sarawak (holotype KEP; isotypes K, L, MO, SAN, SAR).
Small tree to 15 m tall, 10 cm diameter. Twigs brownish, c. 5 mm thick with whitish lenticels.
Terminal bud oblong, c. 12 mm long, rusty hairy. Leaves glabrous, with 2 pairs of leaflets;
rachis grey, not swollen at base; petiolules of lateral leaflets 815 mm long, of terminal
leaflet c. 4.5 cm long, the ends black on drying; blade greenish brown on drying, leathery,
with 1-mm-wide circular galls, elliptic to ovate or oblong, 7.515.5 x 2.56 cm, strongly
asymmetric; base broadly cuneate, apex pointed; midrib flattened above, sharply keeled
below; lateral veins 10 pairs, faint on both surfaces; intercostal veins finely reticulate,
visible on both surfaces. Inflorescence (male) glabrous, axillary, 48 cm long, often
terminating in a vegetative shoot. Flowers (male): calyx-lobes triangular with pointed tip;
petals oblong; stamens 6 with short stout filaments; disc with wavy margin; rudimentary
ovary with 3 distinct stigma. Fruits unknown.
Distribution. Endemic to Borneo. Uncommon, known only by the type specimen from the
Sabal Forest Reserve in Sarawak.
Ecology. Hill forest at about 360 m.
79
4. Haplolobus kapitensis Kochummen
Fig. 4BD.
(of Kapit, Sarawak)

l.c. (1994) 78. Type: Wright S. 23966, Sarawak, Kapit (holotype KEP; isotypes A, BO, K, L, SAN,
SAR, SING).
Emergent tree to 40 m tall, 80 cm diameter. Twigs greyish white or grey-brown, c. 1 cm

thick, lenticellate, youngest strongly angled. Leaves glabrous, with 23 pairs of leaflets;
rachis c. 26 cm long, channelled above near base; petiolules of lateral leaflets 1.52 cm
long, of terminal leaflets c. 4.5 cm long, strongly swollen at both ends; blade thickly
leathery, drying to reddish brown below and greenish brown above, oblong to lanceolate,
11.520 x 4.59 cm; base asymmetric, almost rounded to broadly cuneate, apex pointed;
midrib raised above; lateral veins 912 pairs, distinctly looping near margin, distinct on both
surfaces; intercostal veins reticulate, more distinct above than below. Inflorescence (male)
axillary, to 21 cm long, on leafless vegetative shoot. Flowers (male): calyx cup-shaped with
truncate apex; petals yellow, oblong; stamens 3 or 6, with short filaments; disc cup-shaped
with wavy margin; pistil rudimentary. Fruits unknown.
Distribution. Endemic to Borneo. Uncommon, known by four collections, S. 23966 and S.
29162 from Kapit, Sarawak, and SAN 22391 and SAN 62657 from Ranau in Sabah.
Ecology. Lowland and hill forests to 600 m.
5. Haplolobus leenhoutsii Kochummen
Fig. 4A.
(P.W. Leenhouts, botanist at Rijksherbarium, Leiden, the Netherlands)

l.c. (1994) 81. Type: Othman et al. S. 41304, Sarawak, Kapit (holotype SAR; isotypes K, KEP, L,
MO, SAN).
Small tree to 20 m tall, 20 cm diameter. Bark brownish green, scaly. Twigs brown, c. 1 cm
thick. Leaves with 3 pairs of leaflets; rachis c. 23 cm long; petioles channelled with incurled
edges above, with many shallow grooves on the remaining part; petiolules of lateral and
terminal leaflets c. 2 cm long, strongly swollen at both ends and with horizontal cracks;
blade drying to greenish brown, glabrous, leathery, lanceolate to elliptic, 1423 x 4.57.5
cm, with many circular (3-mm-diameter) domatia holes; base rounded or broadly cuneate,
apex pointed; midrib raised above, channelled below; lateral veins 1417 pairs, distinct
below, faint above; intercostal veins reticulate, visible on both surfaces. Inflorescences and
flowers unknown. Infructescences axillary, c. 15.5 cm long, with many branches. Fruits
(immature) red when fresh, black on drying, oblong, c. 1 cm long, with prominent apical
stigma, surface slightly rugose, with remains of floral parts at base, stalk c. 1 cm long.
Distribution. Endemic to Borneo. Uncommon, known only from the type specimen from
Ulu Balleh, Kapit, Sarawak.
Ecology. Mixed dipterocarp forest at about 500 m, on ridge.
80
6. Haplolobus sarawakanus Kochummen

(of Sarawak)
l.c. (1994) 81. Type: Othman et al. S. 41339, Sarawak (holotype SAR; isotypes K, KEP, L, SAN,
MO).
Small tree to 11 m tall, 15 cm diameter. Bark greyish brown, scaly. Twigs c. 5 mm thick,
pale brown, shallowly grooved, with pale lenticels. Leaves glabrous, with 34 pairs of
leaflets; rachis c. 25 cm long, slightly swollen at base; petiolules of lateral leaflets c. 1 cm
long, of terminal leaflet c. 3 cm long, the ends swollen and whitish or blackish; blade thinly
leathery, drying to yellowish brown, elliptic or oblong, 1215 x 3.55 cm; base cuneate,
slightly asymmetric, apex pointed; midrib flattened above; lateral veins 89 pairs, distinct
below, faint above, curving and joining near margin; intercostal veins reticulate, very faint
on both surfaces. Inflorescences and flowers unknown. Infructescences c. 4 cm long.
Fruits (immature) c. 8 x 5 mm, calyx funnel-shaped.
Distribution. Endemic to Borneo. Uncommon, known only by the type from Ulu Balleh,
Kapit, Sarawak. A collection from Sabah, SAN 97191, probably belongs to this species but
better material is needed to confirm this.
Ecology. Mixed dipterocarp forest at about 500 m, on ridge.
5. PROTIUM Burm. f., nom. cons.

(Greek, proteion = first-class; probably
referring to the hardness and strength of the wood)
Fl. Ind. (1768) 88; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 318; Leenhouts l.c. (1952) 154, l.c.
(1956) 213; Backer & Bakhuizen f. l.c. 113.
Trees or shrubs. Pith of branchlets and petioles withouts vascular strands. Leaves without
stipules, tips of the leaflets usually distinctly mucronulate. Inflorescences paniculate, axillary,
rarely pseudo-terminal. Flowers 45-merous, structurally not always completely unisexual;
sepals united; petals valvate with inflexed margins, glabrous; stamens double the number of
petals, free; disc intrastaminal, annular, truncate to undulate, glabrous; ovary 35-celled,
stigma subsessile. Fruits drupaceous, pericarp fleshy, calyx persistent, not enlarged, with
reflexed lobes. Cotyledons plicate, lobed to palmatifid.
81
82
Distribution. About 85 species; mainly in the American tropics, Madagascar, Mascarenes,

India and Malesia. Only one species has been recorded in Sabah; not yet found in Sarawak.
Protium connarifolium (Perkins) Merr.
Fig. 5.
(with the leaflets resembling those of Connarus)

l.c. (1915) 30; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 320; Leenhouts l.c. (1952) 155, l.c.
(1956) 215. Basionym: Canarium connarifolium Perkins, Fragm. Fl. Philip. 2 (1904) 92. Type:
Merrill 787, Philippines, Palawan (GH, K, L). Synonym: P. philippinensis Elmer, Leafl. Philip. Bot.
7 (1915) 2571.
Small tree. Leaves with up to 3 pairs of leaflets; petiolules of lateral leaflets to 2 cm long,
swollen at both ends; blade ovate to elliptic, 311 x 25 cm; base cuneate, margin entire,
apex pointed with mucronate tip; midrib slightly sunken above; lateral veins 57 pairs,
looping, visible below, faint above; intercostal veins reticulate, very faintly visible on both
surfaces. Inflorescences axillary, or female inflorescences sometimes pseudo-terminal.
Flowers tomentose. Fruits 78 x 513 mm, sparsely pubescent to glabrous, with 14
pyrenes.
Distribution. Borneo, Philippines (Palawan). Very uncommon, known by a single
collection SAN 89964 from Semporna in Sabah. According to Leenhouts l.c. (1956), this
species is rather isolated taxonomically from the rest.
6. SANTIRIA Blume
(Santir, Blume's native guide to G. Salak, Java)
kerantai (Sabah), seladah (Sarawak)
l.c. (1850) 209; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 367; Ridley l.c. (1922) 376; Kalkman l.c.
(1954) 522; Leenhouts l.c. (1956) 229; Kochummen l.c. (1972) 145, Sandakania 5 (1994) 83;
Cockburn l.c. 47; Anderson l.c. 157; Wong l.c. 212; Whitmore, Tantra & Sutisna l.c. 38; Ng l.c. 38.
Synonyms: Trigonochlamys Hook. f., Trans. Linn. Soc. 23, 1 (1860) 170; Icicaster Ridl., J. Str. Br. R.
As. Soc. 75 (1917) 15.
Medium-sized to large dioecious trees. Bark brown, fawn or grey, smooth to fissured, scaly
or flaky, with many small or large lenticels; inner bark either pink, soft, and laminated or
yellowish white, hard mottled and granular (S. griffithii, S. rubiginosa), with white to
colourless exudate darkening on drying. Pith of twigs and petioles with or without vascular
strands. Leaves without stipules, imparipinnate; leaflets entire, petiolules only slightly
swollen at both ends; petiole usually flat or channelled on the upper surface. Inflorescences
usually axillary, rarely terminal panicles. Flowers unisexual, 3-merous; sepals free or
united; petals usually with thickened inflexed apex; stamens 6 or 3, anthers basifixed or
83
dorsifixed; disc intrastaminal, glabrous; ovary 3-celled, glabrous, in male flowers reduced;
stigma (sub)-sessile. Fruits brightly coloured drupes, irregularly globose or ellipsoid, more
or less oblique, in big bunches, usually seated on persistent calyx; stigma off-centre,
sometimes nearly basal; rind thin and firm, almost smooth when dry; stone thinly woody
containing one fertile and two sterile cells. Seeds almost round, not angled; germination in
S. laevigata and S. oblongifolia epigeal, cotyledons 5-lobed, fleshy, first two leaves
opposite, subsequent leaves alternate, leaves in young seedling simple; in S. griffithii and S.
rubiginosa germination hypogeal, first two leaves opposite, subsequent leaves alternate and
then spiral, leaves in young seedlings pinnate.
Distribution. About 22 species, with 6 species restricted to W Africa and the rest in
Malesia (mainly Sumatra, Peninsular Malaysia, Borneo). 15 species in Sabah and Sarawak.
Ecology. Lowland forest, including swamp forest, to submontane forest to 1650 m.

Uses. The timber is commonly grouped together with other members of the family and sold
as kedondong. The wood is light hardwood. It is essentially similar in anatomical structure
to the timber of Canarium and Dacryodes but is more variable in colour, weight and
hardness. Silica is present in all species. S. laevigata and S. tomentosa are important sources of
kedondong timber in Peninsular Malaysia. The buttresses of S. tomentosa are interlocked
and frequently used for parang sheaths.
Taxonomy. The genus is sub-divided into two sections: section Santiria and section
Icicopsis. Apart from the differences in floral characters by which these sections were
established, evidence from wood anatomy and germination supports the distinction of these
sections. In Sabah and Sarawak, section Icicopsis is represented by S. griffithii and S.
rubiginosa.
Key to Santiria species

1.
Leaflets hairy on the lower side, at least on the midrib..............................................2

2.
Leaflets densely velvety or woolly hairy on the lower side............................................3

Leaflets sparsely hairy on the lower side......................................................................5
3.
Intercostal veins scalariform................................................................15. S. tomentosa

Intercostal veins reticulate.........................................................................................4
4.
Leaflets concave, hairs dark brown; petiolule not swollen at ends. Twigs covered with
lenticels. Stigma on fruit more than 90o excentric......................................10. S. mollis
Leaflets not concave, hairs yellowish brown; petiolule swollen at ends. Twigs not so.
Stigma prominently excentric, near pedicel.............................................1. S. apiculata
5.
Twigs stout, c. 1.5 cm thick. Terminal bud c. 2 cm long. Petiole strongly channelled
above with deep groove.......................................................................4. S. grandiflora
Twigs slender, c. 6 mm thick. Terminal bud up to 0.5 cm long. Petiole not channelled or
grooved .......................................................................................14. S. sarawakana
84
6.
Lateral veins sunken above..........................................................6. S. impressinervis

Lateral veins not sunken above..................................................................................7
7. Twigs stout, 1.53 cm thick.........................................................................................8

Twigs slender, to 1 cm thick........................................................................................9
8.
Terminal bud c. 6 cm long with curved tip. Leaflets 79 cm wide; petiolules of lateral
leaflets to 2.5 cm long. Stigma of fruit more than 90o excentric........7. S. kalkmaniana
Terminal bud c. 1.5 cm long, tip not curved. Leaflets 1322 cm wide; petiolules of
lateral leaflets 45 cm long. Stigma of fruit near pedicel.....................9. S. megaphylla
9.
Stigma of fruit more than 90o excentric, near pedicel.....................................................10

Stigma of fruit less than 90o excentric.......................................................................12
10. Terminal bud c. 2 cm long. Petiole strongly channelled at base with sharp edges. Midrib
raised above..................................................................................2. S. conferta
Terminal bud up to 1 cm long. Petiole not so. Midrib flattened to slightly sunken
above.........................................................................................................................11
11. Midrib below channelled. Leaflet margin not curled inwards; intercostal veins distinct
below......................................................................................................1. S. apiculata
Midrib below not channelled. Leaflet margin curled inwards; intercostal veins invisible
below.........................................................................................................3. S. dacryodifolia
12. Terminal bud 0.52.5 cm long, often resin-coated. Leaflets drying reddish brown or
dark brown..............................................................................................8. S. laevigata
Not this combination of characters..........................................................................13
13. Rachis blackish, oily, shiny. Lateral veins of leaflets very faint. Stigma on fruit only
slightly excentric..................................................................................11. S. nigricans
Rachis not so. Lateral veins of leaflets distinct. Stigma in fruit 90o excentric.............14
14. Lateral veins distinctly looping near margin; intercostal veins parallel to lateral veins.
Fruits to 1.3 cm long..................................................................................................15
Lateral veins faintly looping near margin; intercostal veins not parallel to lateral veins.
Fruit to 3 cm long.............................................................................12. S. oblongifolia
15. Leaflets not asymmetric at base, petiolules of lateral leaflets to 5 mm long, not swollen
at ends.....................................................................................................5. S. griffithii
Leaflets asymmetric at base, petiolules of lateral leaflets to 2 cm long, swollen at both
ends....................................................................................................13. S. rubiginosa
1. Santiria apiculata A.W. Benn.

(Latin, apiculatus = furnished with a little point; the pointed leaflets)
in Hooker f. l.c. 537; Ridley l.c. (1922) 378; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 375;
Masamune l.c. 367; Kalkman l.c. (1954) 538; Leenhouts l.c. (1956) 234; Burgess l.c. 62; Kochummen
l.c. (1972) 146; Cockburn l.c. 48; Anderson l.c. 157; Whitmore, Tantra & Sutisna l.c. 38. Type:
Maingay 303, Malacca (holotype K; isotype L).
85
Small to medium-sized tree rarely more than 30 m tall. Bark grey or brown, smooth to
scaly; inner bark pinkish with droplets of white sap and with strong resinous smell.
Sapwood pale yellow. Twigs pale or whitish, glabrous or densely brownish hairy (var.
pilosa). Leaves trifoliolate or with 25 pairs of leaflets; blade glabrous, drying greenish or
hairy below and drying brownish (var. pilosa), elliptic or lanceolate, 3.516 x 1.57 cm;
base broadly cuneate, apex pointed with long tip; midrib flattened to slightly sunken above,
distinctly grooved below; lateral veins 514 pairs, distinct below, faint above, curving and
joining a few millimeters away from margin; intercostal veins reticulate, faintly visible
below, faint to invisible above; with a strong resinous smell when freshly crushed;
petiolules usually whitish, glabrous or hairy, 0.53 cm long. Flowers yellow or reddish,
usually in short axillary inflorescences, glabrous or hairy (var. pilosa). Fruits globose or
ellipsoid, 12 x 1 cm, stigma usually more than 90o excentric, sometimes near the pedicel.
Key to varieties
Twigs, leaflets and flowers densely hairy..............................................................................
var. pilosa (Engl.) Kalkman
in Leenhouts l.c. (1956) 236. Basionym: Santiria pilosa Engl. in DC. l.c. 159.
Endemic to Borneo. In Sabah and Sarawak widely distributed from lowland to

submontane forest but less common compared to var. apiculata.
Twigs, leaflets and flowers glabrous.....................................................................................
var. apiculata
Synonyms: S. beccarii Engl. in DC. l.c. 159; S. glabra Merr. l.c. (1915) 30; S. minutiflora Ridl.
l.c. (1922) 377; S. brachystachys Ridl. l.c. (1925) 79; Canarium pauciflorum Ridl. l.c. (1930) 80;
Haplolobus borneensis H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 418.
Sumatra, Peninsular Malaysia, Borneo, Philippines, Celebes, Moluccas. Common and

widely distributed throughout Sabah and Sarawak. Mixed dipterocarp to submontane
forests to 1300 m.
2. Santiria conferta A.W. Benn.

(Latin, confertus = crowded; the flowers)
in Hooker f. l.c. 537; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 381; Kalkman l.c. (1954) 528;
Leenhouts l.c. (1956) 233; Burgess l.c. 62; Kochummen l.c. (1972) 146; Whitmore, Tantra & Sutisna
l.c. 38. Type: Griffith 1150, Malacca (holotype K; isotype P). Synonyms: S. wrayi King l.c. (1893)
259; S. conferta var. wrayi (King) H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 381.
Small to medium-sized tree to 30 m tall, 50 cm diameter. Bark grey-brown; inner bark

yellow. Terminal bud c. 2 cm long. Twigs greyish yellow, c. 1 cm thick. Leaves: petiole
strongly channelled with sharp edges; leaflets thickly leathery, oblong, ovate or elliptic,
sometimes glaucous below, 1117 x 5.58.5 cm; base rounded to broadly cuneate, apex
pointed; midrib raised above; lateral veins 1114 pairs, arching near margin, raised on both
surfaces; intercostal veins reticulate, faintly visible on both surfaces. Flowers in axillary
inflorescences. Fruits reddish when fresh, subglobose, 1.52 x 1.21.5 cm, stigma strongly
excentric, near pedicel; surface warty.
Distribution. Sumatra, Peninsular Malaysia, Borneo. In Borneo uncommon, known from a
few collections from Sabah and Sarawak.
Ecology. Submontane forest at 12001650 m. Fruiting in November and December.
86
3. Santiria dacryodifolia Kochummen

(with leaves resembling those of Dacryodes)
l.c. (1994) 83. Type: Tong & Jugah S. 32944, Sarawak (holotype KEP; isotypes K, L, SAN, SAR,
SING).
Small tree to 12 m tall, 25 cm diameter; bole fluted. Bark light brown, flaky. Twigs
brownish, finely lenticellate, c. 7 mm thick. Leaves with 2 pairs of leaflets; rachis pustular,
rounded, not swollen at base; blade coriaceous, glabrous, drying to brownish, elliptic to
rhomboid, 8.510.5 x 3.55.5 cm; base cuneate, margin curled inwards, apex pointed;
midrib sunken above; lateral veins 57 pairs, faintly visible on both surfaces; intercostal
veins reticulate, visible above, invisible below; petiolules of lateral leaflets 11.5 cm long,
of terminal leaflet c. 4.5 cm long, strongly swollen at both ends. Inflorescences and flowers
unknown. Infructescence axillary and terminal, c. 14 cm long. Fruits yellowish to deep red
when fresh, drying to dark brown, obliquely oblong, 78 x 56 mm, surface slightly rugose,
stigma more than 90o excentric, near the pedicel.
Distribution. Endemic to Sarawak. Uncommon, known from the type only from Ulu Sg.
Maria, Lawas.
Ecology. Ridge tops at about 1400 m.
Appears close to S. apiculata differing in the stout twigs, thick leaflets with recurved
margins and in the petiolules with strongly swollen ends.
4. Santiria grandiflora Kalkman

(Latin, grandis = large, florus = flower)
l.c. (1954) 525; Leenhouts l.c. (1956) 232; Anderson l.c. 157; Whitmore, Tantra & Sutisna l.c. 38.
Type: Richards 1278, Sarawak (holotype K; isotype SING).
Medium-sized tree to 30 m tall. Bark grey-brown, scaly; inner bark pink, with droplets of
white sap. Sapwood pale. Twig stout, c. 1.5 cm thick, lenticellate, dark brown powdery
hairy towards the tip. Terminal bud c. 2 cm long. Leaves with 46 pairs of leaflets;
petiolules of lateral leaflets 12 cm long, powdery hairy; blade glabrous above except the
midrib, powdery hairy below, oblong-elliptic, 1629 x 59 cm; base rounded or broadly
cuneate, apex pointed; midrib raised below, flat and faint above; lateral veins 1524 pairs;
intercostal veins scalariform-reticulate, distinct below, faint above; petiole strongly
channelled above with deep groove, powdery brown-hairy. Flowers (male) in axillary
inflorescences. Infructescences 918 cm long. Fruits obliquely ellipsoid, yellowish green,
ripening pink, with white waxy bloom on drying, 1.51.9 x 1.11.3 cm, with faintly
wrinkled surface; stigma slightly excentric; persistent calyx reflexed.
87
Distribution. Endemic to Borneo. Uncommon, recorded from Mt. Dulit, Mulu National
Park, and Lambir Hills in Sarawak. Also known from Brunei.
Ecology. Mixed dipterocarp forest to 300 m, on ridges or yellow sandy clay soils. Fruiting
in March and April.
This species is somewhat similar to S. tomentosa but the strongly channelled petiole will
distinguish it.
5. Santiria griffithii (Hook. f.) Engl.

(W. Griffith, 181045, surgeon at Malacca)
Bot. Jahrb. 1 (1881) 43; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 394; Masamune l.c. 368;
Kalkman l.c. (1954) 545; Leenhouts l.c. (1956) 236; Kochummen l.c. (1972) 147; Burgess l.c. 62;
Anderson l.c. 157; Whitmore, Tantra & Sutisna l.c. 38. Basionym: Trigonochlamys griffithii Hook. f.
l.c. (1860) 170. Type: Griffith 1148, Malacca (holotype K; isotype L). Synonym: Santiria borneensis
Engl. l.c. 43.
Medium-sized tree to 30 m tall, 55 cm diameter; buttresses to 2 m tall. Bark grey-brown,

smooth to scaly; inner bark brownish. Sapwood pale. Twigs reddish brown-hairy when
young. Leaves with 511 pairs of opposite or subopposite leaflets; rachis minutely hairy;
petiolules of leaflets c. 5 mm long, hairy, not swollen; blade glabrous or sparsely hairy
below, lanceolate or elliptic, 510.5 x 1.53 cm; base broadly cuneate to rounded, slightly
unequal, apex pointed with long tip; midrib flattened above; lateral veins 1518 pairs,
faintly visible below, faint to inconspicuous above, distinctly curving and joining near
margin; intercostal veins reticulate, visible below, faint above. Inflorescences axillary,
pubescent. Flowers tomentose; calyx deeply divided; petals pubescent; stamens 6, filaments
adnate to disc, pistil in male flowers reduced. Fruits obliquely globose, 813 x 816 mm;
stigma almost 90o excentric.
Distribution. Sumatra, Peninsular Malaysia, Borneo. In Sabah uncommon, recorded from
Keningau and Tawau only. In Sarawak more common and widely distributed. Also occurs
Ecology. Mixed dipterocarp forest on yellow sandy soils to 300 m. Flowering in
MarchNovember and fruiting in SeptemberOctober.
6. Santiria impressinervis Kochummen

(Latin, impressus = pressed in, nervus = nerve; the sunken veins on the upper leaflet
surface)
l.c. (1994) 83. Type: Nooteboom & Chai 2113, Sarawak (holotype KEP; isotype SAR).
Small tree to 15 m tall, 15 cm diameter. Bark yellowish, smooth. Twigs c. 4 mm thick,

brownish, with small lenticels. Leaves with 23 pairs of leaflets; rachis slender, slightly
flattened above towards the base; petiolules of lateral leaflets 0.81.5 cm long, that of
terminal leaflet c. 2 cm long, strongly swollen at both ends; blade glabrous, coriaceous,
drying to yellowish brown, ovate to lanceolate, 69 x 24 cm; base broadly cuneate, slightly
asymmetric, apex pointed, tip 510 mm long; midrib faintly sunken above; lateral veins 79
pairs, sunken above, curving and joining near margin; intercostal veins invisible.
Inflorescences and flowers unknown. Infructescences axillary, to 11 cm long. Fruits
88
glaucous green when fresh, drying to brownish, smooth, ovoid, c. 10 x 7 mm, stigma
prominently off-centre, near pedicel.
Distribution. Endemic to Sarawak. Uncommon, known only from the type from the
Pamerario river at Bario in the Kelabit Highlands.
Ecology. Old secondary forest at 1000 m.
Appears close to S. apiculata but differing in the thickly coriaceous leaflets with invisible
intercostal veins and in the sunken lateral veins.
7. Santiria kalkmaniana Kochummen
Fig. 6.
(C. Kalkman, former director of the Rijksherbarium, Leiden, Netherlands)

l.c. (1994) 86. Type: Saikeh SAN 72293, Sabah (holotype SAN; isotypes A, K, SAR, SING).
Medium-sized tree to 30 m tall, 45 cm diameter; buttresses to 3 m tall. Bark greyish,

smooth, lenticellate; inner bark yellowish. Sapwood pale yellow. Twigs very stout, 1.52
cm thick, grey-brown, lenticellate. Terminal bud lanceolate, rusty brown, c. 6 cm long with
curved tip. Leaves with 5 pairs of leaflets, glabrous; rachis c. 54 cm long, petiole channelled
above; petiolules of lateral leaflets 22.5 cm long, slightly swollen at both ends, petiolules
of terminal leaflets 58.5 cm long; blade thinly leathery, oblong to lanceolate, 1835 x 79
cm; base rounded or broadly cuneate, asymmetric, apex pointed; midrib raised above; lateral
veins 1619 pairs, raised below, faint above; intercostal veins reticulate, more distinct on the
upper surfaces. Inflorescences (male) axillary, glabrous, panicles to 27 cm long, axes
sharply angled. Flowers (male) glabrous; calyx red, cup-shaped; petals deep scarlet, oblong;
stamens 6, inserted on rim of disc; disc annular; ovary rudimentary. Infructescences c. 6 cm
long, with short branches. Fruits light yellow when fresh drying black, subglobose or
hemispherical, 1.72 x 1.01.8 cm, flattened above; stigma more than 90o excentric.
Distribution. Endemic to Borneo. Uncommon, known from few collections from Sabah
(SAN 72293, Sipitang; SAN 49754, Ranau), and Sarawak (S. 14727, Kapit; S. 39782, G.
Mulu, and S. 4030 from Baram).
This species appears close to S. laevigata but differs in the long rusty brown curved terminal
bud and in the fruit stigmas that are more than 90o off-centre.
8. Santiria laevigata Blume

(Latin, laevigatus = smooth and polished; the leaflets)
l.c. (1850) 211; King l.c. 257; Ridley l.c. (1922) 378; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932)
382; Masamune l.c. 368; Kalkman l.c. (1954) 535; Leenhouts l.c. (1956) 232; Burgess l.c. 62;
Type: Korthals, s.n. (= Leiden no. 898.321201), Sumatra (L). Synonym: Canarium laevigatum
(Blume) Miq. l.c. (1859) 648.
89
Medium-sized tree to 30 m tall, 80 cm diameter; buttresses to 4 m tall. Bark grey-brown,

fissured and scaly, lenticellate; inner bark pinkish, laminated, with droplets of white sap.
Sapwood pale white. Terminal bud 0.52.5 cm long, often resin-coated. Twigs c. 1 cm
thick, lenticellate, dark brown. Leaves with 14 pairs of leaflets; petiole flattened above
with sharp edges (forma laevigata) or not flattened (forma glabrifolia); petiolules of lateral
leaflets 11.5 cm long, slightly swollen at both ends, sometimes the swollen parts drying
black; blade drying reddish or dark brown, elliptic, ovate or oblong, 712 x 2.56.5 cm;
base rounded to broadly cuneate, sometimes unequal, apex pointed; midrib raised above;
lateral veins 715 pairs, fairly raised on both surfaces, curving and joining near margin;
intercostal veins reticulate, very faintly visible on both surfaces. Flowers glabrous, in
axillary inflorescences. Fruits rounded to oblong, 12 cm across; stigma less than 90o
excentric.
Key to forms
Petiole flattened above, at base with sharp edges...................................................................
forma laevigata
Synonyms: Santiria laevigata forma typica H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 384; S.
violacea H.J. Lam, Ann. Jard. Bot. Btzg. 42 (1932) 205.
Sumatra, Peninsular Malaysia, Borneo, and Celebes. In Sarawak uncommon, known

from two collections (S. 32817 and S. 41457). Not yet recorded from Sabah.
Petiole rounded at base, without sharp edges........................................................................
forma glabrifolia (Engl.) H.J. Lam
Ann. Jard. Bot. Btzg. 42 (1932) 205. Basionym: Santiria glabrifolia Engl. in DC. l.c. 164. Type:
Beccari PB 3756, Sarawak (FI).
Sumatra, Peninsular Malaysia, and Borneo. Common and widely distributed in Sabah
and Sarawak.
Ecology. Mixed dipterocarp, mixed peat swamp and kerangas forests on low-nutrient soils
to submontane forests to 1200 m.
90
91
9. Santiria megaphylla Kalkman

(Greek, mega = large, phyllon = leaf)
l.c. (1954) 533; Leenhouts l.c. (1956) 236; Burgess l.c. 62; Anderson l.c. 157; Whitmore, Tantra &
Sutisna l.c. 38. Type: Beccari PB 3059, Sarawak (FI).
Medium-sized tree to 25 m tall. Twigs very stout, c. 3 cm thick, grey with circular lenticels
and with stipule-like undeveloped leaves. Terminal bud c. 1.5 cm long. Leaves with 35
pairs of leaflets; petiole stout, strongly channelled at base; petiolules of lateral leaflets 45
cm long, petiolules of terminal leaflets c. 11 cm long, strongly swollen at both ends; blade
thickly leathery, very large, oblong, 2561 x 1322 cm; base broadly cuneate, apex
pointed; midrib raised above; lateral veins 814 pairs, raised on both surfaces, curving and
joining near margin; intercostal veins reticulate, faintly visible on both surfaces. Flowers in
axillary inflorescences. Fruits pink when fresh, asymmetric, ellipsoid, 1.51.7 x 11.25
cm; stigma excentric, near pedicel.
Distribution. Endemic to Borneo. Uncommon, known from few collections from Sabah
(SAN A 1746, Beaufort; SAN 15152, Sipitang) and Sarawak (S. 38747 and S. 43958 from
Semengoh Arboretum and Mt. Dulit), and ecological plots at Bako and Lambir NP, and
Segam FR. Also known from Brunei.
Ecology. Mixed dipterocarp forests on deep sandy humult ultisols.
The stout twigs and very large leaflets are good diagnostic features for the species.
10. Santiria mollis Engl.

(Latin, mollis = soft-hairy; the indumentum of young twigs, leaves and flowers)
in DC. l.c. 156; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 388; Masamune l.c. 368; Kalkman l.c.
(1954) 530; Leenhouts l.c. (1956) 231; Burgess l.c. 62; Anderson l.c. 157; Whitmore, Tantra &
Sutisna l.c. 38. Type: Beccari PB 3497, Sarawak (FI). Synonym: Canarium hirtipetalum Ridl. l.c.
(1930) 84.
Medium-sized tree to 24 m tall, 70 cm diameter; buttresses to 3 m high. Bark brown,

smooth; inner bark reddish brown, with white latex. Sapwood pale yellow. Twigs densely
reddish brown-hairy with abundant small lenticels when young. Terminal bud stout, c. 1 x
0.5 cm, hairy as the twig. Leaves with 24 pairs of leaflets; rachis densely reddish brownhairy; petiolules of lateral leaflets to 1 cm long, not swollen at both ends, hairy; blade
densely velvety hairy below and on midrib above, obovate, oblong or elliptic, often concave,
7.515.5 x 37 cm; base cuneate or rounded, often unequal, margin curled inwards, apex
pointed with long tip; midrib raised above; lateral veins 1013 pairs, raised below, faint to
sunken above; intercostal veins reticulate, raised below, faint to invisible above. Flowers
(male) in hairy axillary inflorescences; sepals almost free, sparsely hairy outside; petals
densely hairy on both sides. Fruits subglobose, 1.51.8 cm across; stigma more than 90o
excentric.
Distribution. Endemic to Borneo. Uncommon, in Sabah recorded from Beaufort and
Tawau. In Sarawak collected from Semengoh Arboretum and Lambir National Park. Also
known from Brunei and Kalimantan.
92
Ecology. Mixed dipterocarp forest to 200 m, often by streams. Flowering in March and
November and fruiting in February and March.
Very atypical of Santiria in vegetative features.
11. Santiria nigricans Kochummen

(Latin, nigricans = becoming black; the leaf rachis)
l.c. (1994) 89. Type: Ilias & Yeo S. 38319, Sarawak (holotype KEP; isotypes K, L, MO, SAN, SAR).
Large tree to 30 m tall, 100 cm diameter. Bark brownish, scaly; inner bark reddish brown.
Sapwood whitish. Twigs brown or greyish brown, 34 mm thick. Leaves with up to 3 pairs
of leaflets, glabrous; rachis slender, dark brown, surface oily, slightly swollen at the very
base; petiolules of lateral leaflets slender, 11.5 cm long, of the terminal leaflets to 4 cm
long, slightly swollen at both ends, drying to dark brown; blade thinly leathery, drying to
grey-brown, elliptic to lanceolate, 6.511.5 x 2.84.7 cm; base cuneate, strongly asymmetric, apex pointed; midrib raised above; lateral veins 711 pairs, very faint on both
surfaces; intercostal veins reticulate, faintly visible on both surfaces. Inflorescences (male)
axillary and terminal, glabrous. Flowers (male) with green calyx and white petals. Fruits
green, ripening deep purple to blackish, drying to pale brownish, subglobose, 1114 mm
across, surface with shallow reticulate veins; stigma only slightly off-centre.
Distribution. Endemic to Sarawak; of scattered distribution in Lambir National Park (S.
38319 and S. 46599), Mulu National Park (S. 42406), Balingian (S. 23687), Selampit (S.
24939) and Sg. Jelalong (S. 48804).
Ecology. Mixed dipterocarp forest from lowlands to 1600 m.
Somewhat close to S. oblongifolia but the dark brown rachis, very faint lateral veins and the
subglobose fruits distinguish this species.
12. Santiria oblongifolia Blume

(Latin, oblongus = rather long, folium = leaf; the leaflet shape)
l.c. (1850) 211; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 372; Masamune l.c. 369; King l.c. 257;
Ridley l.c. (1922) 379; Kalkman, l.c. (1954) 537; Leenhouts l.c. (1956) 233; Burgess l.c. 62;
Kochummen l.c. (1972) 150; Anderson l.c. 157; Whitmore, Tantra & Sutisna l.c. 39. Type:
Praetorius, s.n. (= Leiden no. 898.321221), Sumatra (L). Synonyms: S. caesia Engl. in DC. l.c. 166;
S. latifolia Stapf. ex Ridl. l.c. (1930) 86.
Medium-sized tree 30 m tall, very rarely to 50 m tall, 80 cm diameter; buttresses to 1.5 m

high. Bark grey-brown, smooth, lenticellate, rarely scaly or flaky; inner bark pink, laminated.
Sapwood pale white. Leaves with 24 pairs of leaflets; petiolules of lateral leaflets 13.5
cm long, swollen at both ends; blade thinly to thickly leathery, oblong to lanceolate, 821.5
x 37.5 cm; base broadly cuneate, apex pointed; midrib raised above; lateral veins 810
pairs, visible on both surfaces, curving and joining near margin; intercostal veins reticulate,
faint on both surfaces. Flowers in axillary or terminal inflorescences, glabrous or sparsely
hairy. Fruits oblong-ellipsoid, 1.23 x 1.12.2 cm; stigma slightly excentric.
93
Distribution. Sumatra, Peninsular Malaysia, Borneo. Widely distributed in Sabah. In

Sarawak collected mainly from the Bako National Park and Semengoh Forest Reserve area.
Also found in Kalimantan.
Ecology. Mixed dipterocarp forest on yellow sandy and clay soils, heath forest and
submontane forest to 1000 m. Flowering in March, May and September and fruiting in
January, MarchMay and September.
13. Santiria rubiginosa Blume

(Latin, rubiginosus = rusty coloured; the indumentum)
l.c. (1850) 213; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 399; Masamune l.c. 369; Kalkman l.c.
(1954) 542; Leenhouts l.c. (1956) 237; Burgess l.c. 62; Kochummen l.c. (1972) 151; Anderson l.c.
158; Whitmore, Tantra & Sutisna l.c. 39. Type: Praetorius, s.n. (=Leiden no. 898.321229),
Sumatra (L). Synonym: Canarium rubiginosum (Blume) Miq. l.c. (1859) 651.
Medium-sized tree to 28 m tall, 40 cm diameter; buttresses to 1.5 m high. Bark grey,

smooth, lenticellate. Twigs brownish, glabrous, except the tip. Leaves with 14 pairs of
leaflets, sometimes mixed with simple leaves; petiolules of lateral leaflets 0.52 cm long,
either rounded or strongly flattened, swollen at both ends; blade glabrous or very sparsely
powdery hairy below, elliptic to lanceolate, 4.513.5 x 1.56 cm; base cuneate, often
unequal, apex pointed with long tip; midrib raised above; lateral veins 913 pairs, visible
below, faint above, arching and joining near margin; intercostal veins reticulate, faintly
visible on both surfaces. Flowers in axillary inflorescences, glabrous; sepals almost free;
stamens 3. Fruits globular or ellipsoid, 1014 x 810 mm; stigma less than 90o excentric.
S. rubiginosa is very similar to S. griffithii in vegetative characters but the asymmetric
leaflets of S. rubiginosa are distinctive.
Key to varieties
1.
Pedicels 13 mm long, shorter or as long as the flowers...............................................2

Pedicel 28 mm long, longer than the flowers................................................................
var. pedicellata (Ridl.) Kalkman
l.c. (1954) 544. Basionym: Santiria pedicellata Ridl. l.c. (1930) 86. Synonym: S.
minimiflora Ridl. l.c. (1930) 87. Type: Haviland 1866, Sarawak, Sibu, Rejang (SING).
Endemic to Borneo. Common in Sarawak but not yet reported from Sabah. Freshwater, heath and peat swamp forest.
2.
Leaflet apex caudate-acuminate, more than 1 cm long; petiolule flattened, the ends
blacken on drying..........................................................................................................
var. latipetiolata Kochummen
l.c. (1994) 89. Type: George S. 40254, Sarawak (holotype KEP; isotypes E, K, L, MO,
SAN, SAR).
Endemic to Sarawak, known from two collections from the Lambir National Park
(S. 40254 and S. 47190). Mixed dipterocarp forest on ridges at 250 m.
Leaf apex not so, petiolule not flattened.........................................................................
var. rubiginosa
Synonyms: Santiria planchonii A.W. Benn. in Hooker f. l.c. 536; C. planchonii (A.W.
Benn.) King l.c. 240; Icicaster planchonii (A.W. Benn.) Ridl. l.c. (1917) 15; S. havilandii
Ridl. l.c. (1930) 85.
Sumatra, Peninsular Malaysia, Borneo, and New Guinea. Common and widely
distributed in Sarawak in mixed dipterocarp forest but uncommon in Sabah and
known only by a single collection, SAN 16678, from Sipitang at 750 m.
94
14. Santiria sarawakana Kochummen

(of Sarawak)
l.c (1994) 91. Type: Tong et al., S. 34287, Sarawak (holotype SAR; isotypes K, KEP, L, MO, SAN).
Medium-sized tree to 26 m tall, 30 cm diameter. Bark dark brown, flaky. Twigs whitish,
powdery brown-hairy. Terminal bud ovate, c. 5 x 3 mm. Leaves with 3 pairs of leaflets;
rachis hairy as the twig, rounded; petiolules of lateral leaflets 715 mm long, of terminal
leaflets 1.53 cm long, hairy and swollen at both ends; blade leathery, glabrous above
except the midrib, densely hairy on midrib and lateral veins below, ovate to oblong or
elliptic, 614 x 35 cm; base broadly cuneate, asymmetric, margin curled inwards, apex
pointed; midrib raised above; lateral veins 912 pairs, curving and joining near margin,
raised below, visible above; intercostal veins scalariform-reticulate, visible on both surfaces.
Inflorescences and flowers unknown. Infructescences axillary, to 11.5 m long, axes hairy.
Fruits glaucous black when fresh, black on drying, oblong, 1113 x 710 mm; stigma less
than 90o excentric.
Distribution. Endemic to Sarawak. Uncommon, known only from Miri and the Sabal Forest
Reserve (S. 24119 and S. 34287).
Close to Santiria tomentosa but the sparsely hairy leaflets and short terminal buds are quite
distinct.
15. Santiria tomentosa Blume

(Latin, tomentosus = densely pubescent with matted, woolly or short hairs; the young twigs,
buds and leaflets)
l.c. (1850) 211; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 391; Masamune l.c. 369; Kalkman l.c.
(1954) 529; Leenhouts l.c. (1956) 231; Burgess l.c. 62; Kochummen l.c. (1972) 151; Cockburn l.c. 49;
Anderson l.c. 158; Whitmore, Tantra & Sutisna l.c. 39. Type: Korthals, s.n. (= Leiden no.
95
96
898.321245), Sumatra (L). Synonyms: Canarium korthalsii Miq. l.c. (1859) 645; Santiria multiflora
A.W. Benn. in Hooker f. l.c. (1875) 538; C. micrantherum Stapf ex Ridl. l.c. (1930) 82; S. mollissima
Ridl. l.c. (1930) 85.
Medium-sized tree to 30 m tall, 50 cm diameter; buttresses to 2 m high. Bark grey-brown,

scaly; inner bark pinkish with droplets of white sap. Sapwood pale yellow. Twigs 515 mm
thick, slightly angled, woolly hairy. Terminal bud to 2 cm long, hairy. Leaves with 14
pairs of leaflets; rachis woolly hairy; petiolules of lateral leaflets 0.52 cm long, hairy;
blade woolly hairy below and on midrib above, ovate to oblong, 1127 x 412 cm; base
rounded or cuneate, sometimes unequal, apex pointed; midrib raised above; lateral veins 10
20 pairs, raised on both surfaces, curving and joining near margin; intercostal veins
scalariform-reticulate, distinct below, faint above. Flowers hairy, in axillary inflorescences.
Fruits globose or oblong, 22.7 x 1.52 cm; stigma less than 90o excentric.
Distribution. Sumatra, Peninsular Malaysia, Borneo. Common and widely distributed in
Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. Mixed dipterocarp forests, rarely in swamps, to 300 m. Flowering in MayOctober
and fruiting in March, April, June, October and November.
There is great variation in the size of leaflets and in the degree of pubescence.
7. SCUTINANTHE Thwaites
(Latin, scutum = shield, anthus = flower; the shape of the flower)
in Hooker f., J. Bot. Kew Misc. 8 (1856) 266; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 420;
Leenhouts l.c. (1952) 160, l.c. (1956) 246; Kochummen l.c. (1972) 152; Anderson l.c. 158; Wong l.c.
215; Ng l.c. 38; Whitmore, Tantra & Sutisna l.c. 39.
Dioecious trees; bole smooth with horizontal ring or occasionally scaly; inner bark raspberryred with large beads of creamy to colourless exudate. Pith of branchlets and petioles without
vascular strands. Leaves without stipules; leaflets entire. Inflorescences axillary, paniculate.
Flowers unisexual, 5-merous, receptacle cup-shaped; sepals free; petals free with slightly
thickened apex; stamens 10, confluent at base, episepalous ones longer, in female flowers
slightly reduced in size, sterile; disc intrastaminal, adnate to receptacle; ovary hairy, 3celled, only slightly reduced in male flowers. Fruits drupaceous, stigmatic scar nearly
apical, pericarp fleshy; pyrene hard and bony, usually 2 cells strongly reduced; persistent
calyx not enlarged. Seeds one; germination in S. brunnea epigeal; cotyledons leafy, entire;
leaves of young seedling simple, alternate to spiral.
Distribution. Two species in Sri Lanka, S Burma, W Malesia and Celebes. One species in
Sabah and Sarawak.
Ecology. Lowland forests to 100 m.
Uses. The light hardwood timber is commonly grouped together with other members of the
family and sold as kedondong.
97
98
Scutinanthe brunnea Thwaites
Fig. 7.
(Latin, brunneus = brown; the indumentum on the inflorescence)

in Hooker f. l.c. (1856) 267; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932) 420; Leenhouts l.c. (1952)
162, l.c. (1956) 247; Burgess l.c. 62; Kochummen l.c. (1972) 152; Anderson l.c. 158; Whitmore,
Tantra & Sutisna l.c. 39. Type: Thwaites 1149, Ceylon (K). Synonyms: Garuga brunnea (Thwaites)
Marchand, Adansonia 8 (1867) 34, 66; Canarium brunneum (Thwaites) Beddome, Fl. Sylv. 1 (1868) t.
127.
Medium-sized tree to 30 m tall, 30 cm diameter. Sapwood pale. Twigs brownish,

lenticellate. Leaves with 36 pairs of leaflets; petiolules of lateral leaflets 12.5 cm long,
swollen at both ends; blade elliptic to obovate, 821 x 48 cm; base broadly cuneate,
unequal, apex pointed; midrib sunken above; lateral veins 810 pairs, raised below, faint
above, curving and joining near margin; intercostal veins reticulate, visible on both surfaces.
Inflorescences rusty red-pubescent. Flowers densely pubescent with greenish white inner
face. Fruits ellipsoid, 46.5 x 23 cm, green, ripening yellow, with persistent calyx.
Distribution. Sri Lanka, Sumatra, Peninsular Malaysia, and Borneo. In Sabah and Sarawak
uncommon, known only by a few collections. Also in Brunei and Kalimantan.
8. TRIOMMA Hook. f.
(Greek, tri = 3, omma = eyes or openings;
the triangular seed showing 3 openings in cross-section)
Trans. Linn. Soc. 23 (1860) 171; Ridley l.c. (1922) 369; H.J. Lam, Bull. Jard. Bot. Btzg. 3, 12 (1932)
331; Kalkman l.c. (1954) 499; Leenhouts l.c. (1956) 218; Burgess l.c. 62; Kochummen l.c. (1972) 154;
Cockburn l.c. 49; Anderson l.c. 158; Wong l.c. 248; Ng l.c. 38; Whitmore, Tantra & Sutisna l.c. 39.
Dioecious trees. Pith of branchlets without vascular strands. Stipules absent. Leaflets with
entire margin, strongly asymmetrical at base; stalk not swollen. Inflorescences axillary
panicles. Flowers unisexual, 5-merous; sepals and petals free; stamens 5, episepalous, base
of filaments adnate to disc; disc extrastaminal, 5-lobed, lobes emarginate; ovary triangular,
3-celled, in male flowers entirely reduced. Fruits 3-winged, dry, woody capsule, splitting
into 3 valves. Seeds 3, broadly winged; germination epigeal; cotyledons shallowly 5-lobed,
leafy; leaves of young seedling alternate to spiral, simple for the first few nodes, later with 3
and 5 leaflets, margin initially toothed becoming entire later.
Distribution. One species confined to W Malesia.
Uses. The timber is commonly grouped with other members of the family and sold as
kedondong. The wood is light hardwood, and used for indoor construction. The resin is
sometimes used for torches.
99
Triomma malaccensis Hook. f.
Fig. 8.
(of Malacca)
l.c. (1860) 171; King l.c. 236; Ridley l.c. (1922) 369; Merrill l.c. (1929) 119; H.J. Lam, Bull. Jard.
Bot. Btzg. 3, 12 (1932) 332; Kalkman l.c. (1954) 499; Leenhouts l.c. (1956) 218; Burgess l.c. 62;
Type: Maingay 299, Malacca (holotype K; isotype L).
Tree to 60 m tall, 100 cm diameter; buttresses to 5 m high, spreading. Bark smooth to scaly,
grey-brown; inner bark pink or reddish brown, with a strong mango smell. Sapwood pale
yellow. Twigs dark brown. Leaves with 25 pairs of leaflets; petiolules of leaflets not
swollen; blade withering yellow with pink midrib and veins, oblong or ovate, 412 x 28
cm; base strongly unequal with one half rounded, apex pointed; midrib raised above; lateral
veins 611 pairs, raised on both surfaces; intercostal veins reticulate, distinct on both
surfaces. Inflorescences many-branched, hairy. Flowers: sepals and petals densely
tomentose. Fruits 57 x 22.5 cm. Seeds 3 with broad membranous wing. Saplings with
densely hairy rachis and leaflets.
Vernacular name. Sabahkedondong asam (Malay).
Distribution. Sumatra, Peninsular Malaysia, Borneo. In Sabah a common emergent tree of
the lowland forests, especially by streams. Widespread but not common in Sarawak, known
from only two collections from Lambir National Park (S. 46465) and Limbang district (S.
42833). Also occurs in Kalimantan.
Ecology. Mixed dipterocarp forest on yellow sandy clay soils. Flowering in AprilSeptember
and fruiting in MayDecember.
100
CAPPARACEAE
Deborah Kennard
Department of Botany,
University of Florida, U.S.A.
Merrill, EB (1921) 280; Masamune, EPB (1940) 318; Jacobs, FM 1, 6 (1960) 61, FM 1, 7 (1976) 822;
Whitmore, TFM 2 (1973) 25; Anderson, CLTS (1980) 158; Ashton, MNDTS 2 (1988) 83; Corner,
WSTM 1 (1988) 201; Whitmore, Tantra & Sutisna, CLK 1 (1989) 40.
Trees, shrubs, woody climbers, or herbs. Leaves simple and entire, or trifoliolate or deeply
57-lobed, alternate or spirally arranged, petioled; stipules thorny, minute or absent.
Inflorescences racemose or paniculate, terminal or lateral, sometimes individual flowers
axillary or serial; bracts, if present, small and caducous, rarely with stipular bracteoles.
Flowers bisexual, regular or slightly irregular, sometimes big and showy, mostly in bud
until anthesis but in Crateva opening at a very early stage; sepals 4, either equal or in two
whorls of 2, free; petals 4 or (in Stixis) absent, equal or sometimes 2 of the petals slightly
asymmetric and joined at the base, often free, usually stalked; receptacle more or less
conical; stamens (4)6 to many, all fertile, either free or their bases connate with the stalk of
the ovary (gynophore) and forming an androgynophore; anthers dorsifixed; ovary superior, 1
6-loculed, generally on a long gynophore, sometimes sessile, ovoid to cylindrical, with a
small, simple sessile stigma. Fruit a leathery berry or a capsule. Seeds many, rarely 1
(Stixis), mostly kidney-shaped; endosperm scanty.
Distribution. Approximately 45 genera and 700 species in tropical and subtropical regions.
In Sabah and Sarawak represented by 4 genera; Crateva with 2 species of trees, the other 3
genera are mainly shrubs, woody climbers, and herbs.
Ecology. Mostly in open places in the lowlands, a few ascend to 1000 m. Although species
of the Capparaceae are usually found in dry regions, both Bornean species of Crateva are
riverine trees. The flower of this family is characterised by long-radiating stamens,
suggesting pollination by butterflies and moths. Several species are known to be nightflowering. Birds, bats, and ants have all been cited as dispersal agents. Trees of Crateva,
however, have water-dispersed fruits.
Uses. The familiar capers are from Capparis spinosa originating in the Mediterranean
region. Several species are noted for their medicinal properties (cf. Perry, MPESA (1980)
6870). Some tree species of Crateva are deemed to have religious significance and are
planted for ornamental purposes as well.
Taxonomy. The link between Capparaceae and Brassicaceae (the cabbage family) has long
been recognised. The distinct mustard-like taste common in Brassicaceae is also present in
many Capparaceae, e.g. in the capers. Recent cladistic studies (Judd et al., Harvard Papers
101
in Botany 5 (1994) 151) based on morphological and genetic data strongly suggest that the
Capparaceae is paraphyletic, and the subfam. Cleomoideae is more closely related to the
Brassicaceae than to the Capparaceae.
Key to genera
1.
Trees. Leaves trifoliolate...................................................................................Crateva

Shrubs, woody climbers, or herbs. Leaves simple or deeply 37-lobed...........................2
2.
Herbs. Leaves palmately, deeply 37-lobed. Fruit a long capsule....................................

Cleome L.
Gen. Pl. ed. 5 (1754) 302; Jacobs l.c. (1960) 99; Whitmore l.c. 25.
A pantropical and subtropical genus with over 150 species, well represented in
America; 4 species in Sabah and Sarawak.
Annual or perennial herbs. Flowers in leafy, terminal racemes or panicles; sepals 4,
free; petals clawed, normally 4; stamens 6 to many. Capsule linear and cylindrical.
Seeds many.
Shrubs or woody climbers. Leaves simple. Fruit a leathery berry .................................3
3.
Plants with stipular thorns. Flowers on long stalks. Fruit with one to many seeds..........
Capparis Tourn. ex L.
Gen. Pl. ed. 5 (1753) 222; Jacobs l.c. (1960) 70, Blumea 12, 3 (1965) 405; Whitmore l.c. 25.
About 250 species in the tropics and subtropics, half of which are in the New
World; 7 species in Sabah and Sarawak.
Shrubs, woody climbers, or rarely small trees. Leaves simple, spiral, with a pair of
stipular thorns which are occasionally wanting. Flowers single or in small, sessile
groups just above the leaf axils, feathery with long stamens; sepals biseriate,
mostly imbricate; petals 4; stamens many, rarely (7)8(12), free, radiating, longer
than the petals; ovary 1-locular. Berry globular to ellipsoid with a corky or leathery
pericarp. Seeds 1 to many, kidney-shaped.
Plants without thorns. Flowers on short stalks. Fruit with 1 seed ...................................
Stixis Lour.
Fl. Coch. (1790) 295, ed. Willdenow (1793) 361; Jacobs l.c. (1960) 95, Blumea 12 (1964) 5;
Whitmore l.c. 25.
7 species distributed throughout India, Burma, Cambodia, Hainan, and W Malesia.

One species endemic to Borneo, S. ovata, occurs in Sabah and Sarawak.
Rather small unarmed, woody climbers. Branches lenticellate. Leaves simple,
rather large. Inflorescences racemose or paniculate. Flowers many, axillary or
terminal; petals absent; sepals mostly 6, in two whorls of 3; stamens 2050(100),
about as long as sepals; ovary subglobular. Fruits few, on a thick woody stalk,
ellipsoid, 2.55 cm long. Seed one, large.
102
CAPPARACEAE (KENNARD)
CRATEVA L.
(Kratevas, 13263 B.C., an ancient Greek medicine man or root gatherer)
l.c. (1754) 203; Jacobs l.c. (1960) 63, l.c. (1964) 186; Whitmore l.c. 26; Anderson l.c. 158; Corner l.c.
203; Whitmore, Tantra & Sutisna l.c. 40.
Small to medium-sized trees. Bark grey or brown, smooth. Twigs round, glabrous, with
distinct leaf-scars, stipules small, not persistent; axillary buds small. Leaves spiral,
palmately lobed or trifoliolate, sessile to shortly stalked, base of stalks bearing small
appendages to 1 mm. Racemes terminal, corymbiform, either with arrested growth or
growing through and developing into a leafy twig with lateral flowers; pedicels leaving
distinct scars on the rachis. Flowers large and showy, to 5 cm across, opening while green
and young, subtended by bracts; sepals 4, clawed, green; petals 4, clawed, first white then
cream-coloured; stamens (8)1230, filaments at the very base connate with the gynophore,
57 cm, thread-like, pinkish violet; gynophore approximately as long as the stamens;
receptacle dish-shaped, floral parts not persistent. Fruits large, round or oblong berries,
with tough, sometimes papillate skin, hanging on a thickened woody pedicel and
androgynophore, together to 14 cm long; torus leaving a distinct scar. Seeds numerous,
heart- or horseshoe-shaped, embedded in pulp.
Distribution. About 6 species distributed throughout the tropics, except in Australia and
New Caledonia. Two species in Sabah and Sarawak.
Ecology. Found mostly in periodically flooded lowland forest near rivers, usually below
700 m. Fruits are dispersed by water, possibly also by fish.
Key to Crateva species

Leaflets with 1015 pairs of lateral veins, paler beneath than above..................1. C. magna
Leaflets with 511 pairs of lateral veins, same colour beneath as above..........2. C. religiosa
1. Crateva magna (Lour.) DC.
Fig. 1AC.
(Latin, magnus = great, large; referring to the stature)

Prod. 1 (1824) 243; Merrill, Comm. Lour. (1935) 172; Masamune l.c. 318; Jacobs l.c. (1964) 206, l.c.
(1976) 822; Whitmore l.c. 26; Corner l.c. 203; Whitmore, Tantra & Sutisna l.c. 40. Basionym:
Capparis magna Lour., Fl. Cochinch. 1 (1790) 331. Type: Loureiro, s.n., Indochina (BM).
Synonyms: Crateva nurvala Buch.-Ham., Trans. Linn. Soc. 15 (1827) 121; C. lophosperma Kurz, J.
Bot. 12 (1874) 195, t. 147, f. 46.
Tree to 20 m tall, 40 cm diameter. Twigs terete, lenticellate. Petiole 311 cm. Leaflets
firmly herbaceous to slightly coriaceous, paler beneath, mostly lanceolate; apex gradually
acuminate with an acute tip; lateral leaflets symmetrical, 817 x 26 cm, usually subsessile,
sometimes with stalk 36 mm long; lateral veins 1015 pairs, prominent beneath.
Inflorescences terminal on leafy twigs to 12 cm, bearing 20 to 100 flowers; pedicels 47 cm
long; petals 4, ovate-spathulate with a 512 mm stalk, lobes 1530 x 1522 mm; sepals 4,
ovate, 23.5 x 11.5 mm; stamens 1525, 3.54.5 cm long, filaments purple; ovary oblong
to cylindrical, gynophore 3.44.5 cm. Fruits ellipsoid, sometimes globose, c. 5 x 4 cm;
103
pericarp covered with a yellow-greyish crust which eventually breaks off. Seeds horseshoeshaped, 69 mm across, dull dark-brown.
Vernacular names. Sabahmempulak (Murut), pangos (Dusun Kinabatangan). Sarawak
serang (Iban, in confusion with Melicope). Bruneikebuan (Iban). Kalimantansasgah,
sebelu, tigarun (Iban).
Distribution. India, Burma, China, Hainan, Indo-China, Thailand, Sumatra, Peninsular
Malaysia, Java and Borneo. In Sabah, recorded from the Kalabakan, Lahad Datu, Lamas,
Sandakan and Telupid districts. In Sarawak from the 3rd, 4th and 7th Div., in all cases near
limestone or basalt exposures. Also in Brunei and Kalimantan.
Ecology. Found along streams and rivers or the edge of swamps; rarely found on ridges
although it can occur up to 1000 m. Flowering and fruiting irregular.
Uses. In Sabah and Sarawak, poultices made from the root, leaf, or bark are used externally
for sakit angin (wind), poultices made from the bark alone are used externally to reduce
high fever. The wood is sometimes used as timber. Elsewhere, the juice from the bark has
been used to increase the appetite and as a laxative. The fruits are used in Thailand as fish
bait.
2. Crateva religiosa G. Forst.
Fig. 1D.
(alluding to its common cultivation around temples in Tahiti)

Pl. Escul. Ins. Oc. Austral. (1786) 45; Masamune l.c. 318; Jacobs l.c. (1960) 65, l.c. (1964) 191;
Whitmore l.c. 26; Anderson l.c. 158; Corner l.c. 203; Ashton l.c. 84; Whitmore, Tantra & Sutisna l.c.
40. Type: G. Forster, s.n., Pacific Society Isl. (K). Synonyms: C. membranifolia Miq., Fl. Ind. Bat.,
Suppl. (1861) 387; C. brownii Korth. ex Miq., Illustr. (1870) 21; C. macrocarpa Kurz l.c. 195, t. 148,
f. 810.
Shrub or tree, to 30 m tall and 30 cm diameter. Twigs with many large lenticels. Petiole (3
)610 cm. Leaflets thin herbaceous, oblong to ovate, top rather abruptly acuminate, (5)8
16(27) x (3)410 cm, mostly subsessile, sometimes stalked to 5 mm, concolorous when
dried; lateral veins 711 pairs. Flowers few to 25, toward the end of leafy twigs which do
not grow through; pedicels 29 cm; sepals ovate, 47 x 13 mm; petals 4, ovate, upper pair
23 x 12 mm, lower pair smaller, 520 mm stalked; sepals ovate, 47 x 13 mm; stamens
1030, to 11 cm long; ovary subcylindrical to ovoid. Fruits subglobose to ovoid, 620 x 5
9 cm; pericarp papillate. Seeds embedded in cream-coloured pulp, somewhat heart-shaped,
1019 x 517 mm.
Distribution. India, Burma, China, Indo-China, Thailand, Sumatra, Peninsular Malaysia,
Java and Borneo. In Sabah, recorded in the Kudat, Sandakan and Kinabatangan districts,
and once on a rocky beach on Pulau Tiga off the southwest coast. In Sarawak, collections
are known from Sibu district in the 3rd Div., near limestone.
104
CAPPARACEAE (KENNARD)
105
U
Ecology. Common in periodically inundated forest along rivers; rarely in secondary or
primary dryland forest. Mostly found below 100 m but occasionally also at about 700 m.
Flowering and fruiting irregular.
Uses. Occult power has been ascribed to C. religiosa in India and Polynesia where it is
planted around temples. In the Solomon Islands the leaves are used for ear-ache and the
fruit is used to relieve constipation. In Borneo, C. religiosa is occasionally planted for
ornamental purposes, and the fruits are also used for fish bait.
106
CELASTRACEAE
K.M. Kochummen
Kepong, Malaysia
Merrill, EB (1921) 354; Ridley, FMP 1 (1922) 443; Masamune, EPB (1942) 417; Browne, FTSB
(1955) 76; Ding Hou, FM 1, 6 (1962) 227, FM 1, 6 (1964) 389, FM 1, 6 (1972) 930, Blumea 17 (1969)
97; Backer & Bakhuizen f., FJ 2 (1965) 53; Smythies, CST (1965) 38; Burgess, TBS (1966) 73;
Kochummen & Whitmore, TFM 1 (1972) 156; Cockburn, TS 1 (1973) 51; Anderson, CLTS (1980)
159; Wong, DMT (1982) 33; Corner, WSTM 1 (1988) 212; Ashton, MNDT 2 (1988) 87; Whitmore,
Tantra & Sutisna, CLK 1 (1989) 42; Ng, MFR 34 (1991) 41.
Trees, erect or scandent shrubs or woody climbers; bole sometimes buttressed, rarely with
pneumatophores. Leaves simple, decussate, opposite, sub-opposite, spiral or rarely alternate,
sometimes black-dotted especially beneath, margin faintly toothed, wavy or entire. Stipules
small, falling off early or absent. Incipient bracts absent except in Microtropis. Inflorescences
axillary and/or terminal, sometimes extra-axillary or borne on the branches, thyrsoid,
paniculate, racemose, cymose or fasciculate, usually with bracts. Flowers regular, bisexual
or unisexual (plants dioecious, rarely polygamous); calyx 45-lobed, lobes imbricate, rarely
valvate, persistent; petals 4 or 5, imbricate, contorted or rarely valvate, falling off early,
rarely persistent, free or united at base, sometimes united with the staminal ring, always
larger than calyx-lobes except in Perrottetia; stamens 25, alternate with petals, filaments
inserted on or on the inner side of the disc or on a basal ring (Microtropis), anthers 2-celled,
longitudinally or laterally dehiscent, introrse or extrorse; disc often conspicuous, fleshy or
membranous, cup-shaped, saucer-shaped or flat, entire, toothed, angled or lobed,
extrastaminal or intrastaminal, rarely absent (Microtropis); ovary superior or rarely semiinferior, partly or fully immersed in the disc, usually glabrous, sometimes with a tuft of
hairs at top (Bhesa), rarely with papilla-like or fleshy subulate processes at the base
(Euonymus), (1)25-celled, rarely many-celled (Siphonodon), very rarely hollow at the top
(Siphonodon), style distinct, or obscure or lacking, simple or divided to the base (Bhesa),
stigmas simple or lobed; ovules usually 2 in each cell, sometimes 1 or 318, anatropous.
Fruits drupes, capsules, or berries, surface smooth or spiny. Seeds erect or pendulous,
sometimes winged, aril present or absent, when present usually partly or entirely enveloping the seed or forming a cushion-like structure situated at the base of seed or with long,
filamentous appendages (Sarawakodendron), often orange or orange-red, rarely white;
endosperm present or absent; cotyledons flat, foliaceous.
Distribution. About 90 genera and over 1000 species distributed in both hemispheres,
predominant in the tropics and subtropics. In Malesia, 18 genera and c. 115 species, of
which 15 genera and 70 species are recorded in Sabah and Sarawak.
Ecology. From lowland including mangrove and peat swamp, to montane forests to about
3200 m. Most members of the family have nectariferous disc in the flowers which are
visited by various kinds of insect-pollinators. Seeds of species having brightly coloured arils
107
are dispersed by birds; the winged seeds of Kokoona, Lophopetalum and other taxa are
dispersed by wind. The fruits of the mangrove tree, Cassine viburnifolia, have a corky
mesocarp that facilitates dispersal by sea water.
Uses. Timbers of the genera Kokoona, Lophopetalum, and Siphonodon have commercial
value. Some species contain alkaloids. Aborigines in Peninsular Malaysia make use of the
bark of Lophopetalum pallidum in the preparation of dart poison. The ochre yellow middle
bark of some Lophopetalum and Kokoona species burn readily and is useful tinder in the
forest.
Key to genera
1.
Woody climbers or scandent shrubs.................................................................................2

Trees, or erect shrubs.............................................................................................................6
2.
Leaves spirally arranged........................................................................................................3

Leaves opposite, decussate, or rarely spiral.........................................................................4
3.
Woody climber. Ovary free from the disc.......................................................................

Celastrus L.
Gen. Pl. ed. 5 (1754) 91; Ridley l.c. 450; Ding Hou l.c. (1963) 233.
About 30 species, tropics and subtropics; 2 species in Sabah and Sarawak.

Leaves spirally arranged, margin toothed. Inflorescences cymose or paniculate.
Flowers unisexual, 5-merous, pedicel articulated; calyx bell-shaped; disc membranous
or fleshy, cup-shaped; ovary superior, 3-celled; style columnar, stigma 3-lobed;
ovules 2 in each cell. Fruit a 3-valved capsule, tipped by persistent style. Seeds 1
6, enveloped by crimson aril.
Scandent shrubs. Ovary partially immersed in the disc..................................................
Maytenus Mollina
Saggio Chile (1782) 177; Ding Hou l.c. (1962) 238.
About 225 Species; tropics and subtropics of both the Old and New World. In
Malesia 5 species, one of which, M. emarginatus (Willd.) Ding Hou, is present in
Sabah; uncommon, known only by a single collection (Sugau 34) from Balambangan
island.
Scandent shrubs with spines on leafy shoots. Leaves spirally arranged, margin
toothed. Flowers in fascicles, unisexual, 45-merous; calyx-lobes deltoid; petals
obovate or oblong, entire; disc rounded; stamens inserted on the margin of the disc;
ovary partly immersed in the disc, 3-celled, style cylindric, stigma 3. Fruits a
capsule. Seeds ellipsoid, red, subtended by white fleshy aril at the base.
4.
Flowers with distinct disc......................................................................................................5

Flowers without or with inconspicuous disc...................................................................
Reissantia Hall
Mus. Hist. Nat. Paris 30 (1958) 466; Ding Hou l.c. (1964) 400.
7 species, Old World tropics; 4 species in Sabah and Sarawak.

Leaves decussate, rarely subopposite. Inflorescences axillary. Flowers bisexual; calyxlobes 5, imbricate; petals 5, imbricate; disc inconspicuous; stamens 3; ovary semi108
CELASTRACEAE (KOCHUMMEN)
superior, 3-celled; style short; stigma obscure; ovules 2 (rarely 48) in each cell.
Fruits capsular consisting of 3 divergent separate follicles which split into 2 valves.
Seeds with basal more or less transparent membranous wing.
5.
Fruits capsular with 3 follicles. Seeds winged. Petals with inflexed tip...........................
Loeseneriella A.C. Smith
Am. J. Bot. 28 (1941) 438; Ding Hou l.c. (1964) 397.
26 species, Tropical Africa, Asia, Malesia, New Hebrides. In Malesia, 4 species of

which 2 are present in Sabah and Sarawak.
Leaves decussate. Inflorescences axillary, cymose. Flowers bisexual; calyx deeply
5-lobed; petals 5, thick with inflexed tip; disc fleshy; stamens 3; ovary superior or
semi-inferior. Fruits capsular, with 3 divergent follicles, each dehiscing into 2
valves. Seeds usually with basal membranous wing.
Fruits drupaceous. Seeds embedded in pulp. Petals without inflexed tip.........................
Salacia L.
Mant. (1767) 159; Ridley l.c. (1922) 456; Ding Hou l.c. (1964) 404.
About 150 species, pantropical; 33 species in Malesia of which 17 are found in

Sabah and Sarawak. Leaves decussate, subopposite or rarely spiral. Flowers bisexual, axillary, in clusters or organised in cymes or panicles; calyx deeply 5-lobed
or 35-lobed in apical part, and circumscissile at the base or lengthwise splitting or
not lobed; petals usually 5, rarely 4 or 7; disc fleshy, ring-like, truncate-conical or
flattened, occasionally cup-like; stamens 3, rarely 2; ovary semi-inferior, 3-celled,
rarely 2-celled, ovules 28 in each cell; style distinct or obscure; stigma obscure.
Fruits drupaceous, 13-celled. Seeds 1 to several, embedded in mucilaginous pulp.
6.
Leaves alternate or spirally arranged............................................................................7

Leaves decussate, opposite, subopposite, or very rarely alternate or spiral............10
7.
Leaf margin toothed.....................................................................................................8

Leaf margin entire................................................................................................................9
8.
Petals usually similar in size to calyx-lobes. Ovary not hollow in apical part...8.
Perrottetia
Petals always larger than calyx-lobes. Ovary hollow in apical part.........10. Siphonodon
9.
Leaves and twigs with pustules; petiole to 1.5 cm long, not swollen towards apex;
stipules not prominent, leaving no stipular scars on twigs.............9. Sarawakodendron
Leaves and twigs without pustules; petioles longer than 1.5 cm, swollen at the apex or
both ends; stipules prominent, leaving scars on twigs.......................................1. Bhesa
10. Incipient bracts prominent. Disc absent..................................................7. Microtropis

Incipient bracts absent. Disc present...............................................................................11
11. Trees confined to tidal rivers and mangroves. Ovary 2-celled........................2. Cassine
Trees or shrubs of inland forests. Ovary 3 or 45-celled.............................................12
109
110
12. Ovary 45-celled. Seeds not winged...........................................................................13

Ovary 3-celled. Seeds winged.....................................................................................14
13. Ovules 2 in each cell.................................................................................3. Euonymus
Ovules 1 in each cell.................................................................................4. Glyptopetalum
14. Petals twisted and overlapping. Wing attached to the apex of seed.........5. Kokoona
Petals imbricate. Wing surrounding seed..............................................6. Lophopetalum
1. BHESA Buch.-Ham. ex Arn.

(origin unknown, probably an English plant name)
Edin. New Phil. J. (1834) 315; Ding Hou, Blumea Suppl. 4 (1958) 149, l.c. (1962) 280; Kochummen
& Whitmore l.c. 158; Cockburn l.c. 54; Anderson l.c. 159; Wong l.c. 22; Corner l.c. 213; Ashton l.c.
89; Whitmore, Tantra & Sutisna l.c. 42; Ng l.c. 43. Synonym: Kurrimia Wall. ex Arn., Nov. Act. Ac.
Caes. Leop.-Car. 18 (1836) 328.
Medium-sized to tall trees; bole with buttresses, often fluted at base. Bark grey-brown,
smooth to cracking and scaly; inner bark mottled cream and orange. Sapwood pale yellow.
Twigs smooth, with distinct stipular scars. Stipules prominent, lanceolate, soon falling off
and leaving scars on the twigs. Leaves spiral, entire, with silky sheen; intercostal veins fine,
scalariform; petioles slender, longer than 1.5 cm, swollen at both ends or at the apex only.
Inflorescences axillary, solitary or paired, paniculate or racemose; bracts falling off early;
pedicels jointed. Flowers bisexual, 5-merous (rarely 4-merous); calyx deeply lobed, lobes
imbricate or valvate; petals twisted; disc fleshy, entire or lobed; stamens 45, seated on or
just below the disc; ovary superior, usually with terminal tuft of hairs, 2-celled, with 2
ovules in each cell, styles 2. Fruit a capsule, entire or 2-lobed, splitting into 2 valves or on
one side only. Seeds 12 per cell, with bright orange-red or pink basal aril; endosperm
copious; germination epigeal.
Distribution. 5 species; Sri Lanka and Malesia; 2 species in Sabah and Sarawak.
Ecology. Widely distributed from lowland to submontane forests to 1500 m.
Key to Bhesa species

Stipules to 3 cm long. Petioles strongly swollen at both ends. Intercostal veins distinct.
Inflorescences usually paniculate. Disc deeply 5-lobed. Fruits two-lobed.....1. B. paniculata
Stipules to 1 cm long. Petioles only slightly swollen at apex. Intercostal veins faint to
invisible. Inflorescence racemose. Disc entire or obscurely notched. Fruits not lobed, with
pointed tip.......................................................................................................2. B. robusta
111
1. Bhesa paniculata Arn.
Fig. 1.
(Latin, paniculatus = having a loose, branched flower-cluster or inflorescence)

l.c. (1834) 315; Ding Hou l.c. (1958) 151, l.c. (1962) 282; Burgess l.c. 73; Kochummen & Whitmore
l.c. 160; Cockburn l.c. 55; Anderson l.c. 159; Corner l.c. 213; Ashton l.c. 92; Whitmore, Tantra &
Sutisna l.c. 42. Type: Wallich no. 4336, Penang (BO, K). Synonyms: Kurrimia paniculata Wall. ex
Arn. l.c. 328; K. luzonica Vidal, Rev. Pl. Vasc. Filip. (1886) 88; K. minor Ridl., Kew Bull. (1938) 235.
Medium-sized tree to 35 m tall, 50 cm diameter; bole often fluted at base. Bark grey-brown,
smooth to cracking; inner bark orange-yellow. Sapwood pale white. Twigs pale brown to
dark brown, 510 mm thick, with prominent stipular and leaf scars. Stipules lanceolate, c. 3
cm long. Leaves elliptic to oblong, rarely obovate, 5.527 x 2.213 cm; base cuneate or
rounded, apex pointed or blunt; midrib flattened or raised above; lateral veins 520 pairs,
prominently raised below, visible above; intercostal veins scalariform, very close, faint
above, distinct beneath; petioles 19.5 cm long, strongly swollen at both ends.
Inflorescences panicles, 1037 cm long or racemes to 20 cm long, from axils of upper
leaves. Flowers greenish yellow or dark purplish red, pedicels 23 mm long; calyx-lobes
hairy outside; disc deeply 5-lobed; petals oblong or ovate, hairy inside; stamens 1.52 mm
long, anthers triangular; ovary ellipsoid, styles free, about half as long as ovary. Fruits 2lobed, red when fresh, drying dark brown, 12 cm long, with the biggest lobe c. 0.5 cm
wide. Seeds 24, more than half covered with pink aril; cotyledons leafy.
Vernacular names. Sabahbiku-biku (Malay). Sarawaksimun (Iban). Bruneiserunai
(Malay).
Distribution. S India, S Thailand, Sumatra, Peninsular Malaysia, Borneo and the
Philippines. Common and widely distributed in Sabah and Sarawak. Also in Brunei and
Kalimantan.
Ecology. Lowland, including mixed dipterocarp, heath, and peat swamp, to submontane
forests to 1500 m. Submontane samples have comparatively smaller and thicker leaves.
Flowering in MarchMay and AugustOctober, fruiting in AprilDecember.
2. Bhesa robusta (Roxb.) Ding Hou

(Latin, robustus = strong-growing, robust; the habit)
l.c. (1958) 152, l.c. (1962) 283; Kochummen & Whitmore l.c. 161; Anderson l.c. 159; Corner l.c. 214;
Ashton l.c. 93; Whitmore, Tantra & Sutisna l.c. 42. Basionym: Celastrus robustus Roxb., Fl. Ind. 2
(1824) 395. Type: Roxburgh, Icones no. 2185 (K). Synonyms: Kurrimia pulcherrima Wall. ex Laws.
in Hooker f., Fl. Br. Ind. 1 (1875) 622; Kurrimia maingayi Laws. in Hooker f. l.c. 622.
Medium-sized tree to 18 m tall and 40 cm diameter. Bark grey-brown; inner bark yellow.
Sapwood pale. Stipules lanceolate, 510 mm long. Leaves elliptic or oblong, 79 x 34 cm;
base cuneate, apex blunt or pointed; midrib raised above; lateral veins 1115 pairs;
intercostal veins very faint to invisible; petioles 13 cm long, slightly swollen at apex.
Flowers in racemes, subsessile; calyx lobed, lobes broadly ovate to rounded; petals oblong
to elliptic; disc cup-shaped, subentire or obscurely notched; stamens c. 2 mm long, attached
beneath the outer margin of disc, anthers deltoid; ovary subglobose with tuft of hairs at
apex, styles free, longer than ovary. Fruits ovoid, not lobed, c. 2 x 1 cm, apex pointed.
Seeds 1, enveloped by aril, sometimes only the lower half; cotyledons fleshy.
112
Distribution. NE India, Bhutan, Chittagong, Burma, Andaman Islands, Thailand, Indo-China,

Sumatra, Peninsular Malaysia, and Borneo. In Sabah and Sarawak, uncommon; in Sabah,
known from 3 collections (SAN A 6613, SAN 50057, and SAN 76690) from Beaufort and
Sandakan districts, and in Sarawak by one collection (Beccari 2624).
Ecology. Mixed dipterocarp forests at 300600 m, on clay-rich soils.
Uses. Though of no commercial importance in Sabah and Sarawak, the timber is used in
house-building in other countries where it occurs.
2. CASSINE L.
(origin and meaning unknown)
Gen. Pl. (1737) 338; Ding Hou l.c. (1962) 284; Backer & Bakhuizen f. l.c. 55; Kochummen &
Whitmore l.c. 161; Anderson l.c. 159; Kostermans, Gard. Bull. Sing. 39 (1986) 177; Ashton l.c. 93.
Synonym: Elaeodendron Jacq. f. ex Jacq., Ic. Pl. Rar. 12 (1782) t. 48.
Shrubs or small trees. Leaves decussate. Inflorescences axillary or extra-axillary cymes,

with distinct peduncles. Flowers bisexual, 45-merous; calyx-lobes imbricate; petals free,
imbricate; stamens 45, inserted on the disc or on its outer margin, filaments subulate,
anthers introrse; disc prominent, fleshy, flat, orbicular or lobed; ovary semi-inferior, 2celled, conical or flask-like, the base slightly united with the disc or partly immersed in it;
style very short or obscure, stigma obscure or 2-lobed; ovules 2 in each cell, erect, attached
at the base. Fruit a drupe, 12-celled. Seeds 12, not winged, without aril, with endosperm.
Distribution. About 80 species throughout the tropics, mainly in Africa; 2 species in
Malesia of which only one is present in Sabah and Sarawak.
Ecology. In Sabah and Sarawak, confined to mangroves and banks of tidal rivers.
Cassine viburnifolia (Juss.) Ding Hou
Fig. 2.
(with leaves resembling those of Viburnum)

l.c. (1962) 286; Kochummen & Whitmore l.c. 161; Cockburn l.c. 54; Anderson l.c. 159; Ashton l.c.
93. Basionym: Aegiphila viburnifolia Juss., Ann. Mus. Hist. Nat. Paris 7 (1806) 76. Type: Sine coll.,
s.n., Philippines (P). Synonyms: Euonymus viburnifolius (Juss.) Merr., Philip. J. Sc. 9 (1914) Bot.
312; Elaeodendron subrotundum King, J. As. Soc. Beng. 65, 2 (1896) 356.
Shrub or small tree to 10 m tall, 20 cm diameter. Bark yellowish grey, smooth. Twigs black.
Leaves broadly obovate, 410 x 26 cm; base cuneate, margin curled inwards, remotely
minutely toothed, apex blunt to rounded; lateral veins 46 pairs, faint; intercostal veins
reticulate, equally prominent as the lateral ones; petioles 813 mm long. Inflores-cences
cymose, axillary. Flowers white, 4-merous; calyx-lobes broadly ovate to rounded, almost
free; petals oblong to ovate-oblong. Fruits obovoid, often rhomboid in cross-section;
mesocarp thick and corky. Seed 1, c. 6 x 3 mm, obovate to oblong.
113
Vernacular name. Sarawakbarat-barat (Malay).

Distribution. Andaman Islands, Thailand, Sumatra, Peninsular Malaysia, Borneo, Philippines,
and Celebes. In Sabah known from Kudat, Sandakan and Tawau districts, and in Sarawak
from the Rejang delta in 3rd. Div. Also in Brunei.
Ecology. Common on the banks of tidal rivers and mangrove channels near the inland limits
of salinity. The fruits are dispersed by water.
3. EUONYMUS Tourn. ex L.
(Greek, eu = good, onuma = name; a plant name)
Gen. Pl. ed. 5 (1754) 91; Ridley l.c. (1922) 445; Masamune l.c. 417; Blakelock, Kew Bull. (1951) 232;
Ding Hou l.c. (1962) 245; Backer & Bakhuizen f. l.c. 53; Kochummen & Whitmore l.c. 162; Cockburn
l.c. 55; Anderson l.c. 159; Ashton l.c. 94; Whitmore, Tantra & Sutisna l.c. 42.
Shrubs or small trees. Leaves opposite-decussate sometimes with dark spots on the under
surface; stipules lanceolate, falling off early. Inflorescences axillary, cymose; rarely flowers in
fascicles (E. javanicus); pedicels jointed. Flowers bisexual, 5- or 4-merous; calyx deeply
lobed, lobes imbricate, entire or minutely toothed; petals free, imbricate, spreading or
reflexed; disc prominent, fleshy or thin, flat, 5- or 4-angled or 5- or 4-lobed or rounded,
smooth or covered with fleshy papilla-like or subulate processes; stamens 45, inserted on
the disc, anthers 2-celled, lateral or introrse, dehiscence apical, filaments obscure or
distinct; ovary semi-inferior 45-celled, partly or wholly immersed in the disc, stigma
obscure or discoid, ovules mostly 2 in each cell. Fruits capsules, usually (3)45-angular or
lobed, smooth or armed with rigid prickles, apex obtuse, acute, truncate or concave. Seeds
usually black, not winged, partly or completely covered by orange aril.
Distribution. About 180 species, mainly in the tropics and subtropics. In Malesia, 12
species of which 5 are recorded in Sabah and Sarawak.
Ecology. In primary and secondary forests from lowlands to mountains to 3200 m.
Key to Euonymus species

(based on inflorescences and flowers)
1.
Flowers 5-merous. Leaves without black gland-dots below............................................2

Flowers 4-merous. Leaves with black gland-dots below................................................4
2.
Petals fimbriate................................................................................................................3
Petals entire.....................................................................................1. E. acuminifolius
3.
Peduncles distinct........................................................................3. E. cochinchinensis

Peduncles obscure..................................................................................5. E. javanicus
4.
Lateral veins of the leaves sunken above..........................................2. E. castaneifolius

Lateral veins of the leaves not sunken above.......................................4. E. glandulosus
114
115
116
Key to Euonymus species

(based on leaves and twigs)
1.
Leaf margin toothed.....................................................................................................2

Leaf margin entire (sometimes faintly toothed in E. cochinchinensis).............................3
2.
Leaf lower surface with scattered black gland-dots. Lateral veins sunken above.............
.......................................................................................................2. E. castaneifolius
Leaf lower surface without black gland-dots. Lateral veins not sunken above.................
........................................................................................................1. E. acuminifolius
3.
Twigs usually 4-angled. Leaf lower surface with scattered black gland-dots; margin
strongly recurved................................................................................4. E. glandulosus
Twigs not 4-angled. Leaf lower surface without scattered black gland-dots; margin not
strongly recurved........................................................................................................4
4.
Leaf margin entire. Peduncle obscure.......................................................5. E. javanicus

Leaf margin faintly toothed. Peduncle distinct..............................3. E. cochinchinensis
1. Euonymus acuminifolius Blakelock

(Latin, acuminatus = long-pointed, folius = leaves)
l.c. 253; Ding Hou l.c. (1962) 251; Whitmore, Tantra & Sutisna l.c. 43. Type: Bnnemeijer 581,
Sumatra, Ophir district, NW slopes of Mt. Talamau (BO). Synonym: E. acuminifolius Blakelock var.
borneensis Blakelock l.c. 253.
Shrub to 4 m tall. Twigs 4-angled. Leaves membranous to chartaceous, ovate-oblong to

lanceolate or elliptic to oblong, 710.5 x 24 cm, lower surface without black gland-dots;
base cuneate, margin toothed, apex pointed, tip c. 2 cm long; lateral veins 46 pairs, arching
near margin, not sunken above; petioles 26 mm long. Flowers in cymes, purplish red, 5merous; calyx-lobes rounded, inner two usually larger; petals almost rounded, entire; disc
obscurely 5-angular; stamens with very short filaments, anthers c. 0.5 mm long; ovary
faintly 5-angled, stigma discoid, obscurely 5-angled. Fruits obcordate, distinctly 5-lobed,
1.52 x 11.7 cm. Seeds with aril at base.
Distribution. Sumatra, Borneo and Celebes. In Sabah uncommon, known only from Mt.
Kinabalu (Clemens 30350, type of E. acuminifolius var. borneensis, and Clemens 34478).
Not recorded from Sarawak.
Ecology. Montane forest at about 2700 m.
117
2. Euonymus castaneifolius Ridl.
Fig. 3.
(Latin, castaneus = chestnut-coloured, folius = leaves)

Kew. Bull. (1931) 36; Masamune l.c. 417; Ding Hou l.c. (1962) 251; Cockburn l.c. 56; Anderson l.c.
159; Ashton l.c. 94; Whitmore, Tantra & Sutisna l.c. 43. Type: Haviland 827, Sarawak, Kuching
(holotype K; isotype BO). Synonym: E. moultonii Ridl l.c. (1931) 36, Masamune l.c. 417.
Small tree rarely reaching 12 m tall and 10 cm diameter. Bark grey-brown, smooth. Young
twigs angled. Leaves drying to greenish grey, lower surface with scattered black glanddots, elliptic to lanceolate, 717.5 x 35.5 cm; base cuneate, margin distantly faintly
toothed, apex pointed; midrib raised above; lateral veins 610 pairs, sunken above, faint or
rarely prominent below; intercostal veins invisible; petioles 1015 mm long, drying to pale
yellowish and wrinkled. Flowers in cymes, white, 4-merous; calyx-lobes rounded, inner pair
largest; petals with few longitudinal veins; stamens inserted on the disc, anthers slightly
depressed to oblong, ovary 4-angular towards the base. Fruits red on ripening, obovoid
with truncate apex, 4-lobed at apex with 4 ridges, 1018 x 710 mm, with persistent calyx.
Seeds with cup-shaped aril at base.
Distribution. Sumatra and Borneo. Widely distributed in Sabah and Sarawak; also known
in Kalimantan and Brunei.
Ecology. Lowland mixed dipterocarp forests on fertile clay-rich soils, near limestone and on
basic volcanic rock to montane forests to 2700 m. Flowering in FebruarySeptember and
fruiting in JanuaryMarch.
3. Euonymus cochinchinensis Pierre

(of Indo-China)
Fl. For. Coch. 4 (1894) t. 309 A; Blakelock l.c. 255; Ding Hou l.c. (1962) 248; Whitmore, Tantra &
Sutisna l.c. 43. Type: De Perry 4073, Cochinchina, Trans Province (holotype P; isotype K).
Synonyms: Glyptopetalum scortechinii King l.c. 345; Euonymus philippinensis Merr., Philip. J. Sc. 3
(1908) Bot. 238; E. pahangensis Ridl., FMP 5 (1925) 299.
Small tree to 12 m tall. Leaves chartaceous to subcoriaceous, elliptic, oblong or obovate,

4.516 x 2.57 cm, lower surface without black gland-dots; base cuneate to rounded,
margin entire or remotely toothed, apex pointed; lateral and intercostal veins obscure or
slightly elevated on both surfaces; petioles 38 mm. Inflorescences 310.5 cm long. Flowers
greenish yellow, 5-merous; calyx-lobes fimbriate; petals broadly obovate with fimbriate or
toothed margins; disc rounded and faintly 5-angled; stamen filaments flat, subulate; ovary
narrowed towards apex, stigma obscure. Fruits broadly obovoid to subglobose, concave at
apex, deeply 5-lobed. Seeds ellipsoid, 56 x 34 mm.
Distribution. Thailand, Indo-China, China (Hainan), Sumatra, Peninsular Malaysia, Borneo,
Philippines, Celebes, Lesser Sunda Is., Moluccas and New Guinea. Uncommon in Borneo,
known only by two collections (Merrill 9644 and Clemens 17445) from Sabah.
Ecology. Coastal forests.
118
4. Euonymus glandulosus (Merr.) Ding Hou

(Latin, glandulosus = gland-bearing; black gland-dots on the leaf)
l.c.(1962) 251; Cockburn l.c. 56; Whitmore, Tantra & Sutisna l.c. 43. Basionym: Glyptopetalum
glandulosum Merr., Philip. J. Sc. 12 (1917) Bot. 279. Type: Merrill 9547, Philippines, Palawan
(holotype UC; isotypes BO, K).
Small tree to 5 m tall. Twigs terete to 4-angled. Leaves papery to leathery, elliptic to
lanceolate or obovate, 4.512.5 x 24.5 cm, with scattered black gland-dots below; base
cuneate, margin strongly recurved, almost entire, apex pointed; midrib raised above; lateral
veins 45 pairs, very faint to obscure; intercostal veins invisible; petioles 58 mm long.
Flowers white or dark purple, 4-merous; calyx-lobes almost rounded with faintly
irregularly toothed margin, inner lobes thinner, larger and transparent; petals rounded to
broadly obovate, with few longitudinal veins; stamens with very short filaments; disc
obscurely 4-angled; ovary slightly 4-angled towards base. Fruits broad-obovoid, 1018 x
1015 mm, 4-lobed, concave at apex, smooth, yellow, ripening red. Seeds ellipsoid with
disk-like aril at base.
Distribution. Borneo and Philippines. In Sabah, restricted to Mt. Kinabalu Park, but in
Sarawak widely distributed.
Ecology. Hill and montane forests at 9002700 m.
Ding Hou (l.c.) stated that the flowers are dark purplish but SAN 37988 and SAN 87659
have greenish or whitish flowers. SAN 79981 from Mt. Kinabalu has thinner leaves with
strongly toothed margins and with reddish flowers.
5. Euonymus javanicus Blume

(of Java)
Bijdr. (1826) 1146; Ridley l.c. (1922) 445; Blakelock l.c. 257; Ding Hou l.c. (1962) 248; Backer &
Bakhuizen f. l.c. 53; Kochummen & Whitmore l.c. 162; Cockburn l.c. 56; Anderson l.c. 160; Ashton
l.c. 95; Whitmore, Tantra & Sutisna l.c. 43. Type: Blume, s.n., Java (holotype L; isotypes BO, K,
SING.). Synonyms: E. alatus Elmer, Leafl. Philip. Bot. 4 (1912) 1484; E. elmeri Merr. l.c. (1917)
281; E. coriaceus Ridl. l.c. (1925) 299; E. micropetalus Ridl. l.c. (1925) 299.
Small tree to 15 m tall and 20 cm diameter. Bark grey-brown, smooth; inner bark pinkish.
Sapwood yellowish. Twigs brownish. Leaves elliptic to oblanceolate, 715 x 36.5 cm,
lower surface without black gland-dots; base cuneate, margin entire, not recurved, apex
pointed; midrib raised above, pale yellow below on drying; lateral veins 57 pairs, very faint
on both surfaces; intercostal veins invisible; petioles 0.51 cm long. Flowers yellowish, in
clusters, 5-merous; peduncle obscure; calyx-lobes unequal, rounded; petals obovate or
rounded with fimbriate margins; disc 5-angled; stamens with triangular anthers; ovary
conical, style short, stigma obtuse. Fruits dark green, ripening red, broadly obovoid, 23.5
x 1.52.5 cm, with pointed tip and 5 well-developed ridges, stalk 1.52 cm long. Seeds
black.
119
120
Distribution. Nicobar Is., Burma, Thailand, Indo-China, Java, Borneo, Natuna Is., Philippines,
Celebes, Lesser Sunda Isl., Moluccas, and New Guinea. In Sabah, known from Lahad Datu,
Ranau, Tawau and Tenom districts. In Sarawak, uncommon, recorded from the Bukit
Numpang and Tai Ton forests.
Ecology. Lowland (including limestone) to montane forests to 2400 m.
4. GLYPTOPETALUM Thwaites
(Greek, glypto = incised, petalum = petals)
in Hooker, J. Bot. Kew Misc. 8 (1856) 267; Ridley l.c. (1922) 446; Ding Hou l.c. (1962) 254;
Kochummen & Whitmore l.c. 162; Anderson l.c. 160; Ashton l.c. 95.
Shrubs or small trees. Leaves opposite. Inflorescences cymose, axillary or extra-axillary.

Flowers bisexual, 4-merous; calyx spreading, inner pair larger; petals free, fleshy with
small appendage or depression inside; disc conspicuous, fleshy; stamens 4, inserted on the
disc, near base of ovary, anthers opening apically; ovary semi-inferior, immersed in the
disc, 4-celled, style obscure, stigma obscure, ovule one in each cell. Fruits capsules, when
splitting leaving a persistent central columella. Seeds with fleshy aril at lower half.
Distribution. c. 20 species; India, Sri Lanka, Burma to Hainan, and Malesia. 8 species are
present in Malesia; 2 in Sabah and Sarawak.
Taxonomy. The genus is closely allied to Euonymus from which it differs in having only
one ovule per ovary-cell.
Key to Glyptopetalum species

Leaf apex blunt or rounded. Twigs rounded............................................1. G. palawanense
Leaf apex pointed. Twigs sharply 4-angled..........................................2. G. quadrangulare
1. Glyptopetalum palawanense Merr.

(of Palawan)
Philip. J. Sc. 26 (1925) 466; Ding Hou l.c. (1962) 257. Type: Forestry Bureau 29181, Philippines,
Palawan (BO, K).
Small tree to 5 m tall. Twigs grey-green, rounded. Leaves leathery, drying to greenish
yellow, obovate or elliptic, 1014 x 58 cm; base cuneate, margin distantly toothed towards
the upper half, apex blunt or rounded; midrib raised above; lateral veins 68 pairs, almost
invisible below, faintly raised above; intercostal veins invisible; petioles 812 mm long.
Flowers unknown. Fruits globose to depressed globose, c. 15 x 8 mm, 14-celled.
Distribution. Borneo and Philippines (Palawan). Uncommon in Borneo, known only from a
sterile collection (Wong & Payne, s.n., 27 April 1993) from Balambangan Island, off Kudat
Peninsula, Sabah.
Ecology. Coastal limestone ridge.
121
2. Glyptopetalum quadrangulare Prain ex King
Fig. 4.
(Latin, quadrangularis = 4-angled; the twigs)

l.c. 345; Ridley l.c. (1922) 446; Ding Hou l.c. (1962) 257; Kochummen & Whitmore l.c. 162;
Anderson l.c. 160; Ashton l.c. 95. Type: King's collector 7106, Perak (lectotype K; isolectotype BO).
Shrub or small tree to 5 m tall, 5 cm diameter. Twigs strongly 4-winged. Leaves ellipticlanceolate, 930 x 314 cm; base cuneate to rounded, margin toothed from base to apex,
apex pointed; lateral veins 812 pairs, raised below, sunken above; intercostal veins
reticulate; petioles 510 mm long. Flowers greenish yellow, in 12 cm long cymes; calyx
almost divided to base, lobes reniform; petals suborbicular, fleshy, margin thinner, wavy;
disc flat; stamens inserted near the base of ovary; ovary pyramidal, style and stigma
obscure. Fruits c. 2 x 11.3 cm, depressed globose, sulcate, 34-celled.
Distribution. Burma, Sumatra, Peninsular Malaysia, and Borneo. In Sarawak uncommon,
known only by two collections (S. 16363 from Serian, and S. 12549 from Bekup); not yet
recorded from Sabah. Also known from Kalimantan.
Ecology. Lowland forests on limestone.
5. KOKOONA Thwaites
(kokoon = a Ceylonese plant name)
mata ulat (Malay), bajan (Iban)
in Hooker, J. Bot. Kew Misc. 5 (1853) 379; King l.c. 346; Merrill l.c. (1921) 354; Masamune l.c. 417;
van Steenis, Sarawak Mus. J. 8 (1958) 437; Ding Hou l.c. (1962) 258, l.c. (1969) 105; Balan Menon,
MF 17 (1964) 18; Backer & Bakhuizen f. l.c. 54; Smythies l.c. 38; Kochummen & Whitmore l.c. 163;
Jansen et al., Blumea 21 (1973) 153; Cockburn l.c. 56; Anderson l.c. 160; Wong l.c. 109; Ashton l.c.
97; Whitmore, Tantra & Sutisna l.c. 43; Ng l.c. 43; Kochummen, Sandakania 5 (1994) 51.
Small to very large trees; bole often with short buttresses. Bark grey to chocolate-brown,
often with horizontal rings, smooth, cracking or fissured, with large lenticels; middle bark
ochre or orange; inner bark pinkish or yellowish, fibrous. Sapwood white to yellow-brown
with prominent pale bands. Twigs flattened at nodes, drying black. Stipules small. Leaves
decussate, occasionally subopposite or alternate, margin entire, wavy or toothed. Flowers
bisexual, in axillary panicles or racemes, pedicels jointed; sepals 5; petals 5, free, overlapping and twisted (contorted); disc conspicuous, fleshy, cup-shaped, corrugated or 5lobed; stamens 5, inserted on the inner edge of the disc, filaments abruptly narrowed
towards the apex and transparent at the upper end, anthers usually with prominent
connective; ovary superior or semi-inferior, 3-celled, style obscure, stigma capitate, ovules
616 in each cell, in two rows down the central axis. Fruits capsules, 3-angled, 3-valved,
splitting loculicidally. Seeds overlapping, flat, with conspicuous membranous wing at the
apical end; endosperm absen; germination durian-type; cotyledons fleshy; seedlings with
opposite leaves, without or with fine hair-like, deciduous stipules.
122
Distribution. 10 species; Southern India, Sri Lanka, Burma, Malesia. 8 species in Malesia
of which 7 are present in Sabah and Sarawak.
Ecology. Found scattered in a wide range of inland forests including swamps to 1500 m.
Uses. The timber is a medium hardwood. It is easy to saw and cross-cut; suitable for heavy
construction if treated and can be used as posts, beams, joints and railway sleepers. Also
suitable for heavy-duty furniture, parquet-flooring, veneers, window and door-frames.
Taxonomy. The genera Kokoona and Lophopetalum are difficult to distinguish on vegetative
characters alone. However, some species of Kokoona have wavy to toothed leaf margins and
they dry with greenish tinge, while in Lophopetalum the leaves have entire margins and they
usually dry to a dark brown colour. The important diagnostic character is in the seed which
is surrounded by a membranous wing in Lophopetalum, while in Kokoona the wing is
apical. There is a sharp difference in pollen grains between the two genera; pollen grains of
Kokoona are single while that of Lophopetalum are in tetrads or polyads.
Key to Kokoona species

(based on flowers and fruits)
1.
Fruit surface with abundant pustules. Twigs whitish............................2. K. leucoclada

2.
Anthers with distinctly prolonged connective...............................................................3

Anthers without or with obscure connective....................................................................5
3.
Connective longer than anther-cells.............................................................4. K. ochracea

Connective shorter or as long as anther-cells......................................................................4
4.
Flower buds ovoid to globose; apex of calyx-lobes rounded or truncate....3. K. littoralis

Flower buds broad-ellipsoid; apex of calyx-lobes acute...............................1. K. coriacea
5.
Stamen filaments not broadened at base. Leaf margin strongly wavy and toothed toward
apex......................................................................................................................6. K. reflexa
Stamen filaments with broad thickened base. Leaf margin recurved, entire or only
slightly wavy......................................................................................................................6
6.
Inflorescences many-branched. Stigmas not papillose. Leaf margin recurved; intercostal veins equally prominent as the lateral ones...........................5. K. ovatolanceolata
Inflorescences with few short branches only. Stigmas papillose. Leaf margin entire or
slightly wavy; intercostal veins very faint.....................................................7. K. sabahana
123
Key to Kokoona species

1.
Leaf-margin entire to faintly wavy.......................................................................................2

Leaf-margin toothed..................................................................................................................5
2.
Twigs whitish. Leaves 6.510 cm wide; lateral veins 1012 pairs....2. K. leucoclada
Twigs blackish or dark-brown. Leaves to 6 cm wide; lateral veins 59 pairs............3
3.
Leaf-base almost rounded; margin strongly recurved. Common trees of peat and
freshwater swamp forests.........................................................................5. K. ovatolanceolata
Leaf-base not rounded; margin not recurved. Except for K. sabahana, usually not found
in swamp forests.................................................................................................................................4
4.
Small pole-sized tree. Petiole to 7 mm long, channelled above.........................7. K. sabahana

Medium-sized tree. Petiole to 2 cm long, wrinkled on drying.............................3. K. littoralis
5.
Twigs strongly 4-angled. Leaves drying to dark brown above and purplish below.................
...................................................................................................................................1. K. coriacea
Twigs not angled. Leaves drying pale yellow on both surfaces or pale greyish below.............6
6.
Leaves when dry pale yellow on both surfaces; margin strongly wavy and distinctly
toothed toward the apex..........................................................................................6. K. reflexa
Leaves when dry pale greyish below; margin shallowly toothed from the base to apex
.................................................................................................................................4. K. ochracea
1. Kokoona coriacea King

(Latin, coriaceus = leathery; the leaves)
l.c 347; Ding Hou l.c. (1962) 261; Kochummen & Whitmore l.c. 164. Type: Kunstler 4226, Perak
(holotype K; isotype BO). Synonym: Lophopetalum coriacea (King) Ridl. l.c. (1922) 450.
Medium-sized tree to 25 m tall, 35 cm diameter. Bark grey-white. Twigs 4-angled (somewhat like that of Glyptopetalum quadrangulare). Leaves leathery, drying to dark brown
above, and purplish below, oblong, 1113 x 57.5 cm; base cuneate, margin toothed,
slightly recurved, apex pointed; midrib sharply keeled below; lateral veins 57 pairs, faint;
intercostal veins invisible below; petioles c. 1 cm long. Flowers 5-merous, greenish yellow,
fragrant; buds broad ellipsoid; calyx-lobes triangular, apex acute; petals with pale margin;
connective of anthers prolonged, almost as long as anthers; free part of the ovary ovoid,
gradually narrowed into an obscure style, stigmas globose. Fruits unknown.
Distribution. Peninsular Malaysia, Borneo. In Sarawak uncommon, known by a single
collection (S. 40130) from Niah Cave National Park. Not yet recorded from Sabah.
Ecology. Lowland forest below 100 m.
This is the second collection of this species and a new record for Borneo. This species was
until now known only from the type collected from Perak in Peninsular Malaysia.
124
2. Kokoona leucoclada Kochummen

(Greek, leuco = white, cladus = branch)
l.c. (1994) 51. Type: Amin & Francis SAN 129399, Sabah, Ranau (holotype SAN; isotypes K, L).
Medium-sized to large tree to 35 m tall, 20 cm diameter. Bark grey, smooth; inner bark pale
brown. Sapwood ochre. Twigs whitish, youngest flattened, grooved in the centre. Leaves
leathery, drying to grey-brown, elliptic to oblong, 16.520 x 6.510 cm; base cuneate,
margin faintly wavy, slightly recurved, apex pointed; midrib raised above, keeled below;
lateral veins 1012 pairs, visible on both surfaces, distinctly looping near margin; intercostal
veins reticulate, very faint; petioles 11.5 cm long. Inflorescences axillary, to 14 cm long,
rachis rectangular, greyish. Flowers (immature) sessile, subtended by 5 bracteoles. Fruits c. 16
x 6 cm, with large pustules outside, on 20 cm long stalk. Seeds c. 12 x 2.5 cm.
Distribution. Endemic to Sabah. Uncommon, and known only from two collections, SAN
129399 (the type) from Ranau and SAN 61073 from Sandakan.
The flowers are too young to give details of anther and ovary characters. The pustulate fruits
and the white twigs distinguish this species from the others.
3. Kokoona littoralis Laws.

(Latin, littoralis = of the sea-shore)
in Hooker f. l.c. 617; Ding Hou l.c. (1962) 261; Burgess l.c. 73; Kochummen & Whitmore l.c. 164;
Cockburn l.c. 57; Anderson l.c. 160; Ashton l.c. 100; Whitmore, Tantra & Sutisna l.c. 43. Type:
Maingay 396/2, Malacca (holotype K; isotype BO).
Medium-sized tree to 27 m tall, 30 cm diameter, or rarely pole-sized treelet to 6 m tall and 2

cm diameter; bole slightly fluted with irregular hoops. Bark greyish or grey-brown with
yellowish tinge, smooth to scaly. Twigs blackish on drying, slightly swollen at the nodes,
youngest ones rectangular. Leaves thickly leathery, drying to dark green with purplish
blotches above and pale yellowish below, elliptic-lanceolate to oblong or narrowly obovate,
520 x 1.56.5 cm; base broadly cuneate to rounded, apex pointed; midrib faintly raised
above; lateral veins 69 pairs, very faint below, faint to inconspicuous above, looping and
joining near margin; intercostal veins visible or invisible; petioles 0.52 cm long, wrinkled
on drying. Flowers with yellow petals, pedicels 12.5 mm long; flower buds ovoidsubglobose, calyx-lobes semi-orbicular, apex rounded or truncate; petals broadly ovate,
elliptic or suborbicular; anthers with protruding connective to 1 mm long, slightly shorter
125
126
than the anthers; ovary triangular, style obscure, stigma capitate. Fruits 1318 x 35.5 cm,
base tapered. Seeds 7.512.5 x 2.5 cm (including wing).
Key to varieties
1.
Leaves c. 20 x 6.5 cm, margin pinkish...........................................................................

var. longifolia Kochummen
l.c. (1994) 53. Type: Wright & Ismawi S. 32289, Sarawak, Limbang (holotype SAR; isotype
K).
Small tree to 18 m tall, 20 cm diameter. Bark greyish, smooth to scaly. Leaves

thick-leatherly, elliptic to lanceolate, c. 20 x 6.5 cm; petiole c. 2 cm long, channelled
above; base cuneate, margin pinkish, apex pointed; midrib raised above; lateral
veins 79 pairs, faint on both surfaces; intercostal veins laxly reticulate, very faint.
Uncommon, known by a single collection (S. 32289) from Limbang, Sarawak.
Lowland forest.
Leaves to 11 x 6 cm, margin not pinkish......................................................................2
2.
Leaf apex blunt or rounded; intercostal veins invisible...................................................

var. bakoensis Kochummen
l.c. (1994) 53. Type: Ding Hou 534, Sarawak, Kuching (holotype SAR; isotype L).
Small pole-sized tree to 6 m tall, 2 cm diameter. Leaves coriacous, oblong with

rounded or blunt apex, 511 x 1.54 cm; lateral and intercostal veins invisible.
Locally frequent, known only from Bako National Park, Sarawak. Heath forest.
Leaf apex pointed; intercostal veins visible....................................................................
var. littoralis
Synonyms: Lophopetalum dubium Laws. in Hooker f. l.c. 616; L. maingayi Ridl. l.c. (1922)
450; L. littoralis (Laws.) Ridl. l.c. (1922) 450; K. scortechinii King l.c. 347; K. lanceolata
Ridl., Kew Bull. (1938) 237, Masamune l.c. 417; Solenospermum littorale (Laws.) Loes.,
Notizbl. Berl. Dahl. 13 (1936) 223.
Sumatra, Peninsular Malaysia, Borneo. In Sarawak, known from Semengoh Arboretum, Balingian, Belaga and Lundu. One sample was collected from G. Matang at
1000 m. Uncommon in Sabah with only 4 collections.
Ecology. Mixed dipterocarp forest on leached soils and heath forest to submontane forest to
1000 m, including limestone. Flowering in AugustOctober and fruiting in MayJuly.
4. Kokoona ochracea (Elmer) Merr.

(Latin, ochraceus = yellow or yellowish brown; the colour of the flowers)
En. Philip. 2 (1923) 484, PEB (1929) 171; Masamune l.c. 417; van Steenis l.c. 438; Ding Hou l.c.
(1962) 260; Burgess l.c. 74; Cockburn l.c. 57; Anderson l.c. 160; Ashton l.c. 101; Whitmore, Tantra &
Sutisna l.c. 43. Basionym: Ardisia ochracea Elmer, Leafl. Philip. Bot. 5(1913) 1819. Type: Elmer
12881, Philippines, Palawan (holotype K; isotype BO).
Medium-sized tree to 30 m tall, 50 cm diameter. Twigs black, terete. Leaves elliptic to ovate or
lanceolate, 713.5 x 3.56 cm, drying pale greyish below; base broadly cuneate, margin
shallowly toothed, apex pointed; midrib raised or flattened above; lateral veins 58 pairs,
127
slightly raised on both surfaces; petioles c. 1 cm long. Flowers almost sessile, in 12 cm long
panicles; calyx-lobes semi-orbicular; petals fleshy, ovate or elliptic, margin thin and
transparent; anthers subglobose, connective prolonged, longer than the anthers; ovary
conical, style obscure, stigma cylindrical. Fruits unknown.
Vernacular name. Sabahperupok kuning (Dusun, Malay).
Distribution. Peninsular Malaysia, Borneo, and Palawan Is. In Sabah, reported from
Kinabatangan, Lahad Datu, Tenom and Tawau districts. In Sarawak, collections were made
from northeastern parts.
Ecology. Scattered in mixed dipterocarp forests at low altitude.
5. Kokoona ovatolanceolata Ridl.
Fig. 5.
(Latin, ovatus = egg-shaped, lanceolatus = tapering toward both ends; the leaf shape)
l.c. (1938) 236; Masamune l.c. 417; Ding Hou l.c. (1962) 261; Smythies l.c. 39; Cockburn l.c. 57;
Anderson l.c. 160; Ashton l.c. 100; Whitmore, Tantra & Sutisna l.c. 43. Type: Beccari PB 3471,
Sarawak (holotype FI; isotypes BO, K). Synonym: K. scortechinii (non King) Steenis l.c. 438.
Tree to 36 m tall. Bark smooth to scaly, hoop-marked. Twigs dark brown to blackish.
Leaves ovate to ovate-lanceolate or elliptic, 711.5 x 45 cm; base rounded to broadly
cuneate, margin strongly recurved, apex pointed; midrib flattened above; lateral veins 56
pairs, very faint on both surfaces and hardly distinguishable from the equally prominent
reticulate intercostal veins; petioles 11.5 cm long, wrinkled and yellowish on drying.
Inflorescences many-branched panicles, to 14 cm long. Flowers yellowish, fragrant, buds
globose, c. 2 mm across; calyx-lobes suborbicular or reniform; petals ovate; anthers ovoid,
connective not prolonged, filaments with broad thickened base; ovary ovoid, style distinct,
stigma obtuse, not papillose. Fruits narrow-oblong, 1017 x 35 cm. Seeds 711 x 22.5
cm (including wing).
Vernacular name. Sarawakbajan paya (Iban, Malay).
Distribution. Endemic to Borneo. In Sabah, collected from Papar district and in Sarawak
from Baram, Marudi and Sibu. Common in Brunei.
Ecology. Known from heath, peat and freshwater swamp forests.
6. Kokoona reflexa (Laws.) Ding Hou

(Latin, reflexus = recurved; the leaf tip)
l.c. (1962) 262; Burgess l.c. 74; Kochummen & Whitmore l.c. 164; Cockburn l.c. 57; Anderson l.c. 40;
Ashton l.c. 101; Whitmore, Tantra & Sutisna l.c. 44. Basionym: Lophopetalum reflexum Laws. in
Hooker f. l.c. 616. Type: Maingay 383/2, Malacca (holotype K; isotype BO). Synonym: Hippocratea
maingayi Laws. in Hooker f. l.c. 625.
Emergent tree, to 50 m tall and 130 cm diameter; bole with stout buttresses. Bark dull greybrown, smooth, hoop-marked, becoming scaly. Leaves elliptic to lanceolate, 5.511 x 2.54
cm; base cuneate, margin strongly wavy and toothed towards the apex; apex pointed; midrib
128
raised above; lateral veins 5 pairs, slightly raised on both surfaces; petioles slender, 715
mm long, pale yellow on drying. Flowers pale yellowish green; calyx suborbicular; petals
ovate, or broadly elliptic; anthers oblong, connective not protruding, filaments not
broadened at base; ovary ovoid, narrowed to an obscure style, stigma rounded with flat top.
Fruits c. 12 x 3 cm. Seeds 911 x 2.5 cm including the wing.
Distribution. Sumatra, Peninsular Malaysia, and Borneo. Uncommon in Sabah, known
from a single collection (SAN 64149) from Ranau. In Sarawak, locally frequent, recorded
from Lundu, Semengoh, G. Gaharu, G. Matang, and Bako National Park.
Ecology. Mixed dipterocarp forests.
7. Kokoona sabahana Kochummen

(of Sabah)
l.c. (1994) 55. Type: Asik SAN 129928, Sabah, Nabawan (holotype SAN; isotypes A, BO, K, L, OX,
SAR, SING).
Small tree to 12 m tall, 10 cm diameter. Bark pale grey; inner bark brownish. Sapwood
yellowish. Twigs dark brown to black, swollen at the nodes. Leaves elliptic to oblong, 711
x 46 cm, shiny, drying to yellowish brown; base broadly cuneate, margin entire or slightly
wavy, apex pointed or blunt; midrib very faintly visible above; lateral veins 67 pairs,
looping near margin, visible below, faint above; intercostal veins reticulate, faintly visible
below; petioles 57 mm long, channelled above. Inflorescences axillary or terminal panicles, to
6 cm long, with a few branches. Flowers 5-merous; buds subglobose, c. 3 mm across;
pedicels c. 2 mm long, articulated; sepals triangular; petals slightly obovate with pale
margin; disc 5-lobed; stamens inserted within the disc, filaments broad and flat below,
tapering towards the apex, anthers without protruding connective; ovary immersed in the
disc, style distinct, stigma capitate, with papillae at the top. Fruits unknown.
Distribution. Endemic to Borneo. Uncommon, known only from 3 collections from Sabah,
SAN 129928 (the type) from Nabawan, SAN 61465 from Tawau, and SAN 43778 from
Sandakan.
Ecology. Lowland (including swamp) to hill forests to 600 m.
Close to K. ovatolanceolata from which it differs by the few-branched, short panicles, larger
flowers, papillose stigma, and shorter petiole.
6. LOPHOPETALUM Wight ex Arn.

(Greek, lophos = crested, petalum = petal)
perupok (Malay, Iban)
Ann. Mag. Nat. Hist. 1, 3 (1839) 150; Merrill l.c. (1921) 354; Ridley l.c. (1922) 447; Masamune l.c.
418; Browne l.c. 77; Ding Hou l.c. (1962) 262, l.c. (1969) 108; Backer & Bakhuizen f. l.c. 54; Balan
Menon l.c. 18; Smythies l.c. 38; Burgess l.c. 76; Kochummen & Whitmore l.c. 165; Jansen et al. l.c.
153; Cockburn l.c. 58; Anderson l.c. 160; Wong l.c. 121; Ashton l.c. 102; Whitmore, Tantra & Sutisna
129
l.c. 44; Ng l.c. 44. Synonym: Solenospermum Zoll., Nat. Tijd. Ned. Ind. 14 (1857) 168, Masamune l.c.
419.
Small to very large trees to 45 m tall and 180 cm diameter; bole with or without buttresses,
occasionally with pneumatophores. Bark similar to Kokoona. Twigs pale whitish, dark
brown to blackish, often flattened at the nodes. Leaves opposite or subopposite, margin
entire. Inflorescences axillary, peduncles distinct or obscure, pedicels jointed. Flowers
bisexual, 5-merous (except ovary); calyx-lobes spreading, inflexed or reflexed; petals free,
imbricate, inner surface partly covered with appendages or without; disc conspicuous,
usually fleshy and flat, surface smooth or denticulate, 5-angular, rounded or 5-lobed;
stamens 5, inserted on the disc, anthers introrse, pollen grains in clusters of fours (tetrads);
ovary usually semi-inferior, 3-celled, style short, stigma obscure, ovules 418 in each cell,
arranged in two series. Fruit a capsule, 3-angled. Seeds flat, surrounded by membranous
wing; endosperm absent; germination as in Kokoona; cotyledons very thin, seedlings leaves
minutely bistipulate, the first two leaves opposite, subsequent leaves alternate on the leader
shoots but opposite on the branches.
Distribution. About 18 species; India, Burma, Thailand, Indo-China, Malesia, and Australia. In
Malesia 15 species, 10 of which are in Sabah and Sarawak.
Ecology. Lowland (including swamp) to submontane forests to 1500 m.
Uses. The timber is a light hardwood. It is suitable for interior finishing, panelling, partitioning,
furniture, veneers, plywood, boxes, crates and mathematical instruments.
Key to Lophopetalum species

1.
Leaves short-petioled or almost sessile.........................................................................2

Leaves with distinct petioles..............................................................................................3
2.
Twigs rounded, not winged. Lateral veins 57 pairs, sunken above..........2. L. glabrum
Twigs winged, lateral veins 1015 pairs, not sunken above.................8. L. sessilifolium
3.
Leaves drying to greenish brown above, lower surface with black gland-dots.................
...................................................................................................................1. L. beccarianum
4.
Intercostal veins invisible below...................................................................................5

Intercostal veins faint to distinctly visible below..............................................................7
5.
Blade thinly leathery, apex rounded or notched; lateral veins 46 pairs..........................

..........................................................................................................9. L. subobvatum
Blade thickly leathery, apex pointed, blunt or rounded; lateral veins 411 pairs...........6
130
6. Petioles 22.5 cm long. Blade densely papillose below, margin curled inwards..............
......................................................................................................5. L. pachyphyllum
Petiole 38 mm long. Blade not papillose below, margin not curled inwards..................
................................................................................................................7. L. rigidum
7.
Intercostal veins prominently raised below.......................................10. L. wightianum

Intercostal veins faint below.........................................................................................8
8.
Leaves pale whitish below. Twigs pale whitish.........................................6. L. pallidum

Leaves not pale whitish below. Twigs dark brown to blackish......................................9
9.
Lateral veins 810 pairs. Petiole 11.5 cm long....................................3. L. javanicum

Lateral veins 1015 pairs. Petiole 1.53 cm long.............................4. L. multinervium
Key to Lophopetalum species

(based on flowers and leaves)
1.
Flower buds almost flat, or wider than long; petals without appendages on the inner
surfaces..............................................................................................................................2
Flower buds conical or subglobose, longer than wide; petals with appendages on the
inner surfaces....................................................................................................................5
2.
Twigs distinctly 4-angled to winged. Inflorescences 1850 cm long; pedicels 715 mm

long...................................................................................................8. L. sessilifolium
Twigs usually rounded. Inflorescences to 15 cm long; pedicels to 9 mm long...............3
3.
Disc dish-shaped, 5-lobed............................................................................................4

Disc suborbicular, flat or obscurely 5-angled...........................................7. L. rigidum
4.
Petioles 715 mm long. Leaves with black gland-dots below; lateral veins not sunken
above.....................................................................................................1. L. beccarianum
Petioles to 3 mm long. Leaves without black gland-dots below; lateral veins sunken
above.......................................................................................................2. L. glabrum
5.
Disc 4.59.5 mm in diameter at anthesis.........................................................................6

Disc 13 mm in diameter at anthesis........................................................................7
6.
Leaves densely papillose beneath, intercostal veins invisible............5. L. pachyphyllum

Leaves not papillose beneath, intercostal veins prominent below.......10. L. wightianum
7.
Disc with fleshy subulate processes around the base of filaments. Leaves pale whitish
below......................................................................................................6. L. pallidum
Disc smooth or minutely papillose. Leaves not pale whitish below................................8
8.
Leaf-apex rounded or notched; lateral veins 46 pairs..........................9. L. subovatum

Leaf-apex pointed; lateral veins 815 pairs.........................................................................9
9.
Petals 11.5 cm long. Lateral veins 810 pairs.........................................3. L. javanicum

Petals 1.53 cm long. Lateral veins 1015 pairs..............................4. L. multinervium
131
1. Lophopetalum beccarianum Pierre
Fig. 6AD.

Fl. For. Coch. 4 (1894) sub t. 307; Merrill l.c. (1921) 354; Masamune l.c. 418; Ding Hou l.c. (1962)
266; Smythies l.c. 39; Kochummen & Whitmore l.c. 165; Cockburn l.c. 59; Anderson l.c. 160; Ashton
l.c. 103; Whitmore, Tantra & Sutisna l.c. 44. Type: Beccari PB 2475, Sarawak (holotype FI; isotypes
BO, K). Synonyms: Lophopetalum scortechinii King l.c. 350; L. havilandii Ridl. l.c. (1931) 37,
Masamune l.c. 418.
Small to medium-sized tree, rarely to 36 m tall and 60 cm diameter. Bark grey-brown,

smooth, hoop-marked; inner bark pale brown. Sapwood pale yellow. Twigs dark brown to
black, rounded. Leaves ovate, elliptic or oblong, drying to greenish brown above, with
abundant black gland-dots below, 430 x 2.59 cm; base cuneate, apex pointed or blunt;
midrib raised above; lateral veins 58 pairs, visible below, faint above, often pinkish when
dried; intercostal veins very faint; petioles 715 mm long. Flowers yellowish, in axillary
panicles to 15 cm long; buds flat; pedicels c. 5 mm long; calyx-lobes triangular or
suborbicular; petals suborbicular or broadly ovate, usually without appendages on inner
side, sometimes with distinct veins; disc dish-shaped, 5-lobed; stamen filaments with
cushion-like thickening at base; ovary pyramidal, style and stigma obscure. Fruits 5.211 x
33.2 cm. Seeds 4.5 x 1.51.2 cm (including wing).
Distribution. Peninsular Malaysia and Borneo. In Sabah and Sarawak frequent.
Ecology. Mixed dipterocarp forest to submontane forest to 1800 m, mainly on ridges and
hillsides on clay-rich soils. Flowering in FebruaryNovember.
2. Lophopetalum glabrum Ding Hou
Fig. 6E.
(Latin, glabrus = smooth; without pubescence)

l.c. (1962) 266; Anderson l.c. 160; Ashton l.c. 105; Whitmore, Tantra & Sutisna l.c. 44. Type: Rutten
83, E Borneo (holotype U; isotype BO).
Small tree to 15 m tall, 15 cm diameter. Bark grey-white; inner bark yellowish. Sapwood
white. Twigs grey-brown, rounded. Leaves elliptic or oblong, 816 x 2.55.5 cm, without
black gland-dots below; base cuneate, apex pointed; midrib raised above; lateral veins 57
pairs, raised below, usually sunken above, curving and joining near margin; intercostal
veins reticulate, faint below, visible above; petioles to 3 mm long, cracking. Flowers
yellowish, in branched panicles, to 9 cm long; buds flat; pedicels 57 mm long; calyx-lobes
deltoid with pointed tips; petals suborbicular, without appendages inside; disc dish-shaped,
5-lobed; anthers suborbicular; ovary pyramidal, style and stigma obscure. Fruits reddish
brown when fresh, obovoid, 1012.5 x 3 cm, apex rounded, drying brownish, surface
pustulate. Seeds 4 in each locule, c. 7.5 x 1.5 cm.
Distribution. Endemic to Borneo. Widespread in Sabah and Sarawak, also found in Brunei
and Kalimantan.
Ecology. Mixed dipterocarp forests on clay-rich soils to 400 m. Flowering in April and
September and fruiting in March.
132
133
3. Lophopetalum javanicum (Zoll.) Turcz.

(of Java)
Bull. Soc. Nat. Hist. Mosc. 36 (1863) 598; Ding Hou l.c. (1962) 269; Backer & Bakhuizen f. l.c. 54;
Smythies l.c. 39; Burgess l.c. 77; Kochummen & Whitmore l.c. 168; Cockburn l.c. 60; Anderson l.c.
161; Ashton l.c. 104; Whitmore, Tantra & Sutisna l.c. 44. Basionym: Solenospermum javanicum Zoll.
l.c. 169. Type: Zollinger 3254, Java (lectotype L; isolectotype BO). Synonyms: Lophopetalum
fuscescens Kurz, J. As. Soc. Beng. 44, 2 (1875) 202; L. oblongum King l.c. 350; L. oblongifolium King
l.c. 351; L. intermedium Ridl., J. Str. Br. R. As. Soc. 59 (1911) 85; L. paucinervium Merr., Philip. J.
Sc. 20 (1922) 402; Solenospermum paucinervium (Merr.) Loes. l.c. 223; S. oblongifolium (King) Loes.
in Engler & Prantl, Pfl. Fam. 2, 20b (1942) 162.
Medium-sized tree to 30 m tall, 90 cm diameter. Bark yellowish grey, smooth to cracking,

lenticellate; inner bark pink. Sapwood pale. Twigs strongly angled, often glaucous, dark
brown to black. Leaves drying to dark brown, not pale whitish beneath, elliptic to oblong or
obovate, 815 x 47 cm; base cuneate, apex pointed; midrib raised above; lateral veins 8
10 pairs, raised below, faint to inconspicuous or sunken above; intercostal veins reticulate,
faintly visible below, invisible above; petioles 11.5 cm long. Flowers in panicles; buds shortconical or subglobose; calyx-lobes spreading, margin ciliate; petals with lobed appendages
inside; disc 13 mm across at anthesis; stamens with distinct (1.5 mm long) filaments; ovary
triangular, narrowed into a cylindric style, stigma obscure. Fruits 4.811 x 2.5 cm. Seeds
6.06.5 x 1.31.6 cm.
Distribution. Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, Philippines, Celebes
and New Guinea. Common in Sabah and Sarawak.
Ecology. Lowland to hill and montane forests, mainly in mixed dipterocarp forests,
sometimes by river banks and peat swamps. Flowering in MarchJune and fruiting in
September.
Uses. The bark is used in Borneo as a constituent of dart poison.
There is a tendency of "witches broom" developing on the leafy shoots which can be mistaken
for inflorescences. Blackish twigs, shorter petioles, inconspicuous intercostal veins and the
absence of pneumatophores are good field characters to separate L. javanicum from L.
multinervium.
4. Lophopetalum multinervium Ridl.

(Latin, multi = many, nervus = veins; the leaves)
l.c. (1931) 39; Masamune l.c. 418; Ding Hou l.c. (1962) 270; Smythies l.c. 39; Burgess l.c. 77;
Kochummen & Whitmore l.c. 168; Cockburn l.c. 60; Anderson l.c. 161; Ashton l.c. 105; Whitmore,
Tantra & Sutisna l.c. 44. Type: Beccarii PB 3659, Sarawak (holotype FI; isotypes BO, K). Synonym:
Solenospermum aquatile Ridl. l.c. (1938) 236, Masamune l.c. 419.
Tree to 36 m tall, 60 cm diameter; bole with tall buttresses and pneumatophores. Bark
yellowish, smooth, lenticellate; inner bark pinkish. Sapwood whitish. Twigs dark brown,
youngest ones angled. Leaves leathery, not pale whitish below, ovate to oblong or elliptic,
1018 x 48 cm, drying brownish; base rounded, obtuse or broadly cuneate, apex pointed;
midrib raised above; lateral veins 1015 pairs, raised on both surfaces; intercostal veins
134
distinct below; petioles 1.53 cm long. Flowers in panicles, pink to dark reddish to purplish or
yellowish or whitish; buds short-conical or subglobose; calyx triangular, margin ciliate;
petals with lobed appendages at the inner side near base; disc 13 mm across at anthesis;
stamens filaments distinct, c. 1.7 mm long. Fruits 48.5 x 1.52.5 cm, furfuraceous outside.
Seeds c. 5 x 1.25 cm.
Distribution. Sumatra, Peninsular Malaysia, and Borneo. In Sabah and Sarawak widely
distributed.
Ecology. Freshwater and peat swamp forests (very rarely in submontane forests at 1500 m).
Flowering in March to August.
5. Lophopetalum pachyphyllum King

(Greek, pakus = thick, phyllon = leaf; thick-leaved)
l.c. 348; Ridley l.c. (1922) 448; Ding Hou, l.c. (1962) 267; Kochummen & Whitmore l.c. 168. Type:
King's collector 7525, Perak (BO, K).
Medium-sized tree to 25 m tall and 30 cm diameter. Twigs stout, c. 1 cm thick, dark brown.
Leaves thickly leathery, elliptic, drying to greenish grey, densely papillose below, c. 10 x 5
cm; base broadly cuneate, margin curled inwards, apex blunt; midrib raised above, lateral
veins 67 pairs, very faint on both surfaces; intercostal veins invisible; petioles 22.5 cm
long, stout, dark brown. Inflorescences panicles to 10 cm long, pedicels to 1 cm long.
Flowers c. 1.5 cm across; buds short-conical or subglobose; calyx faintly 5-lobed; petals
with small appendage inside; disc obscurely 5-lobed, 4.59.5 mm across at anthesis;
stamens with distinct filaments; ovary ovoid. Fruits unknown.
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sarawak uncommon, known by
two collection (S. 6961 and S. 16212) from Bako National Park; not yet recorded from
Sabah.
Ecology. Lowland forests near the coasts.
6. Lophopetalum pallidum Laws.

(Latin, pallidus = pale; the dry leaves)
in Hooker f. l.c. 615; Ridley l.c. (1922) 449; Ding Hou l.c. (1962) 268; Burgess l.c. 77; Kochummen &
Whitmore l.c. 168; Anderson l.c. 161; Ashton l.c. 106; Whitmore, Tantra & Sutisna l.c. 44. Type:
Maingay 1389, Malacca (BO, K). Synonyms: L. curtisii King l.c. 351; Solenospermum pallidum
(Laws.) Loes. l.c. 225.
Tree to 45 m tall, 80 cm diameter. Bark grey, scaly; inner bark orange-brown. Sapwood
whitish. Twigs pale whitish. Leaves thinly leathery, glaucous below, elliptic to narrowly
obovate, 711 x 2.56.5 cm; base cuneate, apex cuspidate with short tip; midrib raised
above, pale whitish below; lateral veins 810 pairs with short intermediate veins, very faint
on both surfaces; intercostal veins very faint; petioles 1.52 cm long, pale whitish. Flowers
in panicles; buds short-conical or subglobose; calyx semi-orbicular or triangular; petals with
135
fleshy lobed processes in the central part inside; disc 13 mm across at anthesis; stamens
surrounded by fleshy subulate processes of the disc; ovary triangular, style cylindric. Fruits c.
15 cm long. Seeds c. 9 x 2.3 cm.
Distribution. Sumatra, Peninsular Malaysia, Borneo. In Sabah uncommon, known by only
two collection (SAN 74348 and SAN 65428) from Sandakan and Tawau respectively. In
Sarawak known from Bintulu, Bt. Mersing, Bt. Raya, and Kapit.
Ecology. Mixed dipterocarp forests on clay soils.
Uses. In Peninsular Malaysia the bark is used as an ingredient in making dart poison by the
aborigines.
7. Lophopetalum rigidum Ridl.

(Latin, rigidus = stiff; the leaf texture)
l.c. (1931) 38; Masamune l.c. 418; Ding Hou l.c. (1962) 267; Smythies l.c. 39; Burgess l.c. 77;
Cockburn l.c. 58; Anderson l.c. 161; Ashton l.c. 106; Whitmore, Tantra & Sutisna l.c. 44. Type:
Haviland 2236, Sarawak, Kuching (BO, K). Synonym: L. subsessile Ridl. l.c. (1931) 37, Masamune
l.c. 418.
Shrub or small tree, very rarely reaching 30 m tall and 40 cm diameter. Bark grey-brown,
smooth, hoop-marked with vertical line of lenticels; inner bark purplish brown. Sapwood
yellow-brown. Twigs dark brown, rounded. Leaves thickly leathery, drying to greenish
brown or reddish brown, not papillose below, ovate-oblong, 4.521 x 24 cm; base broadly
cuneate to rounded, margin not curled inwards, apex pointed or rounded; midrib raised
above; lateral veins 411 pairs, visible below, faint to invisible above, looping near margin;
intercostal veins invisible; petioles stout, 3(8) mm long. Flowers yellowish green, in 10 cm
long panicles, pedicels to 5 mm long; buds almost flat, suborbicular or obscurely 5-angled;
calyx-lobes ovate or triangular, pointed, with small papilla-like processes outside; petals
triangular or suborbicular, without appendages inside, with papilla-like processes outside;
disc suborbicular, flat or obscurely 5-angled; stamens with small short-apiculate anthers;
ovary pyramidal, style cylindrical. Fruits to 11 cm long, surface shortly tuberculate. Seeds c.
5.5 x 1.5 cm, including wing.
Distribution. Endemic to Borneo. In Sabah, recorded from Lahad Datu (common),
Keningau and Ranau. In Sarawak, locally frequent.
Ecology. Freshwater swamps, peat swamps, kerangas swamps and also in montane forests
to 2400 m.
136
8. Lophopetalum sessilifolium Ridl.

(Latin, sessile = stalkless, folium = leaves)
l.c. (1931) 37; Masamune l.c. 418; Ding Hou l.c. (1962) 265; Anderson l.c. 161; Ashton l.c. 107;
Whitmore, Tantra & Sutisna l.c. 44. Type: Haviland 1744, Sarawak, Kuching (BO, K).
Small tree, rarely reaching 20 m tall, 20 cm diameter. Twigs dark brown, strongly 4-angled
to winged. Leaves elliptic to oblong or lanceolate, 2250 x 4.512 cm, drying to dark brown
with brown gland-dots below and grey-brown above; base broadly cuneate to rounded, apex
pointed, acumen to 2 cm long; midrib raised above; lateral veins 1015 pairs, raised on both
surfaces, with short intermediate veins; intercostal veins reticulate (rarely scalariformreticulate), distinct below, faint above; petioles very short to almost absent, rarely to 1 cm
long. Flowers yellow, in 1850 cm long panicles, pedicels 715 mm long; buds wider than
long; calyx-lobes triangular, pointed; petals suborbicular or deltoid with blunt apex, without
appendages on the inner side; anthers obtuse; ovary immersed in disc without distinct style
or stigma. Fruits with tuberculate surface, 7.510.5 x 3 cm. Seeds 3.84.5 x 11.3 cm.
Distribution. Endemic to Borneo. In Sarawak, locally abundant in the W Baram valley; also
known from Bau, Kapit, Samunsam Wild Life Sanctuary, Semengoh Arboretum, Sg. Jeong,
Sg. Kelawit and Sg. Temulan. Not yet reported from Sabah.
Ecology. Lowland forests by rivers. Flowering in February, September and October and
fruiting in August and September.
9. Lophopetalum subovatum King

(Latin, sub = somewhat, ovatus = egg-shaped; the leaf shape)
l.c. 349; Ridley l.c. (1922) 448; Ding Hou l.c. (1962) 271; Burgess l.c. 77; Kochummen & Whitmore
l.c. 169; Cockburn l.c. 59; Anderson l.c. 161; Ashton l.c. 109; Whitmore, Tantra & Sutisna l.c. 45.
Type: Curtis 1501, Penang (BO, K). Synonym: Solenospermum apiculatum Ridl. l.c. (1938) 235.
Medium to large-sized tree to 36 m tall, 75 cm diameter. Bark pale brown, cracking to

scaly, hoop-marked; inner bark pinkish brown. Sapwood pale yellow. Leaves thinly
leathery, drying to brownish, not pale whitish beneath, obovate to elliptic, 4.513 x 2.57
cm; base tapered, apex rounded to notched; midrib raised above; lateral veins 46 pairs,
curving and joining near margin, faintly visible below, faint to inconspicuous above;
intercostal veins invisible; petioles 515 mm long. Flowers in axillary panicles; buds shortconical or subglobose; calyx-lobes spreading, triangular; petals inside with small appendage on
the upper half; disc 13 mm across at anthesis, smooth or minutely papillose; anthers
deltoid, obtuse; ovary triangular, narrowed towards apex, style cylindrical. Fruits to 7 cm
long. Seeds c. 4 x 0.6 cm.
Distribution. Sumatra, Peninsular Malaysia and Borneo. Frequent in Sabah and Sarawak.
Ecology. In mixed dipterocarp forests on leached clay-rich soils, on periodically inundated
alluvium, undulating lands and ridges to 900 m.
137
10. Lophopetalum wightianum Arn.

(R. Wight, 19th Century botanist in India)
l.c. (1839) 151; Ding Hou l.c. (1962) 267; Kochummen & Whitmore l.c. 169; Cockburn l.c. 59;
Whitmore, Tantra & Sutisna l.c. 45. Type: Wight (Icon. t. 162), S India, Malabar (K). Synonyms: L.
fimbriatum Wight in Ridley l.c. (1922) 448; L. winkleri Loes. l.c. 221.
Emergent tree to 40 m tall, 100 cm diameter. Bark dark grey, fissured; middle bark whitish;
inner bark purplish brown. Leaves elliptic to ovate, 1018 x 57 cm, not papillose beneath;
base rounded to wedge-shaped, slightly peltate, apex pointed; midrib raised above; lateral
veins 612 pairs, pale below; intercostal veins prominent below; petioles 1.52.5 cm long.
Flowers yellow, in panicles, pedicels 59 mm long; buds short-conical or subglobose; calyx
distinctly 5-lobed; petals broadly ovate to rounded with wavy margin, appendages lamellate
to cristate and attached to the lower half of petals; disc 4.59.5 mm across at anthesis;
stamens with oblong apiculate anthers; ovary triangular, narrowed into style. Fruits to 15
cm long. Seeds c. 6 x 1.5 cm.
Distribution. India, Indo-China, Sumatra, Peninsular Malaysia, and Borneo. In Sabah,
uncommon and recorded from Sandakan and Tawau only. Not yet recorded from Sarawak.
7. MICROTROPIS Wall. ex Meisn., nom. cons.

(Greek, micro = small, tropis = keel; a small keel on the inner side of the petal)
Pl. Vasc. Gen. Tabul. Diagn. (1837) 68; Ridley l.c. (1922) 443; Merrill l.c. (1921) 354; Merr. &
Freem., Proc. Am. Acad. Arts. Sc. 73 (1940) 276; Masamune l.c. 418; Ding Hou l.c. (1962) 272;
Backer & Bakhuizen f. l.c. 54; Kochummen & Whitmore l.c. 170; Cockburn l.c. 61; Anderson l.c.
161; Ashton l.c. 109; Kochummen, Sandakania 5 (1994) 55. Synonyms: Microtropia Reichb.,
Nomencl. (1841) 190; Paracelastrus Miq., Fl. Ind. Bat. 1, 2 (1859) 590.
Shrubs or small trees. Terminal node with one or two pairs of prominent, subpersistent,
reduced leaves (incipient bracts). Leaves opposite, glabrous, entire. Inflorescences axillary or
extra-axillary, dichotomous or paniculate cymes, sometimes in sessile clusters. Flowers
bisexual, 5- or 4-merous; calyx-lobes almost free, imbricate, persistent, unequal in size, the
outer 2 or 3 usually smallest, margin often thinner, transparent, entire, gnawed, irregularly
split or fimbriate; petals free or united, imbricate; disc absent; stamens 45, filaments united
at base into a ring or a short tube, the united part free from the petals or adnate to it,
sometimes the stamens inserted in the mouth of corolla, anthers dorsifixed, usually introrse;
ovary superior, cylindrical, conical or flask-shaped, 2-celled, ovules 2 in each cell, style
very short or cylindrical, stigma obscure, rarely 24-lobed. Fruit a capsule, striated
lengthwise, splitting along one side, apex with short beak, calyx persistent. Seeds usually 1,
erect on a knob-like thickened hilum enveloped by aril, with endosperm.
Distribution. About 80 species, C America, SE to E Asia and Malesia. 20 species in Malesia,
of which 14 are in Sabah and Sarawak.
Ecology. Lowland to montane forests to 2700 m, more common in montane forests.
138
Key to Microtropis species

(based mainly on leaves)
1. Leaves thickly leathery, margin distinctly curled inwards...............................................2
Leaves thickly or thinly leathery, margin not distinctly curled inwards (except in M.
grandifolia and M. sarawakensis)................................................................................4
2.
Leaves greyish below; lateral veins and intercostal veins faintly sunken above...............
..............................................................................................................1. M. argentea
Leaves not greyish below; lateral veins and intercostal veins not sunken above............3
3.
Petioles 710 mm long. Intercostal veins invisible below....................2. M. borneensis

Petioles absent or to 2 mm long. Intercostal veins visible below..................9. M. rigida
4.
Leaves drying to greenish yellow. Incipient bracts greenish.........................................5

Leaves drying brownish. Incipient bracts brown..........................................................7
5.
Incipient bracts needle-like, c. 5 x 1 mm...........................................10. M. sabahensis

Incipient bracts elliptic, lanceolate or oblong, at least 4 mm broad..................................6
6.
Intercostal veins equally prominent as the faint lateral veins. Incipient bracts oblong or
lanceolate.......................................................................................6. M. kinabaluensis
Intercostal veins invisible or very faint. Incipient bracts elliptic or lanceolate.................
.....................................................................................................11. M. sarawakensis
7.
Leaves to 4 cm long; base usually rounded to subcordate, apex blunt or rounded..........8

Leaves longer than 4 cm; base cuneate, apex usually pointed.........................................9
8.
Ovary ovoid-oblong.....................................................................................7. M. ovata

Ovary conical.......................................................................................14. M. wallichiana
9.
Incipient bracts foliaceous, 1.73 cm long.............................................4. M. grandifolia

Incipient bracts stiff, not foliaceous, to 12 mm long.......................................................10
10. Leaves uniformly papillose below with scattered pustules..............5. M. keningauensis

Leaves not papillose, without pustules........................................................................11
11. Inflorescences in sessile clusters........................................................3. M. fascicularis
Inflorescences distinctly stalked.................................................................................12
12. Petioles 0.71 cm long......................................................................12. M. sumatrana
Petioles 12 cm long..................................................................................................13
13. Ovary cylindrical, slightly constricted in the middle.................................13. M. valida
Ovary conical, without constriction in the middle................................8. M. platyphylla
139
Key to Microtropis species

(based mainly on flowers)
1. Inflorescences or infructescences in sessile clusters or condensed cymes; peduncle if
present to 1 cm long..........................................................................................................2
Inflorescences and infructescences with peduncle more than 1 cm long.......................6
2.
Lower surface of leaf uniformly papillose and with scattered pustules....5. M.

keningauensis
Lower surface of leaf not papillose, without pustules.......................................................3
3.
Ovary cylindrical or ovoid-oblong................................................................................4

Ovary conical or gradually tapered to apex...............................................................5
4.
Ovary cylindrical with slight constriction in the middle. Petioles 12 cm long...............

...................................................................................................13. M. valida (in part)
Ovary ovoid-oblong. Petioles absent or very short, to 2 mm long.................7. M. ovata
5.
Stigmas 4-lobed; ovary gradually tapered towards apex.....................3. M. fascicularis

Stigmas not lobed; ovary conical.....................................................14. M. wallichiana
6.
Leaves thickly leathery; margin distinctly curled inwards............................................7

Leaves thinly to thickly leathery; margin not distinctly curled inwards (faintly so in M.
grandifolia and M. sarawakensis)..............................................................................10
7.
Petioles very short, to only 2 mm long or almost absent..............................9. M. rigida

Petioles distinct, at least 7 mm long.............................................................................8
8.
Lower surface of leaf greyish; lateral veins and intercostal veins sunken above..............
..............................................................................................................1. M. argentea
Lower surface of leaf greenish brown; lateral veins and intercostal veins invisible
above...........................................................................................................................9
9.
Incipient bracts 1.73 x 1.22 cm; petioles 2.36 cm long....4. M. grandifolia (in part)
Incipient bracts to 5 mm long. Petioles 710 mm long........................2. M. borneensis
10. Petals distinctly united, at least up to lower half................................10. M. sabahensis

Petals free or only slightly united at the very base..................................................11
11. Ovary cylindrical........................................................................13. M. valida (in part)
Ovary conical, flask-shaped or gradually narrowed towards apex...............................12
12. Ovary flask-shaped....................................................................................................13
Ovary conical or gradually narrowed towards apex....................................................14
13. Incipient bracts ovate, 1.73 x 1.22 cm. Leaves 2030 x 8.514.5 cm; petiole 2.36
cm long................................................................................4. M. grandifolia (in part)
Incipient bracts lanceolate or oblong, to 1.5 cm long. Leaves 1124.5 x 3.59 cm;
petiole 12 cm long........................................................................6. M. kinabaluensis
140
14. Stigmas 4-lobed; ovary gradually narrowed towards apex...................12. M. sumatrana

Stigmas or apex of ovary not lobed; ovary conical......................................................15
15. Leaves thinly leathery. Incipient bracts green, lanceolate or elliptic.....11. M. sarawakensis
Leaves thickly leathery. Incipient bracts brown, lanceolate........................8. M. platyphylla
1. Microtropis argentea Kochummen

(Latin, argenteus = silvery; the lower leaf surface)
l.c. (1994) 55. Type: Lee S. 39980, Sarawak, Belaga (holotype SAR; isotypes E, K, KEP, L, SAN).
Small tree to 5 m tall. Incipient bracts lanceolate, stiff, brown, c. 5 mm long. Twigs swollen at
nodes, grey, rounded. Leaves thickly leathery, drying to pale green above, greyish below,
elliptic, 10.512 x 5.5 cm; base cuneate, margin prominently recurved, with thick rim, apex
shortly pointed, incurved; midrib prominently raised above; lateral veins 67 pairs, invisible
below, faintly sunken above; intercostal veins invisible below, reticulate and faintly sunken
above; petioles 0.71 cm long, wrinkled on drying. Inflorescences in cymes to 1.5 cm long;
peduncles c. 1 cm long; buds globose, c. 2 mm across. Flowers white, 5-merous; sepals
rounded with gnawed margins, wrinkled outside; petals united to more than half the length,
lobes oblong; stamens at mouth of corolla tube, filaments very short, anthers globose; ovary
conical, apical part ridged with no distinct style or stigma. Fruit unknown.
Distribution. Endemic to Borneo. Uncommon, known only from the type specimen from
Belaga, Sarawak.
Ecology. Kerangas forest on plateau at 700 m.
2. Microtropis borneensis Merr. & Freem.

(of Borneo)
l.c. 296; Masamune l.c. 418; Kochummen l.c. (1994) 63. Type: Clemens 31742, British North Borneo,
Mt. Kinabalu (holotype A; isotypes B, BO, GE, N, UC).
Shrub or small tree. Twigs grey, youngest ones reddish brown and slightly angled. Incipient
bracts triangular, stiff, c. 5 mm long. Leaves thickly leathery, drying to greenish brown on
both surfaces, elliptic, 5.59 x 2.54.5 cm; base cuneate, margin with thick rim, wavy and
strongly recurved, apex pointed; midrib prominently raised above; lateral veins 57 pairs,
very faintly visible below only; intercostal veins invisible; petioles 710 mm long, wrinkled
on drying. Inflorescences and flowers unknown. Infructescences with a peduncle c. 2 cm
long. Fruits when fresh oblong, c. 2 x 1.5 cm, yellow, drying black.
Distribution. Endemic to Borneo. Locally abundant on Mt. Kinabalu in Sabah. In Sarawak,
known from a single collection (S. 40412), from Lambir National Park.
Ecology. Kerangas and montane forests to 2400 m.
Taxonomy. Ding Hou l.c. 279 considered this a synonym of M. platyphylla Merr. After
studying the types and other recently collected specimens of both species, I have come to
the conclusion that M. borneensis should be reinstated as a distinct species.
141
3. Microtropis fascicularis Kochummen
Fig. 7.
(Latin, fascicularis = in a small bundle; the flower clusters)

l.c. (1994) 57. Type: Lee S. 44277, Sarawak, Lingga (holotype KEP; isotypes A, K, L, SAN, SAR).
Treelet to 2 m tall. Incipient bracts stiff, brown, lanceolate, c.7 x 1 mm, with sharp tip.
Twigs dark brown, rounded. Leaves leathery, surface not papillose nor pustulate, elliptic or
oblong, 10.520.5 x 48.5 cm; base cuneate, margin slightly recurved, apex pointed; midrib
raised above; lateral veins 1012 pairs, looping near margin, very faint on both surfaces;
intercostal veins reticulate, very close, visible on both surfaces; petioles 0.52.2 cm long,
wrinkled on drying. Flowers in sessile fascicles, 4-merous; sepals rotund, margin gnawed,
c. 2 x 1.5 mm; petals free, oblong to obovate, c. 1.5 x 1.2 mm; stamens free, anthers oblong,
c. 1 mm long; ovary gradually tapered towards apex, striate, stigma 4-lobed. Fruits green
when fresh, drying black, oblong or ellipsoid, 1518 x 46 mm, with pointed tip.
Distribution. Endemic to Borneo. Uncommon, known by 5 collections from Sarawak.
Ecology. Mixed dipterocarp forest and kerangas forest, to 500 m, usually by streams.
Flowering in December and fruiting in MarchMay.
4. Microtropis grandifolia Kochummen

(Latin, grandis = large, folius = leaf)
l.c 1994) 59. Type: Awa & Ilias S. 47380, Sarawak, Lundu (holotype KEP; isotypes K, SAR).
Small tree to 5 m tall. Twigs brownish, youngest ones flattened. Incipient bracts prominent,
foliaceous, ovate to elliptic, 1.73 x 1.22 cm. Leaves thinly to thickly leathery, elliptic,
drying greenish brown on both surfaces, 2030 x 8.514.5 cm; base broadly cuneate,
margin faintly wavy, faintly curled inwards, apex pointed; midrib flattened to raised above;
lateral veins 1012 pairs, looping near margin, faintly raised on both surfaces with short
intermediate veins; intercostal veins reticulate, faintly visible above; petioles 2.36 cm long,
wrinkled on drying. Inflorescences paniculate cymes, c. 67 cm long; peduncles 23 cm
long; bracteoles triangular, transparent. Flowers 4-merous; outer two sepals larger than
inner 2, rotund; petals slightly united at base; anthers sessile, jointed at base on a staminal
ring, connective slightly prolonged; ovary flask-shaped, style ridged, stigma 24-lobed.
Fruits unknown.
Key to varieties
Leaves thinly leathery, margin slightly incurved. Petioles c. 3 cm. Stigmas 4-lobed..............
var. grandifolia
Endemic to Sarawak. Uncommon, known only by two collections (S. 47380 from Bukit
Panjo, Lundu, and S. 34033 from Lubok Antu). Lowland and hill forests to 600 m.
Flowering in May.
Leaves thickly leathery, margin strongly incurved. Petioles c. 6 cm. Stigma 2-lobed.............
var. longipetiolatus Kochummen
l.c. (1994) 59. Type: Chai S. 34033, Sarawak, Lanjak Entimau (holotype SAR;
isotypes K, KEP, L, MO, SAN). Uncommon, known only from the type collection.
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143
5. Microtropis keningauensis Kochummen

(of Keningau, in Sabah)
l.c. (1994) 61. Type: Fidilis SAN 118405, Sabah, Keningau (holotype SAN).
Shrub or small tree. Twigs grey-brown. Incipient bracts brown, lanceolate, c. 10 x 2 mm,
with distinct median ridge. Leaves leathery, drying brownish, uniformly papillose below
with scattered pustules, elliptic, 11.523.5 x 4.58.5 cm; base cuneate, margin not curled
inwards, apex pointed; midrib flattened above; lateral veins 912 pairs, looping near margin,
distinct below, faintly visible above; intercostal veins reticulate, distinct below, almost
invisible above; petioles 815 mm long, drying black. Inflorescences and flowers unknown.
Infructescences almost sessile to 1-cm-stalked. Fruits yellowish green when fresh, black
on drying, oblong, 22.3 x 0.70.8 cm, apex with distinct point.
Distribution. Endemic to Borneo. Uncommon, known by two collections only (SAN 118364
and SAN 118405), from Lanas, Keningau in Sabah.
Ecology. Lowland forest by stream and ridge top. Fruiting in October.
6. Microtropis kinabaluensis Merr. & Freem.

(of Mt. Kinabalu, Sabah)
l.c. 304; Masamune l.c. 418; Ding Hou l.c. (1962) 277. Type: Clemens 29518, British North Borneo,
Mt. Kinabalu (holotype A; isotypes B, BO, GE, K, L, N, UC). Synonyms: M. kinabaluensis var.
acuminata Merr. & Freem. l.c. 305, Masamune l.c. 418; M. sterrophylla Merr. & Freem l.c. 305,
Masamune l.c. 419.
Shrub or small tree to 5 m tall. Incipient bracts pale green, to 1.5 cm long, 0.5 cm broad,
lanceolate or oblong. Young twigs slightly angled. Leaves greenish yellow on drying,
leathery, elliptic-oblong, 1124.5 x 3.59.5 cm; base cuneate to tapered, margin slightly
recurved, apex pointed; midrib raised above; lateral veins 612 pairs, looping near margin,
very faint on both surfaces; intercostal veins reticulate, equally prominent as lateral veins;
petioles 12 cm long, drying wrinkled and yellowish. Inflorescences cymose panicles, to
5.5 cm long, peduncles 1.52.5 cm long. Flowers white; calyx-lobes with 35 longitudinal
veins; petals free, ovate or broadly elliptic; stamen filaments united at base; ovary flaskshaped. Fruits red when ripe, oblong or ovoid to ellipsoid, 1.52 x 0.71 cm, faintly
furrowed, crowned by the persistent style.
Distribution. Endemic to Borneo. Widely distributed in Sabah and Sarawak.
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7. Microtropis ovata Merr. & Freem.

(Latin, ovatus = egg-shaped; the leaf shape)
l.c. 297; Masamune l.c. 419; Ding Hou l.c. (1962) 279. Type: Clemens 40046, British North Borneo,
Mt. Kinabalu (BM, BO).
Shrub to 3 m tall. Twigs 4-angled. Leaves thickly leathery, under surface not papillose nor
pustulate, ovate to ovate-oblong, 4.510 x 27 cm; base rounded to cordate, margin not
curled inwards, apex acute; lateral veins 59 pairs; petioles very short to 2 mm long or
leaves sessile. Flowers in condensed cymes, peduncle very short; calyx-lobes suborbicular;
petals free, ovate or ovate-oblong; stamen filaments united at the lower half; ovary ovoidoblong, not constricted at the middle. Fruits unknown.
Distribution. Endemic to Sabah. Uncommon, known from Mt. Kinabalu only.
Ecology. Submontane forests at 12001500 m.
8. Microtropis platyphylla Merr.

(Greek, platus = broad, phyllon = leaf; broad-leaved)
Philip. J. Sc. 10 (1915) Bot. 319; Ding Hou l.c. (1962) 279. Type: Loher 5779, Philippines (BO, L).
Shrub or small tree to 10 m tall, 10 cm diameter. Incipient bracts brown, 412 mm long,
lanceolate. Leaves thickly leathery, not papillose nor pustulate beneath, elliptic to lanceolate or
ovate-lanceolate, 4.524 x 4.511 cm; base cuneate, margin not curled inwards, apex blunt
to pointed; lateral veins 612 pairs; petioles 12 cm long. Inflorescences paniculate,
peduncles 11.5 cm long. Flowers (4)5-merous; calyx-lobes suborbicular or subreniform
with gnawed margins; petals free, elliptic or broadly ovate, obtuse; stamen filaments united
at base; ovary conical, not constricted at the middle, apex obtuse or notched. Fruits ovoid
or ellipsoid, 12.5 x 1 cm.
Distribution. Borneo and the Philippines. Common in Sabah, but uncommon in Sarawak.
Ecology. Hill and montane forests to 2700 m.
9. Microtropis rigida Ridl.

(Latin, rigidus = stiff; the thick leaves)
l.c. 36; Merr & Freem. l.c. 297; Masamune l.c. 419; Ding Hou l.c. (1962) 278. Type: Beccari PB
1702, Sarawak (holotype FI; isotypes BO, K).
Small tree to 20 m tall, 15 cm diameter. Incipient bracts lanceolate, c. 0.5 cm long, brown,
stiff. Twigs swollen at nodes. Leaves thickly leathery, elliptic-oblong, 712 x 2.56 cm,
pustulate below; base obtuse, rarely cuneate, margin curled inwards, apex pointed; midrib
raised above; lateral veins 57 pairs, very faint; intercostal veins reticulate, visible below,
invisible above; petioles very short to 2 mm long to almost absent. Inflorescences to 2.5 cm
long, peduncles 11.5 cm long. Flowers white; calyx-lobes suborbicular, sometimes
irregularly split; petals free, oblong, obtuse; stamen filaments subulate, lower half united;
145
ovary globose, narrowed towards apex, style very short, stigmas obscurely 46-lobed.
Fruits oblong, 1215 x 79 mm, apex rounded with short persistent style.
Distribution. Endemic to Sarawak; uncommon and known only from Bako National Park,
G. Santubong and Mulu National Parks.
Ecology. Lowland to submontane forests to 1000 m, including kerangas.
10. Microtropis sabahensis Kochummen

(of Sabah)
l.c. (1994) 61. Type: Banang SAN 51915, Sabah (holotype KEP; isotype SAN).
Treelet to 5 m tall. Youngest twigs reddish brown. Incipient bracts greenish, needle-like, c.
5 x 1 mm. Leaves thinly leathery, elliptic, 13.520.5 x 3.57 cm; base cuneate, margin
slightly wavy, apex pointed; midrib raised above; lateral veins 810 pairs, looping and
joining near margin, faint on both surfaces; intercostal veins reticulate, very faint; petioles
11.5 cm long, drying greenish yellow. Inflorescences axillary, 34.5 cm long, cymose
panicles, peduncles 11.5 cm long; bracteoles triangular, transparent. Flowers 4-merous;
sepals ovate, transparent, wrinkled outside; petals united half-way, lobes oblong with
transparent margins; stamens seated on the mouth of corolla-tube, filaments flat; ovary
gradually tapered towards apex, slightly ridged towards apex, style indistinct, stigma distinct,
4-lobed. Fruits unknown.
Distribution. Endemic to Sabah. Uncommon, known only from the type collection from
Kinabatangan.
Ecology. Lowland and seasonal swamp forests. Flowering in June.
Very similar to M. kinabaluensis but differing in the united petals, shape of ovary and the
prominent stigma.
11. Microtropis sarawakensis Kochummen

(of Sarawak)
l.c. (1994) 63. Type: Ilias S. 36501, Sarawak (holotype KEP; isotypes K, L, MO, SAN, SAR).
Small tree to 5 m tall, 10 cm diameter. Incipient bracts green, elliptic or lanceolate, c. 10 x

4 mm. Twigs reddish brown, rounded, youngest ones angular. Leaves thinly leathery, drying
to greenish yellow, elliptic or oblong, 1017 x 7.5 cm; base cuneate, margin wavy and
slightly recurved, apex pointed; midrib raised above; lateral veins 79 pairs, very faint on
both surfaces; intercostal veins invisible to very faint; petioles 11.5 cm long, channelled
above, wrinkled and yellowish on drying. Inflorescences cymose panicles, 25 cm long,
peduncles 11.5 cm long. Flowers 4- or 5-merous; calyx-lobes suborbicular, transparent
with few fine veins in the middle; petals free, imbricate, oblong, transparent; stamens with
short filaments which are joined at the bottom by a staminal ring, connective slightly
146
prolonged; ovary short conical, without distinct style, with wavy surfaced, and unlobed
stigma. Fruits (immature) green when fresh, ovoid, 710 x 45 mm, apex with short style
and stigma, calyx persistent.
Distribution. Endemic to Sarawak. Uncommon, known only by the type collection.
Ecology. Lowland forests by streams. Flowering in April and June, and fruiting in October.
The leaves are somewhat similar to those of M. sabahensis, but the shape and size of
incipient bracts of M. sarawakensis distinguishes it from M. sabahensis. Furthermore, in M.
sabahensis the stigma is distinctly lobed, and the corolla is united almost half way. In
contrast, in M. sarawakensis the stigma is unlobed and the petals are free.
12. Microtropis sumatrana Merr.

(of Sumatra)
Pap. Mich. Acad. Sc. 19 (1934) 164; Merr. & Freem. l.c. 304; Ding Hou l.c. (1962) 279. Type:
Rahmat 214, Sumatra (A, BO).
Small tree to 20 m tall and 20 cm diameter. Bark grey, smooth; inner bark pale grey.
Sapwood pale white. Incipient bracts lanceolate, to 12 mm long. Twigs greyish. Leaves
thinly leathery, pale below, dark brown on drying, not papillose nor pustulate beneath,
elliptic, 6.511 x 3.55.5 cm; base cuneate, apex pointed; midrib flattened above; lateral
veins 57 pairs, faint to distinct below, curving and joining near margin; intercostal veins
laxly reticulate, faint; petioles 0.71 cm long. Inflorescences dichotomously branched,
peduncles 13 cm long. Flowers with calyx-lobes irregularly split; petals free, oblong,
obtuse; stamens united at the lower half; ovary gradually narrowed towards apex, stigma 4lobed. Fruits oblong, 1.51.7 x 0.71.0 cm, with pointed tip, calyx persistent at base.
Distribution. Sumatra and Borneo. Uncommon, in Sabah known only from Sipitang,
Tawau, and Mostyn.
147
148
13. Microtropis valida Ridl.

(Latin, validus = robust-growing)
J. Str. Br. R. As. Soc. 75 (1917) 19, l.c. (1922) 445; Merr & Freem. l.c. 304; Ding Hou l.c. (1962)
276; Kochummen & Whitmore l.c. 170. Type: Curtis 1331, Perak (holotype K; isotype BO).
Synonyms: M. bicolor Merr. & Freem. l.c. 298; M. pauciflora Boerl. ex Merr. & Freem. l.c. 303.
Shrub or small tree to 5 m tall. Incipient bracts brown, stiff, 312 mm long. Leaves
leathery, under surface not papillose nor pustulate, ovate to lanceolate or elliptic, 1127 x
4.512 cm; base rounded to cuneate, margin not curled inwards, apex acute or acuminate;
midrib raised above; lateral veins 614 pairs; intercostal veins reticulate, distinct on both
surfaces; petioles 12 cm long. Inflorescences usually paniculate, rarely a simple cyme or
fascicle, peduncles 14 cm long. Flowers 4(5)-merous; calyx-lobes suborbicular, slightly
wrinkled outside, margin transparent; petals free, elliptic or oblong, obtuse; stamen filaments
united at base; ovary cylindric, slightly constricted at the middle, truncate or discoid at the
top. Fruits ellipsoid, c. 2 x 1 cm, apex pointed and furfuraceous with persistent style.
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah and Sarawak uncommon,
known by only two collections (SAN 75680 and SAN 44667) in Sabah, and three (including
the type of M. bicolor by Beccari) collections in Sarawak.
14. Microtropis wallichiana Wight ex Thwaites

(N. Wallich, 17861854, Superintendent, Calcutta Botanic Gardens)
En. Pl. Zeyl. (1858) 71; Ding Hou l.c. (1962) 279. Type: Wight 528, Ceylon (BO, K). Synonym: M.
suborbiculata Merr. & Freem. l.c. 282, Masamune l.c. 419.
Shrub or small tree. Incipient bracts brown, 38 mm long. Twigs rounded to slightly angled.
Leaves thickly leathery, elliptic, lanceolate or rounded, 318 x 16 cm, lower surface not
papillose nor pustulate; base cuneate to rounded, margin not curled inwards, apex pointed,
blunt or rounded and notched; midrib raised above; lateral veins 49 pairs, invisible above;
intercostal veins reticulate, faint to invisible; petioles to 1.5 cm long, rarely very short.
Inflorescences condensed cymes, less than 1 cm long, peduncles very short or absent.
Flowers yellowish; calyx-lobes subreniform; petals free, fleshy; stamen filaments united at
the lower part; ovary conical, longitudinally striate, stigma not lobed. Fruits oblongellipsoid or oblong-obovoid, c. 1 x 0.5 cm, furfuraceous, crowned by persistent style.
Distribution. Sri Lanka, Sumatra and Borneo. In Sabah, known from Mt. Kinabalu only;
not recorded from Sarawak.
Ecology. Submontane to montane forests at 12502500 m.
The types of M. wallichiana and M. suborbiculata have some differences which require to
be substantiated by examining more material. There is no conclusive evidence for their
distinction, although it is doubtful if they are the same species.
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8. PERROTTETIA Kunth
(G.S. Perrotet, 17931870, French botanist)
Nov. Gen. Sp. 7 (1824) 73; Merrill l.c. (1921) 354; Ridley l.c. (1922) 454; Masamune l.c. 419; Ding
Hou l.c. (1962) 288; Backer & Bakhuizen f. l.c. 55; Kochummen & Whitmore l.c. 171; Anderson l.c.
161; Ashton l.c. 112. Synonym: Caryospermum Blume l.c. (1850) 175.
Shrubs or small trees. Leaves alternate. Inflorescences cymose, axillary. Flowers bisexual,
rarely unisexual; calyx-lobes 5 or 4; petals similar to calyx-lobes in size and shape;
stamens 5 or 4, inserted on the margin of the disc; ovary semi-inferior, mostly 2-celled,
style 1; ovules two in each cell. Fruit a berry, globose, 24-seeded. Seeds covered with thin
aril; endosperm thin; embryo small.
Distribution. About 15 species; China, Formosa, Malesia and America. Only one species
Ecology. Primary and secondary hill and montane forests to 2600 m.
Perrottetia alpestris (Blume) Loes.
Fig. 8.
(Latin, alpestris = growing above the limit of forest growth; its habitat)
in Engler & Prantl l.c. 220; Merrill l.c. (1921) 354; Ridley l.c. (1922) 454; Masamune l.c. 419; Ding
Hou l.c. (1962) 288; Kochummen & Whitmore l.c. 171; Ashton l.c. 161. Basionym: Celastrus
alpestris Blume, Bijdr. (1826) 1145. Type: Blume, s.n., Java (BO, L).
subsp. philippinensis (Vidal) Ding Hou

l.c. (1962) 291. Basionym: Caryospermum philippinensis Vidal, Rev. Pl. Vasc. Philip. (1886) 89.
Type: Beccari PB 7770, Sarawak (FI).
Shrub or small tree to 12 m tall. Young twigs and petioles purplish when fresh, blackish on
drying. Leaves thinly leathery, elliptic or oblong, 1020 x 48 cm; base cuneate or rounded,
sometimes subcuneate, margin prominently toothed, apex pointed; midrib flattened above;
lateral veins 812 pairs, raised below, sunken or flattened above; intercostal veins reticulate,
distinct below, faint above; petioles 715 mm long, not swollen. Flowers white or light
greenish, 4-merous; calyx-lobes sparsely puberulous on both surfaces; petals puberulous on
both surfaces; disc cup-shaped; stamen filaments longer than the disc; ovary conical, style
short, stigma distinctly 2-lobed. Fruits globose or subglobose, 2.53.5 mm diameter, red.
Seeds 34.
Distribution. Borneo, Philippines and Celebes. In Sabah, collected from Kinabalu National
Park, Kalabakan and Lamag. In Sarawak, reported from Dulit Range, Mulu National Park,
Bt. Rawan, Kapit and Lubuk Antu.
Ecology. Hill and montane forests to 2700 m, in open screes, usually on moist soils.
This subspecies differs from the other two (alpestris and moluccana) by its 4-merous
instead of 5-merous flowers.
150
151
152
9. SARAWAKODENDRON Ding Hou

(Greek, dendron = a tree; Sarawak tree)
Blumea 15 (1967) 139, l.c. (1969) 97, l.c. (1972) 930; Anderson l.c. 162.
Small trees. Young twigs, petioles, midrib, leaf blades and floral parts with yellow pustules.
Twigs without distinct stipular scars. Leaves alternate, entire; petioles not swollen. Stipules
inconspicuous. Inflorescences axillary, with condensed decussate bracts. Flowers bisexual;
calyx 5-lobed, lobes imbricate; petals 5, imbricate; disc slightly 5-angular; stamens 3,
inserted at the base of ovary, anthers extrorse, transversely dehiscent; ovary semi-inferior,
3-celled, 3-angled, stigma 3, ovules 8 in each cell, in 2 series, placentation axile. Fruit a
capsule, dehiscing loculicidally into 3 valves. Seeds 68 in each cell, with endosperm; aril
fleshy, cushion or caruncle-like at the chalazal end which bears 1.52-cm-long simple or
dichotomously branched, thread-like appendages arising from the base of funicle.
Distribution. A monotypic genus, endemic to Sarawak.
Sarawakodendron filamentosum Ding Hou
Fig. 9.
(Latin, filamentosus = thread-like; the appendages of the seed aril)

l.c. (1967) 141, l.c. (1969) 103, l.c. (1972) 932. Type: Ding Hou 333, Sarawak (holotype L; isotypes
BO, SAR).
Small tree to 10 m tall, 15 cm diameter. Leaves drying to greenish yellow, elliptic to oblong
or oblanceolate, 10.529 x 410.5 cm; base cuneate, apex pointed; midrib flattened above;
lateral veins 48 pairs, faintly raised on both surfaces; intercostal veins reticulate, faintly
visible below; petioles 0.51.5 cm long. Inflorescences 12.5 cm long, unbranched, fewflowered, peduncles to 1.3 cm long, pedicels 1.52 cm long, articulated near base. Flowers
pale orange, floral parts fleshy; calyx-lobes semi-orbicular, slightly toothed along the
margin; petals suborbicular with distinct reticulate venation and light brownish stripes or
dots; disc flat, slightly 5-angular; stamens reflexed at anthesis. Fruits narrow ellipsoid, 3angled, 68.5 x 23.5 cm, gradually narrowed towards both ends. Seeds 68 in each cell, 2
2.5 x 0.5 cm; embryo narrow-lanceolate, c. 18 x 4 mm; cotyledons foliaceous, free.
Distribution. Endemic to Sarawak. Uncommon, known by the following few collections:
Ding Hou 133, S. 24506, S. 24897 and S. 24898 from the Nyabau FR, Bintulu; S. 18685
from Bako National Park; S. 42715 from Sri Aman, and S. 51508 from Bayai, 2nd Div.
Ecology. In mixed dipterocarp forest on yellow sandy humult ultisol soils and in kerangas
forest.
The genus has similar floral characters as that of Salacia and in fruit characters it has
similarities with Kokoona and Lophopetalum.
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10. SIPHONODON Griff.

(Greek, siphonos = tube, odontos = tooth;
the hollow, columnar, toothed upper part of the ovary)
Calc. J. Nat. Hist. 4 (1844) 246; Ding Hou, Blumea 12 (1963) 36, l.c. (1964) 394; Backer &
Bakhuizen f. l.c. 56l; Kochummen & Whitmore l.c. 171; Cockburn l.c. 62; Anderson l.c. 162; Ashton
l.c. 112; Whitmore, Tantra & Sutisna l.c. 45. Synonym: Capusia Lecompte, Bull. Mus. Hist. Nat.
Paris 32 (1926) 95.
Trees. Leaves spiral or alternate. Inflorescences axillary, cymose, sometimes only oneflowered. Flowers 5-merous; calyx-lobes imbricate; petals imbricate, larger than calyxlobes; stamens 5, sometimes alternating with 5 staminodes, united at lower part, anthers
latrorse, connective distinct and broad; ovary semi-inferior, 3-celled, upper half hollow and
with a style-like column arising from the bottom, ovules 1 in each cell, oblique or
pendulous. Fruits drupaceous, with numerous bony 1-seeded pyrenes. Seeds flat, with
endosperm; cotyledons flat, free.
Distribution. 7 species; SE Asia through Malesia to Australia; one species in Sabah and
Sarawak.
Fig. 10.
Siphonodon celastrineus Griff.

(resembling Celastrus)
l.c. 247; Ding Hou l.c. (1964) 394; Backer & Bakhuizen f. l.c. 561; Kochummen & Whitmore l.c.
171; Cockburn l.c. 62; Anderson l.c. 162; Ashton l.c. 112; Whitmore, Tantra & Sutisna l.c. 45. Type:
Griffith 9019, Penang (BO, K). Synonyms: S. pyriformis Merr. l.c. (1908) 240; Xanthophyllum
subglobosum Elmer l.c. (1913) 1676.
Tree to 35 m tall, 70 cm diameter; buttresses to 2 m high. Bark usually grey-brown,
smooth to scaly; inner bark yellowish. Sapwood pale. Twigs brownish, zig-zag. Leaves
drying greenish yellow, lanceolate to elliptic or oblong, 815 x 3.57 cm; base cuneate,
margin toothed, apex pointed; midrib flattened above, yellowish below on drying;
lateral veins yellowish on drying; intercostal veins reticulate, distinct below, faint above; petioles
58 mm long, drying yellowish. Flowers cream-white; calyx-lobes almost rounded, 12
mm long; petals ovate with obtuse apex; stamens c. 1 mm long, filaments flat, united near
base; ovary subglobose to conical, occasionally with 5 ridges towards the apex. Fruits
subglobose to obovoid, 36.5 x 26 cm.
Distribution. India, Burma, Thailand, Indo-China, and Malesia. In Sabah, recorded from
Lahad Datu, Ranau, Sandakan and Tawau. In Sarawak, collected from Belaga, Bau, Bt.
Mersing, and Ulu Melinau.
Ecology. Mixed dipterocarp forests on fertile clay soils, especially near limestone and on
basic volcanic rock. Uncommon. Fruiting in MarchNovember.
This species can be confused with those of Xanthophyllum (Polygalaceae) but the toothed
leaves at once distinguish this.
154
CHRYSOBALANACEAE
Ghillean T. Prance
Royal Botanic Gardens, Kew, England
R. Brown in Tuckey, Narr. Exp. Congo (1818) 433; King, J. As. Soc. Beng. 66, 2 (1897) 275 (under
Rosaceae); Ridley, FMP 1 (1922) 665 (under Rosaceae); Merrill, EB (1921) 287, PEB (1929) 92
(under Rosaceae); Masamune, EPB (1942) 324 (under Rosaceae); Backer & Backhuizen f., FJ 1
(1964) 521 (under Rosaceae); Prance & Whitmore, TFM 2 (1973) 321 (under Rosaceae); Anderson,
CLTS (1980) 292 (under Rosaceae); Cockburn, TS 2 (1980) 79 (under Rosaceae); Corner, WSTM 2
(1988) 615 (under Rosaceae); Prance & White, Phil. Trans. Roy. Soc. B. 320 (1988) 1; Prance, FM 10,
4 (1989) 635; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 298.
Trees or shrubs. Leaves simple, entire, alternate, often coriaceous. Stipules small and
caducous to large and persistent. Inflorescences racemose, paniculate or cymose; flowers
bracteate and usually 2-bracteolate. Flowers actinomorphic (with a regular symmetry) to
zygomorphic (with an irregular symmetry), bisexual or unisexual (in Parastemon, plant
sometimes polygamo-dioecious), markedly perigynous; receptacle short to elongate; disc
always present, forming a lining to the receptacle; calyx-lobes 5, imbricate, often unequal,
erect or reflexed; petals 5, imbricate, usually caducous; stamens variable, 260, inserted on
the margin of the surface of the disc, or basally adnate to it, forming a complete circle or, in
zygomorphic flowers, unilateral, all fertile or some without anthers and then often reduced
to small staminodes; filaments filiform, free, connate at the base or ligulately connate,
included to far exserted; anthers small, dorsifixed, longitudinally dehiscent; gynoecium
(ovary) basically of three carpels and gynobasic (inferior), but usually with only one carpel
fully developed, attached to the base, middle or mouth of the receptacle-tube, sessile or with
a short gynophore (stalk), pubescent or villous; carpel unilocular with two ovules or
bilocular with one ovule in each locule, style filiform, arising from the receptacle at the
base of the carpel, stigma distinctly or indistinctly 3-lobed, ovules erect, epitropous (with
micropyle directed towards the base), tenuinucellate (with thin nucellus). Fruit a dry or
fleshy drupe; endocarp various, thick or thin, fibrous or bony, often with a special
mechanism for seedling escape, often densely hairy inside. Seeds erect, almost without
endosperm; cotyledons plano-convex, fleshy, sometimes ruminate.
Distribution. A pantropical family of 17 genera and 520 species, of which over 400 in the
Neotropics, 60 in Africa and 39 in Asia and the Pacific. In Sabah and Sarawak, the family is
represented by 6 genera and 15 species all of which are trees.
Ecology. Found mainly in lowlands both in swamp and well-drained forests, to 1500 m. The
flowers are pollinated mainly by insects and in a few cases by bats, and the fruits are
dispersed by birds, mammals and water.
Uses. The wood is extremely hard and difficult to cut due to the presence of silica; therefore
it is resistant to decay and to marine borers and is used for posts and marine piles. The wood
of some species is used for general construction and firewood. The fruit of some species is
155
edible. The fruit of Atuna racemosa subsp. racemosa is used as a putty for caulking boats
and the oil from this species is used in the Pacific in hair dressings.
Taxonomy. Although the family Chrysobalanaceae was described by Robert Brown in 1818
(In: J.K. Tuckey, Narrative of an expedition to explore the River Zaire, Appendix 5: 433
434) it has been placed as a tribe or subfamily of Rosaceae in most of the older and widely
used systems of classification such as those of Bentham and Hooker, Engler and Prantl and
Hutchinson. This is in marked contrast to the opinions of nearly all workers with a detailed
knowledge of the group, especially of its anatomy. The Chrysobalanaceae differs from the
Rosaceae in the gynobasic style, the basal, erect ovules which are tenuinucellate, in the
presence of abundant deposits of silica in the wood, stem and leaf and in many wood
anatomical characters such as the large oblique vessels, the banded oblique vessels, the
banded parenchyma which is composed of long strands. The authors of modern evolutionary
systems such as Cronquist, Thorne and Takhtajan consider the Chrysobalanaceae to be a
separate family but still placed in the Rosales. Dahlgren and Thorne (Ann. Missouri Bot.
Gard. 71 (1984) 633699) suggested a relationship between Chrysobalanaceae and Myrtales.
Although the resemblances between Rosaceae and Chrysobalanaceae are superficial, the
family is best left in the Rosales until further evidence such as molecular studies show that it
should be placed elsewhere. Further details about the taxonomy of the group can be found in
Prance and White's paper (l.c.).
Key to genera
(based on flowers)
1.
Flowers actinomorphic with ovary inserted in centre of receptacle; stamens included...2

Flowers zygomorphic with ovary inserted laterally on receptacle; stamens exserted
(except in Parinari).....................................................................................................3
2.
Stamens 710, all fertile..............................................................................3. Licania

Stamens 5, all fertile or 2 fertile and 3 sterile..........................................5. Parastemon
3.
Stamens 610, included. Leaves usually with stomatal crypts.......................6. Parinari

Stamens 1040, exserted. Leaves never with stomatal crypts........................................4
4.
Stamens united into a ligule. Ovary unilocular..................................2. Kostermanthus

Stamens free to base. Ovary bilocular...........................................................................5
5.
Stamens 1020. Inflorescence a raceme or sparsely branched contracted panicle.....1.

Atuna
Stamens 2540. Inflorescence a many-flowered corymbose panicle..........4. Maranthes
Key to genera
(based on fruits)
1.
Fruits dehiscing by two lateral plates........................................................................2

Fruits indehiscent or dehiscing by two basal stoppers...................................................3
156
CHRYSOBALANACEAE (PRANCE)
2. Fruits bilocular (sometimes only 1 locule developing), 34 cm long.........4. Maranthes

Fruits unilocular, usually 11.5 cm long (2.53 cm in P. grandifructus)........5. Parastemon
3.
Fruits dehiscing by two basal stoppers; exocarp lenticellate. Leaves usually with
stomatal crypts beneath................................................................................6. Parinari
Fruits indehiscent or opening by longitudinal lines. Leaves without stomatal crypts.....4
4.
Fruits 11.3 cm long, unilocular, breaking open by longitudinal lines of weakness;

exocarp smooth.......................................................................................................3. Licania
Fruits 36 cm long, uni or bilocular, indehiscent; exocarp verrucose-crustaceous.........5
5.
Fruits unilocular. Stamens united into a ligule..................................2. Kostermanthus

Fruits bilocular. Stamens free...............................................................................1. Atuna
1. ATUNA Rafin.
(an Amboinese vernacular name)
merbatu (Malay)
Sylva Tellur. (1838) 153; Kostermans, Reinwardtia 7, 5 (1969) 421; Prance & Whitmore l.c. 323;
Anderson l.c. 292; Cockburn l.c. 81; Prance l.c. (1989) 665; Whitmore, Tantra & Sutisna l.c. 298.
Synonyms: Cyclandrophora Hassk., Flora Beibl. 1 (1842) 47; Parinari subg. Cyclandrophora
(Hassk.) Blume, Mlang. Bot. 2 (1855) 10; Parinari auct. non Aubl., Ridley l.c. (1922) 666.
Small to large trees, ultimate shoots with complicated system of divaricate branching.
Stipules large, prominently keeled, lateral, persistent or subpersistent. Leaves almost glabrous
on both surfaces, often with minute papillae on venation giving beaded appearance, without
stomatal crypts, with a pair of glands on midrib at or near base of lower surface; petioles
eglandular. Inflorescence a raceme, or sparsely branched, contracted panicle; bracts and
bracteoles persistent, eglandular, not enclosing groups of flower-buds. Flowers bisexual;
receptacle obconical to cylindrical, as long as or exceeding calyx-lobes, hollow, hairy
inside throughout, throad blocked by retrorse hairs; calyx-lobes 5, broadly ovate to
lanceolate, tomentellous on both surfaces; petals 5, glabrous, exceeding calyx-lobes; stamens
1020, posterior, inserted unilaterally on margin of disc, filaments free, exserted; staminodes
forming a barely visible denticular margin to throat; ovary inserted at mouth of receptacle
tube, pilose on exterior, carpel bilocular with 1 ovule in each loculus. Fruits large; epicarp
glabrous, densely verrucose-crustaceous; mesocarp transversely fibrous; endocarp hard,
thick, shortly and sparsely hairy inside, breaking up irregularly at germination; cotyledons
large and strongly ruminate; germination cryptocotylar (with hidden cotyledon), eophylls
alternate.
Distribution. 8 species from Southern India throughout Malesia and to Fiji and Samoa in
the Pacific. 3 species occur in Sabah and Sarawak.
Ecology. Usually found on well-drained mixed dipterocarp forest to 600 m.
Uses. Little used as a timber.
157
Key to Atuna species

1.
Leaves broadly ovate, thickly coriaceous, cordate at base. Stamens c. 10...1. A. cordata
Leaves elliptic or oblong-lanceolate, chartaceous, rounded at base. Stamens 1320......2
2.
Receptacle tube narrowly cylindrical. Leaves oblong-lanceolate. Stamens purple...........

..............................................................................................................2. A. nannodes
Receptacle turbinate-campanulate. Leaves elliptic to oblong-ovate. Stamens white........
..............................................................................................................3. A. racemosa
1. Atuna cordata Cockburn ex Prance
Fig. 1.
(Latin, cordatus = heart-shaped; the leaf base)

Brittonia 39 (1987) 364; Cockburn l.c. 82; Prance l.c. (1989) 667. Type: Ahmad Talip SAN 47687,
Sabah, Lahad Datu (holotype K; isotypes L, SAN).
Tree to 40 m tall, the trunk often with thick buttresses. Bark smooth, grey-green, mottled
white; inner bark hard, reddish brown. Sapwood ochre to red-brown, hard. Young branches
glabrescent, inconspicuously lenticellate. Stipules to 1.7 cm long, very early caducous.
Leaves coriaceous, broadly ovate, 4.512 x 39.5 cm; cordate at base, abruptly acuminate
at apex, the acumen 13 mm long; glabrous and shiny above, glabrous beneath; midrib
prominulous above, prominent beneath; lateral veins 912 pairs, lightly prominulous above,
prominulous and glabrous beneath; petioles 13 mm long, thick, glabrous. Inflorescences
of terminal and subterminal racemes, 48 cm long, borne in single or more often in paired
branches, densely tomentellous on exterior, puberulous within; bracts and bracteoles ovate,
tomentellous, early caducous. Flowers with receptacles 57 mm long, conical to
campanulate, tomentellous on exterior, sessile; calyx-lobes slightly unequal, tomentellous
on both surfaces; petals c. 7 mm long, obovate, glabrous; stamens 10, inserted on one side
of ring, the filaments 1012 mm long; ovary densely pilose, style slender, hirsutulous on
lower portion. Fruits c. 6 x 5 cm, ovoid; epicarp verrucose-crustaceous; mesocarp c. 5 mm
thick, fibrous, hard; endocarp thin.
Distribution. Endemic to Sabah; recorded from Lahad Datu, Karamuak, Mt. Tawai and Mt.
Silam.
Ecology. Confined to hills and locally common on ultramafic rock to 1200 m.
2. Atuna nannodes (Kosterm.) Kosterm.

(Greek, nanno = dwarf; the comparatively small size of the tree)
Reinwardtia 7, 5 (1969) 422; Prance & Whitmore l.c. 325; Prance l.c. (1989) 667. Basionym: Parinari
nannodes Kosterm., Reinwardtia, 7, 1 (1965) 50. Type: Beccari PB 2955, Sarawak, Mt. Matang (BO,
K). Synonym: Cyclandrophora nannodes (Kosterm.) Kosterm. & Prance, Candollea 20 (1965) 122.
Tree to 20 m, usually smaller, unbuttressed. Bark smooth, dark grey. Young branches
sparsely appressed hirsutulous-strigose, soon glabrous, obscurely lenticellate. Stipules narrowly
lanceolate, acute, 612 mm long, strigose to glabrous, subpersistent. Leaves thinly coriaceous,
oblong-lanceolate, 6.719 x 2.56 cm, glabrous on both surfaces, sometimes slightly
158
159
bullate above; rounded at base, long slender acuminate at apex, the acumen 722 mm long;
midrib prominulous above, prominent beneath; lateral veins 1012 pairs, arcuate, prominulous
on both surfaces or sometimes prominent beneath; petioles 24 mm long, glabrescent,
eglandular, the lower part swollen, usually curved. Inflorescences axillary racemes, 37 cm
long, the rachis densely sericeous-tomentellous; bracts and bracteoles lanceolate, 37(13)
mm long, persistent, sericeous. Flowers with cylindrical receptacle, 813 mm long, densely
sericeous on exterior, sessile; calyx-lobes to 6 mm long, unequal, acute, sericeous on
exterior; petals white, spathulate to ovate, 812 mm long, narrowed to the base; stamens 18
20, black to purple, the filaments 1015 mm long, slightly unilateral with tooth-like
staminodes opposite; ovary pilose, style to 15 mm long, glabrous, stigma capitate. Fruits
ellipsoid, 34 x 1.5 cm, slightly tapered to the base, crustaceous-verrucose on exterior;
mesocarp 22.5 mm, fibrous, hard; endocarp thin.
Distribution. Peninsular Malaysia, Borneo (Sabah, Sarawak, Kalimantan). In Sabah known by
collections from localities in the Sandakan, Semporna, Tawau and Tongod districts. In
Sarawak, recorded from Bau and Kuching areas, 1st Div.
Ecology. Forest on well-drained soils to 500 m.
3. Atuna racemosa Rafin.

(Latin, racemosus = with a raceme-like inflorescence)
l.c. 153; Kostermans l.c. (1969) 422; Anderson l.c. 293; Cockburn l.c. 84; Prance l.c. (1989) 669.
Type: Rumphius Herb. Amb. 1 (1741) pl. 66.
Tree to 35 m tall often with short buttresses. Bark smooth, grey to black, mottled; inner bark
orange-brown, red-brown to brown, hard. Sapwood white, ochre to red-brown; heartwood redbrown. Young branches glabrous, dark red-brown, smooth or obscurely lenticellate when
dry. Stipules lanceolate, stiff, to 820 mm long, acute, glabrous to strigose, sub-persistent.
Leaves usually chartaceous, more rarely stiffly coriaceous, broadly ovate, elliptic, ovateoblong, oblong or even lanceolate, 4.525(35) x 211 cm; rounded, subcordate or subcuneate
at base, acuminate at apex, the acumen 325 mm long; glabrous on both surfaces when
mature, sometimes sparsely strigose beneath on lower portion when young; midrib
prominent on both surfaces; lateral veins 913 pairs, prominulous above, prominent
beneath, straight or arcuate; the venation conspicuously papillose and often giving leaf a
scabrous or beaded appearance; petioles slender or thick, 37 mm long, pilose or puberulous,
glabrescent or glabrous. Inflorescences of axillary racemes or little-branched with 23
racemose branches on short main peduncle, 515 cm long, the rachis and branches
tomentellous or densely short sericeous; bracts and bracteoles ovate to oblong, acute, 38
mm long, persistent or caducous. Flowers with receptacle turbinate-campanulate, 410 mm
long, tomentose to sericeous on exterior; pedicels 0.51 mm long; calyx-lobes ovate to
ovate-oblong, 47 mm long, densely tomentellous on both surfaces or sericeous on exterior,
tomentellous within; petals equal, ovate-oblong, c. 10 mm long, blue or white, caducous;
stamens 1320, pale blue, 815 mm long with tooth-like staminodes opposite; ovary pilose
to densely villous, style equalling filaments, glabrous above, stigma small. Fruits ellipsoid,
subglobose to slightly pyriform, 57.5 x 3.54.5 cm; epicarp crustaceous-verrucose;
mesocarp fibrous, 511 mm thick; endocarp thin, 13 mm, densely pilose within.
160
Key to subspecies
Leaves 1025 (35) cm long, usually elliptic, oblong or lanceolate but sometimes ovate,
chartaceous or thickly coriaceous, the apex long finely acuminate, acumen 625 mm long;
petioles thick. Flowers 1017 mm long. Medium-sized to large tree often with fluted bole....
subsp. racemosa
Synonyms: Cyclandrophora glaberrima Hassk. l.c. 47; Parinarium glaberrimum (Hassk.) Hassk.,
Tijd. Nat. Ges. Phys. 10 (1843) 147, Merrill l.c. (1921) 290, l.c. (1929) 92, Masamune l.c. 321; P.
scabrum Hassk. l.c. (1843) 147; P. elatum King l.c. 280; C. elata (King) Prance in Kosterm.,
Candollea 20 (1965) 122; Atuna elata (King) Kosterm. l.c. (1969) 421.
Thailand to Sumatra, Peninsular Malaysia, Singapore, Borneo (Sabah, Brunei, Sarawak),

Philippines, Celebes, Moluccas, New Guinea, New Britain, and the Pacific (Admiralty,
Caroline, Solomon Islands, Fiji, Samoa). In Sabah common and has been recorded in
most districts. In Sarawak uncommon and collected only from Baram district, and
known from ecological plots at Bt. Raya, Rejang, Bt. Iju, Balingian, and Bintulu. Found
mainly in well-drained mixed dipterocarp forests, to 600 m, but also occurs on
riverbanks and freshwater and brackish water swamps, can be very common.
Leaves 4.512 cm long, usually ovate or oblong-ovate, subcoriaceous or coriaceous, the
apex bluntly acuminate, acumen 310 mm long; petioles thin. Flowers 811 mm long. Large
trees with cylindrical bole...........................................................................................
subsp. excelsa (Jack) Prance
l.c. (1989) 670. Basionym: Petrocarya excelsa Jack, Mal. Misc. 2 (7) (1822) 66. Type:
Kostermans & Anta 1136, Bangka, Lobok Besar (neotype K; isoneotypes A, BO, L, SING).
Synonyms: Parinarium asperulum Miq., Fl. Ind. Bat., Suppl. (1861) 307; Cyclandrophora
asperula (Miq.) Prance ex Kosterm., Candollea 20 (1965) 130; Cyclandrophora excelsa (Jack)
Kosterm., Candollea 20 (1965) 128; Atuna excelsa (Jack) Kosterm. l.c. (1969) 422; P. villamilii
Merr., Philip. J. Sc. 10 (1915) Bot. 308, Anderson l.c. 294; C. villamilii (Merr.) Prance ex
Kosterm., Candollea 20 (1965) 126; A. villamilii (Merr.) Kosterm. l.c. (1969) 422.
Sumatra, Peninsular Malaysia, Java, Borneo (Sabah, Sarawak, Kalimantan), N Celebes. In

Sabah known from Beaufort, Lahad Datu, Lamag, Sandakan and Tawau districts. In
Sarawak has been recorded in Lundu, Kuching, Serian, Miri and Kapit districts. Usually
occurring in well-drained mixed dipterocarp forest, to 750 m, on ridges and hillsides,
uncommon.
Uses. The fruit (cotyledon) of subsp. racemosa is grated and made into a putty for caulking
canoes, widely used in the Pacific islands. An oil extracted from the seeds is used variously
in different areas, e.g., to scent coconut oil and for hairdressing. The leaves are used to
thatch the outside walls of houses in Fiji. The wood is used locally for posts and poles but is
not of good quality.
2. KOSTERMANTHUS Prance
(A.J.G.H. Kostermans, 19071994, botanist of the
Forest Research Institute and Herbarium Bogoriense, Bogor, Indonesia)
Brittonia 31 (1979) 91; Prance & Whitmore l.c. 327; Anderson l.c. 293; Cockburn l.c. 85; Prance l.c.
(1989) 675; Whitmore, Tantra & Sutisna l.c. 299. Synonyms: Acioa auct. non Aubl., Kostermans,
Reinwardtia 7, 1 (1965) 9; Parinari auct. non Aubl.: quoad P. heteropetala et P. myriandra, tantum.
161
162
Large trees, ultimate shoots not divaricate. Stipules to 7 mm long, foliaceous, persistent,
lanceolate to ovate. Leaves glabrous on both surfaces with minute papillae on veins giving a
beaded appearance; petioles eglandular. Inflorescence an unbranched or little-branched

terminal or axillary raceme with shortly stalked congested cymules proximally and singly
inserted flowers distally; bracts and bracteoles small, suborbicular, persistent, eglandular,
not enclosing groups of flower buds. Flowers bisexual, strongly zygomorphic; receptacle
broadly obconic-campanulate, shorter than calyx-lobes, asymmetric, hollow, hairy on both
surfaces, but throat not blocked by retrorse hairs; calyx-lobes 5, markedly unequal,
suborbicular to ligulate, strongly imbricate; petals 5, unequal in size and shape, the 2
posterior larger than the others, markedly ungulate and enclosing stamens in bud; stamens
830, inserted unilaterally on margin of disc; filaments united for half to three quarters of
length into a strap; staminodes 58, inserted opposite stamens; ovary inserted laterally at
mouth of receptacle, unilocular with 2 ovules. Fruits large, hard; epicarp glabrous, crustaceousverrucose; endocarp hard, thick, glabrous within, breaking irregularly on germination.
Cotyledons slightly ruminate.
Distribution. 2 species, one (K. malayanus) is confined to Peninsular Malaysia, the other
(K. heteropetalus) is known from Sumatra, Borneo, the Philippines, and Celebes.
Kostermanthus heteropetalus (Scort. ex King) Prance
Fig. 2.
(Greek, hetero = uneven, petalon = petal; the unequal petals)

l.c. (1979) 91, l.c. (1989) 677; Anderson l.c. 293; Cockburn l.c. 85; Prance & White l.c. (1988) 152;
Whitmore, Tantra & Sutisna l.c. 299. Basionym: Parinari heteropetalum Scort. ex King l.c. 283.
Type: Scortechini 2040, Perak (BO, SING). Synonyms: Parinarium myriandrum Merr. l.c. (1929) 93;
Acioa heteropetala (Scort. ex King) Kosterm., Reinwardtia 7, 1 (1965) 11.
Tree to 35 m tall, 2 m diameter, often with low small buttresses. Bark with small fissures
and brittle flakes, grey-brown; inner bark pale red-brown to ochre. Sapwood pale yellow;
heartwood red-brown with a wavy junction between it and sapwood. Young branches
glabrous, lenticellate. Stipules 67 mm long, partly intrapetiolar, carinate, ovate, foliaceous,
acute to acuminate, persistent to subpersistent. Leaves coriaceous, usually elliptic-subovate
to rarely lanceolate, 520 x 2.56 cm; cuneate to rounded at base, bluntly acuminate at apex;
glabrous on both surfaces, minutely papillose on venation of both surfaces giving a beadlike appearance; midrib prominulous above, prominent beneath; lateral veins 610 pairs,
arcuate, slender, prominent beneath; petioles 612 mm long, sometimes lightly alate from
decurrent leaf margins, slightly flattened above, eglandular. Inflorescences little-branched,
to 10 cm long, the rachis and branches lightly tomentellous; bracts and bracteoles ovate, acute,
to 3 mm long, caducous. Flowers with a receptacle broadly campanulate, 23 mm long,
tomentose on both surfaces; calyx-lobes fleshy, unequal, acute, to 7 mm long, pilose on both
surfaces, reflexed in open flowers; petals white-tinged pink, fleshy, elliptic, concave, largest
up to 15 mm long, tomentellous on exterior, enveloping staminal ligule, the others much
smaller to 6 mm long; stamens 2530 united into a unilateral ligule for two thirds of their
length, to 12 mm long, glabrous; anthers pubescent; ovary densely pilose, style densely
appressed pilose, stigma truncate. Fruits ovoid, unilocular, c. 4 x 3 cm; epicarp glabrous,
crustaceous; endocarp hard, thick. Cotyledons slightly ruminate, c. 1.5 x 3 cm.
163
164
Vernacular names. Sabah and Sarawakmerbatu, rasak batu (Malay).

Distribution. Sumatra, Peninsular Malaysia, Borneo (Sabah, Sarawak, Brunei and
Kalimantan), the Philippines (Mindanao), and Celebes. In Sabah found in Beaufort,
Keningau, Kota Marudu, Mostyn, and Tongod districts in the west, and in Lahad Datu,
Sandakan, and Tawau districts in the east. In Sarawak widespread.
Ecology. Mixed dipterocarp forest on sandy clay soils, to 500 m.
Uses. The fruit is eaten in Celebes and Sumatra. The timber is easy to cut and is red when
freshly cut turning brown with age. It is little used because of its tendency to rot.
3. LICANIA Aubl.
(intended to be an anagram of the local name in French Guianacaligni)
Hist. Pl. Guiane Fr. 1 (1775) 119; Prance & Whitmore l.c. 328; Anderson l.c. 293; Cockburn l.c. 86;
Prance l.c. (1989) 645. Synonym: Angelesia Korth., Ned. Kruidk. Arch. 3 (1854) 384.
Small to large trees. Stipules small, free, caducous. Leaves glabrous on both surfaces,
without stomatal crypts; petioles eglandular. Inflorescence a panicle of cymules; bracts and
bracteoles to 1.5 mm long, membranous, eglandular, not enclosing groups of flower buds.
Flowers bisexual; receptacle campanulate, slightly asymmetric, tomentose on exterior and
interior; calyx-lobes 5, acute, unequal; petals 5, small, not exceeding the calyx-lobes, not
clawed; stamens 710, all fertile, inserted on margin of disc; filaments glabrous, included,
slightly united at base; ovary inserted at or near base of receptacle, pilose on exterior;
carpel unilocular, with 2 ovules, style pubescent at base, the stigma capitate. Fruit a small,
fleshy drupe, narrowed to a shortly stipitate base; epicarp smooth, not ridged, glabrous, not
lenticellate; mesocarp thin, fleshy; endocarp thin, hard, bony, breaking up in longitudinal lines
during germination, tomentose within.
Distribution. 200 species in Neotropics, 1 species in West Africa and 3 species in Thailand
and Malesia (Sumatra, Java, Borneo, the Philippines, and Papua New Guinea). In Sabah and
Sarawak only 1 species recorded.
Licania splendens (Korth.) Prance
Fig. 3.
(Latin, splendens = shining; the shiny surface of the dried leaves)

Fl. Neotropica 9 (1972) 172, l.c. (1989) 646; Prance & Whitmore l.c. 328; Anderson l.c. 293;
Cockburn l.c. 86. Basionym: Angelesia splendens Korth. l.c. 384, Merrill l.c. (1921) 290, l.c. (1929)
92, Masamune l.c. 324. Type: Korthals, s.n., Sumatra (L).
Tree to 25 m tall. Bark smooth to scaly, flaking by small scales when old; inner bark redbrown. Sapwood pink, hard. Young branches sparsely lanate, soon glabrous. Stipules
linear-lanceolate, to 3 mm long, caducous. Leaves 411 x 1.84.2 cm, oblong; cuneate at
165
166
base, usually acuminate at apex; glabrous beneath; petioles 25 mm, canaliculate, glabrous
when mature. Inflorescences terminal and axillary panicles of cymules, 1.514 cm long, the
rachis and branches grey-puberulous. Flowers c. 2 mm long; receptacle campanulate,
slightly swollen to one side, grey-tomentellous on exterior, tomentose within; pedicels c. 1
mm long; calyx-lobes acute, tomentellous on both surfaces; petals pubescent on exterior;
stamens 710 , slightly unilateral, the filaments glabrous; ovary at or near base of receptacle,
unilocular, pilose on exterior. Fruits ellipsoid, 11.3 cm long; epicarp smooth, glabrous;
mesocarp thin, fleshy; endocarp thin, hard, bony, breaking open by longitudinal lines of
weakness, tomentose within.
Vernacular names. Sabahsampaluan, tampaluan (Dusun).
Distribution. Thailand, Peninsular Malaysia, Java, Borneo (Sabah, Sarawak, Brunei,
Kalimantan), the Philippines. In Sabah common, recorded in most districts. In Sarawak,
known from Bintulu, Kuching, Lundu and Serian districts.
Ecology. Common tree of primary and secondary mixed dipterocarp forests in well-drained
usually sandy soils on slopes and also in peat swamp and on seashores and in rocky places
below 400 m.
Uses. The timber is strong, durable and resistant to marine borers and is used for saltwater
piles, railroad ties, etc. However, it is extremely hard to work and requires special tools
because of silica. The fruit is edible but is not widely used.
4. MARANTHES Blume
(Greek, maraino = to wither; the long-persistent
withered calyx and stamens below the fruit)
Bijdr. (1825) 89; Kostermans, Candollea 20 (1965) 196; Prance, Bol. Soc. Brot. Ser. 2, 40 (1966) 183,
l.c. (1989) 671; Prance & Whitmore l.c. 329; Anderson l.c. 293; Cockburn l.c. 88; Whitmore, Tantra
& Sutisna l.c. 299. Synonyms: Parinari auct. non Aubl.; Parinari sect. Sarcostegia Benth. in Hooker,
Niger Fl. (1849) 335, pro parte; Parinari subg. Sarcostegia (Benth.) Miq., Fl. Ind. Bat. 1, 1 (1855)
355, pro parte; Parinari subg. Exitelia Blume l.c. (1855) 10.
Medium-sized to large trees. Stipules deltate, intrapetiolar, stiff, caducous. Leaves glabrous
on both surfaces when mature, with dense caducous cobweb-like indumentum when young,
without stomatal crypts; with paired glands at junction of lamina and petiole; petioles
eglandular. Inflorescence a many-flowered corymbose panicle; bracts and bracteoles
eglandular, caducous, not enclosing flower buds in small group. Flowers bisexual;
receptacle obconical, narrowed into pedicel, solid, almost completely filled with nectariferous
tissue, short-tomentose to glabrous on exterior, glabrous within; calyx-lobes suborbicular,
deeply concave, unequal; petals 5, not clawed; stamens 2540, inserted on margin of disc,
unilateral with tooth-like staminodes opposite to almost in a complete circle, filaments far
exserted beyond calyx-lobes, in a tangled mass; ovary inserted laterally at mouth of
receptacle, carpel bilocular with 1 ovule in each locule, style pubescent at base only, curved
upwards, exserted. Fruit a large fleshy drupe; epicarp smooth, glabrous, not lenticellate;
mesocarp fleshy; endocarp very hard, fibrous with a rough exterior, densely tomentose
within, with 2 lateral plates which break away on germination; germination phanero-cotylar
167
(with exposed cotyledons); cotyledons fleshy, pale green; cataphylls absent; first 2 eophylls
opposite, the others alternate or opposite.
Distribution. 10 species in tropical Africa, 1 in Central America, and 1 (Maranthes corymbosa)
widespread in Malesia, Australia and the western Pacific.
Uses. Wood hard and durable, used for posts and house building. Fruits of most species
edible.
Maranthes corymbosa Blume
Fig. 4.
(Greek, corymbos = a cluster; the clustered inflorescences)

l.c. (1825) 89; Kostermans, Candollea 20 (1965) 107; Prance & Whitmore l.c. 330; Anderson l.c. 295;
Cockburn l.c. 88; Prance & White l.c. 127; Prance l.c. (1989) 673; Whitmore, Tantra & Sutisna l.c.
299. Type: Blume, s.n., Java, Prov. Krawang, near Tjiradja (L). Synonyms: Parinarium corymbosum
(Blume) Miq. l.c. (1855) 356, Merrill l.c. (1921) 290, Masamune l.c. 325; Exitelia corymbosa (Blume)
Blume, Fl. Jav. 1, Praef. (1828) 7; Parinarium griffithianum Benth. in Hooker l.c. (1849) 334.
Tree to 30 m tall, 1.5 m diameter, unbuttressed. Bark smooth, grey-brown, lenticellate,

often mottled with patches of lichen; inner bark red, thin. Sapwood pale white to pink;
heartwood red-brown. Young branches red-brown, minutely white-lenticellate, glabrous.
Stipules lanceolate, acute, 510 mm long, sparsely pilose on exterior, glabrous within, early
deciduous. Leaves coriaceous, usually oblong-lanceolate to oblong-elliptic, 6.514 x 2.58
cm; cuneate at base, acuminate at apex, the acumen 820 (30) mm long; glabrous when
mature but often sparsely caducous arachnoid-lanate when young, usually with 2
conspicuous prominent glands at junction of petiole and decurrent lower surface; lateral
veins 710 pairs, arcuate, prominulous on both surfaces; midrib plane above, prominulous
beneath; petioles 49 mm long, glabrous when mature, flattened above. Inflorescences of
flattened many-flowered corymbose panicles; rachis and branches sparsely pilose, glabrescent;
bracts and bracteoles ovate to lanceolate, sparsely pubescent, caducous. Flowers with
receptacle turbinate, tapering into pedicels, 24 mm long, grey-tomentose to glabrous on
exterior, glabrous within; calyx-lobes fleshy, ovate to elliptic, obtuse, 2.54 mm long,
unequal; petals white-tinged pink, glabrous, 36 mm long, caducous; stamens 2535,
inserted in several rows on one side of throat, with tooth-like staminodes opposite; ovary
densely lanate and villous; stigma truncate. Fruits ellipsoid, 34 x 1.52 cm, tapered
towards base; epicarp thin, glabrous on exterior when mature, sometimes lanate when
young; endocarp c. 5 mm thick, densely lanate within, bilocular usually with seed in one
locule only. Cotyledons plano-convex.
Vernacular names. Sabahbangkawang (Dusun, Malay). Sarawakmerbatu (Malay).
Distribution. S Thailand extending east to Solomon and Caroline Islands and Australia. In
Malesia: Sumatra, Peninsular Malaysia, Java, Borneo (Sabah, Sarawak, Brunei, Kalimantan),
the Philippines, Celebes, Moluccas, New Guinea, New Britain and Admiralty Islands. In
Sabah, widespread but in Sarawak, uncommon in mixed dipterocarp forest in the northern
parts.
168
Ecology. In primary and secondary forests from the sea-coast to 1500 m on Mt. Kinabalu,
Sabah. The fruit is eaten by many birds which probably disperse the seed and account for
the wide distribution of this species.
Uses. The wood is used for house building and for posts and the fruit is edible.
5. PARASTEMON A. DC.
(Greek, para = near, stemon = stamen; both fertile and
sterile stamens are located near one another at one side of the flower)
Ann. Sci. Nat. Bot. Ser. 2, 18 (1842) 28; Miquel l.c (1855) 359; Merrill l.c. (1921) 290; Ridley l.c.
(1922) 672; Corner l.c. 617; Masamune l.c. 324; Anderson l.c. 293; Cockburn l.c. 90; Prance l.c.
(1989) 648; Whitmore, Tantra & Sutisna l.c. 299. Synonyms: Diemenia Korth. l.c. (1854) 388;
Trichocarya Miq. l.c. (1855) 537, pro parte.
Trees or shrubs. Stipules small and triangular, caducous. Leaves glabrous on both surfaces,
without stomatal crypts, with 2 small discoid glands at base of lamina; petioles eglandular.
Inflorescences axillary or rarely terminal simple or sparsely branched racemes; bracts and
bracteoles small, eglandular, not enclosing groups of flower buds. Flowers bisexual or
unisexual; receptacle patelliform or shallowly cupuliform, shortly hairy within; calyx-lobes
5, acute, subequal; petals 5, not exceeding calyx-lobes, not clawed; stamens either 5 and all
fertile or 2 fertile with 3 staminodes, the filaments glabrous, shorter than the calyx-lobes;
ovary centrally inserted at base of receptacle, glabrous or densely hairy on exterior, carpel
unilocular, with 2 ovules, style filiform, puberulous towards the base, with 3 large
undivided lobes at apex or 1 obscure lobe and 2 large, sometimes deeply divided lobes.
Fruit a drupe, 1.53 cm long, with 2 large lateral plates which break away on germination
to allow seedling escape; epicarp smooth, not lenticellate; endocarp thin, hard, bony,
smooth on exterior, glabrous within.
Distribution. 3 species in the Nicobar Islands, Sumatra, Peninsular Malaysia, Borneo,
Moluccas, New Guinea, and Admiralty Islands. 2 species in Sabah and Sarawak.
Ecology. Lowland and swamp forests.
Key to Parastemon species

Leaves with 56 pairs of lateral veins. Fruit 2.33.5 cm long.................1. P. grandifructus
Leaves with 811 pairs of lateral veins. Fruit 11.5 cm long.......................2. P. urophyllus
169
170
1. Parastemon grandifructus Prance

(Latin, grandis = large, fructus = fruit)
Brittonia 39 (1987) 366, l.c. (1989) 650. Type: Wright & Ismawi S. 32320, Sarawak (L).
Tree to 30 m tall, 30 cm diameter; trunk lightly buttressed to 1 m high. Young branches

glabrous. Stipules caducous. Leaves coriaceous, glabrous on both surfaces, elliptic to
narrowly ellipticoblong, 58.5 x 1.83.2 cm; cuneate at base, with a long-cuspidate
acumen at apex, the tip 1016 mm long; midrib prominent above, prominulous or plane
beneath; lateral veins 56 pairs, prominulous above, plane beneath; petioles 58 mm long,
glabrous, slightly canaliculate, slightly swollen at base. Inflorescences of axillary and
terminal racemes, the rachis glabrous. Flowers (seen in fruiting specimens): calyx-lobes 5,
acute, glabrous on exterior, glabrous within except for a few hairs around base; receptacle
glabrous on exterior in fruiting condition; style persistent below fruits, the stigma bifid or
trifid. Fruit ellipsoid, 2.33.5 x 1.31.5 cm; epicarp smooth, glabrous; mesocarp thin, c.
0.25 mm; endocarp thin, hard, bony, c. 0.25 mm thick, glabrous within, opening by 2 lateral
plates 1.92 cm long.
Vernacular names. Sabahkayu ajung, mandailas (Dusun). Sarawakngilas (Iban, in
common with Xanthophyllum).
Distribution. Endemic to Borneo (Sabah and Sarawak). In Sabah known from Beaufort Hill
and Lumat Estate in the Beaufort district. In Sarawak found in Bau, Kapit, Kuching and
Limbang districts.
Ecology. Lowland forests, including swamp and heath forests, to 150 m.
2. Parastemon urophyllus (Wall. ex A. DC.) A. DC.
Fig. 5.
(Greek, aura = tail, phyllon = leaf; the cuspidate apex of the leaf)
l.c. 208; Miquel l.c. (1855) 359; Merrill l.c. (1921) 290; Ridley l.c. (1922) 672; Corner l.c. 617;
Masamune l.c. 324; Browne, FTSB (1955) 308; Anderson l.c. 293; Cockburn l.c. 90; Prance &
Whitmore l.c. 331; Prance l.c. (1989) 649; Whitmore, Tantra & Sutisna l.c. 300. Basionym: Embelia
urophylla Wall. ex A. DC., Trans. Linn. Soc. Lond. 17 (1837) 131. Type: Griffith, s.n., Malacca (K).
Synonym: Parastemon spicatum Ridl., J. Str. Br. R. As. Soc. 75 (1917) 29, Merrill l.c. (1921) 290,
Masamune l.c. 324.
Tree to 35 m tall often with small buttresses. Bark brownish, smooth, becoming slightly
cracked and fissured with age. Young branches glabrous, waxy resinous when young.
Stipules triangular, c. 1 mm long, caducous. Leaves thinly coriaceous, narrowly oblong,
2.58 x 1.42.5 cm; cuneate at base, cuspidate-acuminate at apex, the tip 515 mm; midrib
plane above, prominulous beneath; lateral veins 811 pairs; petioles 45 mm long,
canaliculate, glabrous. Inflorescences of axillary and rarely terminal racemes or occasionally
slightly branched, 414 cm long, the rachis glabrous. Flowers unisexual (plant polygamodioecious), c. 1.5 mm long; receptacle broadly cupuliform to flattened saucer-shaped,
glabrous on exterior, tomentose within; pedicels to 2 mm long; calyx-lobes acute, glabrous
on exterior; petals 5; stamens 2 fertile and 3 sterile staminodes opposite; ovary inserted at
base of receptacle, pilose on exterior, unilocular, style pilose at base, glabrous above, the
stigma trifid. Fruits ellipsoid, 11.5 cm long; epicarp smooth, glabrous; mesocarp thin,
hard; endocarp thin, hard, bony, glabrous within, opening by 2 lateral plates.
171
Vernacular names. Sabahmendailas (Dusun). Sarawakmengilas, gilas (Bidayuh).
Distribution. Nicobar Islands, Sumatra, Peninsular Malaysia, Borneo (Sabah, Sarawak,

Brunei, Kalimantan). In Sabah found in Beaufort, Keningau, Papar and Sipitang districts in
the west, and in Lahad Datu and Sandakan districts in the east. In Sarawak widespread and
recorded in most districts.
Ecology. Characteristic component of peat swamp forest where it is a common large tree,
but also wide-ranging into kerangas, more open scrub forest, and secondary forest on poor
soils.
Uses. The wood is hard and very heavy; grain straight or interlocked. It is used for general
construction, posts and as a firewood.
6. PARINARI Aubl.
(a vernacular name in French Guiana)
merbatu (Malay)
l.c. 204; Merrill l.c. (1921) 290, l.c. (1929) 92; Masamune l.c. 325; Backer & Bakhuizen f. l.c. 521;
Kostermans, Reinwardtia 7, 1 (1965) 147; Prance & Whitmore l.c. 332; Anderson l.c. 294; Cockburn
l.c. 90; Prance l.c. (1989) 654; Whitmore, Tantra & Sutisna l.c. 300. Synonym: Parinarium Juss.,
Gen. Pl. (1789) 342; Petrocarya Schreb. in Linnaeus, Gen. Pl. ed. 8, 1 (1789) 245; Parinarium subg.
Petrocarya (Schreb.) Miq. l.c. (1855) 352.
Small or large trees or rarely shrubs. Stipules small to large, persistent or caducous. Leaves
usually with stomatal crypts, filled with pubescence on lower surface or rarely glabrous, or
lanate pubescent without crypts; petioles usually with 2 circular glands above. Inflorescence a
many-flowered complex cyme or cymose panicle; bracts and bracteoles eglandular, usually
concealing flower buds individually and in small groups. Flowers bisexual; receptacle
subcampulate to cupuliform, slightly swollen to one side, tomentose on both surfaces; calyxlobes 5, deltate, acute, densely hairy on both surfaces; petals 5, as long as or shorter than
sepals, caducous; stamens 610, unilateral, the filaments glabrous, included, with c. 6
minute staminodes opposite; ovary inserted on upper half of receptacle-tube below mouth,
pilose on exterior, carpel bilocular with 1 ovule in each locule, style arcuate, included. Fruit
a fleshy drupe; epicarp verrucose; endocarp thick, with a rough fibrous surface, with 2 basal
obturators for seedling escape.
Distribution. Pantropical with 18 species in the Neotropics, 6 in Africa and 15 in tropical
Asia, Malesia and the Pacific. 7 species in Sabah and Sarawak.
Ecology. A wide range of lowland forest and one species in submontane forest.
Uses. The wood is hard and heavy and is little used, the fruits of most species are edible.
172
Key to Parinari species

1. Stomatal crypts absent from leaf underside; leaf underside glabrous or with a persistent
lanate pubescence and then with large persistent stipules 720 mm.............................2
Stomatal crypts present on leaf underside; leaf underside lanate or at least pubescent in
crypts; stipules usually small or if larger then early caducous.......................................4
2.
Leaf underside glabrous; stipules small and caducous..................................................3

Leaf underside densely lanate pubescent, but when removed no stomatal crypts present;
stipules large and persistent, 718 mm long................................................4. P. elmeri
3.
Leaves elliptic to oblong or obovate-elliptic, 9.520.5 x 4.58.5 cm; lateral veins 1116
pairs. Panicles large and silvery pubescent.............................1. P. argenteo-sericea
Leaves ovate, 59 x 245 cm; lateral veins 711 pairs. Panicles small and subsericeous
brown pubescent.................................................................................2. P. canarioides
4.
Leaves with 2033 pairs of lateral veins.............................................6. P. oblongifolia

Leaves with 1019 pairs of lateral veins.......................................................................5
5.
Petioles 1420 mm long. Leaves with metallic sheen above....................5. P. metallica

Petioles 310 mm long. Leaves without a metallic sheen.............................................6
6.
Leaves rigidly coriaceous, often broadest well below middle point; midrib and often
lateral veins lightly impressed on upper surface. Receptacle c. 5 mm. Fruits ovoid, 78
cm long.......................................................................................................7. P. rigida
Leaves chartaceous, broadest at or above middle; midrib and lateral veins prominulous
above. Receptacle 33.5 mm. Fruits ellipsoid, 3.54.5 cm long..................3. P. costata
1. Parinari argenteo-sericea Kosterm.

(Latin, argenteus = silvery, sericeus = silky; the pubescence of the flowers and inflorescence)
Reinwardtia 7, 1 (1965) 47, 158; Cockburn l.c. 91; Prance l.c. (1989) 656; Whitmore, Tantra &
Sutisna l.c. 300. Type: Wood SAN 16175, British North Borneo, Lahad Datu (holotype BO; isotypes
A, BRI, K, KEP, L, SING).
Tree to 35 m tall. Bark brown, lenticellate, hard; inner bark red, hard, c. 1.2 mm thick.
Sapwood yellow. Young branches glabrous, dark purplish brown, with numerous lenticels.
Stipules lanceolate, to 8 mm long, tomentose on exterior, early caducous. Leaves chartaceous,
oblong, elliptic to subovate-elliptic, 9.510.5 x 4.58.5 cm, glabrous on both surfaces,
without stomatal crypts beneath, usually 2 glands beneath at base near junction with
midrib; rounded at base, acute to shortly acuminate at apex, the tip 710 mm long; midrib
lightly impressed above except near base, prominent beneath; lateral veins 1116 pairs,
plane above, prominent beneath, erect-patent; petioles 59 mm long, eglandular, glabrous,
rugulose. Inflorescence a lax, much-branched, terminal panicle, 915 cm long, the rachis
and branches densely grey sericeous-tomentose; bracts and bracteoles ovate, acute, densely
tomentellous on exterior, glabrous within except near apex, caducous. Flowers with
receptacle campanulate, markedly gibbous, densely grey-tomentellous on exterior, 23 mm
long, narrowly ovate, densely grey-tomentose on exterior, tomentellous within; petals
spathulate, c. 2 mm long, caducous; fertile stamens 78, base forming a conspicuous fused
ring with opposite tooth-like staminodes; ovary densely pilose, style pilose, stigma truncate.
Fruits ovoid, 78 x 4.55.5 cm; epicarp densely lenticellate; mesocarp thin, fleshy;
173
endocarp extremely hard and thick (18 cm thick), woody, granular, and very irregularly
ridged, with 2 small locules in centre, densely lanate within.
Distribution. Endemic to Sabah, known from a few collections from the Lahad Datu,
Sandakan, and Tawau districts.
Ecology. Lowland forest to 100 m and forest along rivers.
2. Parinari canarioides Kosterm.

(resembling the genus Canarium)
New & Crit. Mal. Pl. (For. Dept. Bur. of Planning Bogor, Indonesia) 3 (1955) 25; Reinwardtia 7, 2
(1965) 159; Anderson l.c. 294; Cockburn l.c. 93; Prance l.c. (1989) 656; Whitmore, Tantra & Sutisna
l.c. 300. Type: Kostermans 7152, Kalimantan, Tanjong Bangko (holotype BO; isotypes A, BISH,
BRI, CAL, CANB, K, L, LAE, MEL, NY, P, PNH, SAN, SING).
Tree to 60 m tall, trunk often buttressed to 1.52.5 m. Bark smooth, irregularly fissured and
flaking, dark red-brown; inner bark pale red-brown. Sapwood dull yellow to pink;
heartwood brownish red. Young branches sparsely puberulous soon becoming glabrous,
grey-brown. Stipules linear, acute to 5 mm, hirsute, early caducous, present on very young
leaves only. Leaves chartaceous, ovate, 59 x 24.3 cm, glabrous on both surfaces when
mature, without stomatal crypts beneath; rounded to subcordate at base, acuminate at apex,
the tip 512 mm long; midrib lightly impressed above, prominent beneath, sparsely
pubescent when young; lateral veins 711 pairs, plane to prominulous above, prominent
beneath, arcuate; petioles 37 mm, glabrous when mature, eglandular or with small rather
inconspicuous central glands. Inflorescences dense-flowered axillary panicles to 4.5 cm
long, the rachis and branches tomentose; bracts and bracteoles persistent, ovate, puberulous
on exterior, caducous. Flowers with receptacle campanulate, c. 3 mm long, tomentose on
exterior; pedicels 12 mm long; calyx-lobes elliptic, concave, c. 2 mm, acute, sparsely
puberulous on exterior, densely tomentellous on interior; petals elliptic, obtuse, c. 2 mm,
tapered to the base; fertile stamens 78. Fruits ellipsoid, 3.55 x 1.52.5 cm; epicarp
densely to sparsely lenticellate; mesocarp fleshy, c. 1 mm thick; endocarp c. 5 mm thick,
hard, marbled, densely lanate within.
Distribution. Sumatra, Borneo (Sabah, Sarawak, Brunei, Kalimantan), the Philippines
(Palawan), and Celebes. In Sabah found in Kunak and Sandakan districts; widespread but
scattered in Sarawak and Brunei.
Ecology. Primary and secondary mixed dipterocarp forests on well-drained clay-rich soils to
800 m.
Uses. Fruit edible.
174
3. Parinari costata (Kunth) Blume

(Latin; costatus = ribbed; the strongly veined leaf)
l.c. (1855) 10; Miquel l.c. (1855) 354; Merrill l.c. (1921) 290; Ridley l.c. (1922) 666; Masamune l.c.
325; Kostermans, Reinwardtia 7, 2 (1965) 179; Prance & Whitmore l.c. 333; Anderson l.c. 294;
Prance l.c. (1989) 663; Whitmore, Tantra & Sutisna l.c. 300. Basionym: Lepidocarpa costata Korth.
l.c. 387. Type: Korthals, s.n., Sumatra (L).
Tree to 60 m tall, 90 cm diameter; trunk buttressed up to 2 m. Bark smooth to roughish,

cracked, grey or brown, c. 0.5 mm; inner bark pale reddish to reddish brown with pale spots,
610 mm thick. Wood pale brown, darker towards centre. Young branches densely yellowbrown, appressed tomentellous becoming glabrous, with numerous small conspicuous lenticels.
Stipules lanceolate, membranous, 37 mm long, pilose on exterior, early caducous. Leaves
rigidly chartaceous, elliptic, subovate-elliptic, oblong-elliptic to oblong-lanceolate, 411.5 x
1.64.3 cm, glabrous above when mature but with sparse lanate covering when very young,
with stomatal crypts filled with grey lanate pubescence beneath; rounded to subcuneate at
base, acuminate at apex, the tip 35 mm long; midrib prominulous above, tomentellous
towards base, prominent beneath; lateral veins 1019 pairs, arcuate, prominulous above,
prominent beneath; intercostal veins rounded or only slightly flattened; petioles 49 mm
long, slender, thickly tomentose or tomentellous when young, soon glabrous, usually
eglandular or with 2 inconspicuous median glands. Inflorescences of predominantly
axillary or terminal few-flowered lax or dense panicles to 8 cm long, the rachis and
branches grey to brown appressed tomentellous or ferrugineous-villous pubescent; bracts
and bracteoles lanceolate, c. 2 mm long, caducous. Flowers with receptacle campanulate,
slightly gibbous, grey-brown pubescent on exterior, 33.5 mm long; pedicels 0.51 mm
long; calyx-lobes ovate, acute, 1.52 mm long, grey tomentellous on exterior; petals white,
spathulate, 1.52 mm long, caducous, glabrous; stamens 78, with small tooth-like
staminodes opposite, slightly unequal; style glabrous, stigma capitate. Fruits ellipsoid, to
3.5 x 4.5 cm; epicarp usually sparsely verrucose or lenticellate; mesocarp c. 2 mm, fleshy;
endocarp hard, marbled, 35 mm thick, fibrous, densely lanate within.
Key to subspecies
Inflorescences and flowers sparsely to densely grey or brown appressed pubescent. Lowland
plants...................................................................................................................................
subsp. costata
Sumatra, Peninsular Malaysia, Borneo (Sabah, Sarawak, Brunei, Kalimantan). Mixed
dipterocarp forests on well-drained soils, to 300 m. In Sabah and Sarawak uncommon.
Inflorescences and flowers densely ferrugineous-villous pubescent. Usually at high altitudes
subsp. rubiginosa (Ridl.) Prance
Brittonia 39 (1987) 368, l.c. (1989) 663. Basionym: Parinarium costatum var. rubiginosum Ridl.
l.c. (1915) 143; Parinarium rubiginosum (Ridl.) Ridl. l.c. (1917) 29, l.c. (1922) 668. Type: Ridley
16016, Malaya, Padang (holotype K; isotype SING). Synonym: Parinari bicolor Merr., Philip. J.
Sc. 10 (1915) Bot. 309.
Burma, Peninsular Malaysia, Borneo (Sabah, Sarawak, Kalimantan). Submontane

forests at 7501000 m. Uncommon; in Sabah known from Mt. Kinabalu (Clemens
175
176
50081), Sandakan district (Puasa 669), and in Sarawak from Similajau and Segan FR,
Bintulu, 4th Div.
Distribution. Burma, Sumatra, Peninsular Malaysia, Borneo (Sabah, Sarawak and
Kalimantan), and the Philippines.
Ecology. Mixed dipterocarp to submontane forests on well-drained soils at 3001000 m.
Taxonomy. Three subspecies are recognised, two of which occur in Sabah and Sarawak (as
above), and a third in the Philippines.
4. Parinari elmeri Merr.

(A.D.E. Elmer, plant collector with the Bureau of Science, Manila, the Philippines)
l.c. (1929) 92; Masamune l.c. 325; Kostermans, Reinwardtia 7, 2 (1965) 161; Prance & Whitmore l.c.
335; Anderson l.c. 294; Cockburn l.c. 91; Prance l.c. (1989) 657; Whitmore, Tantra & Sutisna l.c. 300.
Type: Elmer 20806, British North Borneo, Tawao, Elphinstone Province (holotype UC; isotypes
BISH, BO, BR, C, DC, DS, F, GH, L, M, MO, P, S).
Tree to 30 m usually much smaller; trunk unbuttressed. Bark pale cream or grey and white
mottled, roughened by green excrescences, soft, thin; inner bark orange, hard, c. 2.5 mm
thick. Sapwood white, thin; heartwood straw coloured, hard. Young branches densely
tomentellous soon becoming glabrous, obscurely lenticellate. Stipules lanceolate, acute, to
18 mm long and 3 mm broad at base, lateral, tomentellous, persistent. Leaves oblong to
oblong-lanceolate, 518 x 1.57 cm, chartaceous to thinly coriaceous, glabrous above,
densely pubescent beneath, without stomatal crypts; subcuneate at base, acuminate at apex,
the tip 513 mm long; midrib plane or slightly impressed and pubescent above when young,
prominent beneath; lateral veins 1421 pairs, prominent beneath, curved at margin;
intercostal veins more or less parallel forming ladder-like reticulation; petioles 1.56 mm
long, tomentellous, glandular, but glands often obscured. Inflorescences of raceme-like
reduced terminal and axillary panicles or cymules, 1.73 cm long, the rachis and branches
densely brown-tomentose; bracts and bracteoles large, c. 2 mm long, ovate, persistent.
Flowers with receptacle conical, gibbous, to 3 mm long, brown-lanate on exterior, pedicels
0.52 mm long; calyx-lobes ovate, acute, 23 mm long, lanate on exterior; petals white,
oblong-ovate, 23 mm long, narrowed to the base; fertile stamens 79, with tooth-like
staminodes opposite. Fruits oblong-ellipsoid, c. 6.7 x 3.7 cm; epicarp sparingly lenticellate.
Distribution. Peninsular Malaysia, Borneo (Sabah, Sarawak, Brunei, Kalimantan), and the
Philippines. Uncommon; in Sabah known from the type collection and from Kelumpang
Hill, Lahad Datu district (SAN 29329), and Tawau district. In Sarawak only two collections
from G. Silantek, 85th mile, Simanggang Rd., 2nd Div. (S. 42636) and Mt. Matang
(Clemens s.n.) are known, but has been recorded from ecological plots at Bt. Mersing,
Anap, and Nyabau FR.
Ecology. Mixed dipterocarp forests on well-drained soils to 900 m, including areas on
ultramafic rock.
Uses. The wood is used for supports of Iban long houses.
177
5. Parinari metallica Kosterm.

(Latin, metallicus = metal-like; the metallic sheen on the upper surface of dried leaves)
Reinwardtia 7, 1 (1965) 49, 160; Cockburn l.c. 91; Prance l.c. (1989) 660. Type: Ashton BRUN 3267,
Brunei, Andulau FR (holotype SAR; isotypes BO, K, L).
Tree to 20 m tall, unbuttressed. Young branches appressed strigose soon becoming

glabrous, conspicuously lenticellate. Stipules ovate-lanceolate, acute, 815 mm long, densely
brown-tomentose, membranous, early caducous. Leaves thickly coriaceous, elliptic, 817 x
49 cm, glabrous and shiny with metallic sheen above when dry, with dense stomatal crypts
filled with hairs; rounded or subcuneate at base, apex rounded to shortly blunt-acuminate,
the tip to 3 mm long; midrib plane above, prominent beneath; lateral veins 1015 pairs,
prominulous to plane above, prominent beneath, erect, curved only at margin; petioles 14
20 mm long, glabrescent, with inconspicuous glands near to lamina base, puberulous,
glabrescent. Inflorescences of axillary little-branched panicles, 410 cm long, the rachis
and branches densely brown tomentellous; bracts and bracteoles ovate, early caducous.
Flowers with receptacle campanulate, slightly gibbous, 23 mm long, ferrugineous,
pubescent on exterior; pedicels c. 0.5 mm long; calyx-lobes lanceolate, acute, c. 1 mm long,
tomentellous; petals lanceolate, glabrous; stamens c. 8 with short tooth-like staminodes
opposite; ovary densely pilose, style glabrous, equalling stamens, stigma truncate. Fruits not
seen.
Distribution. Endemic to Borneo (Sabah, Sarawak, Brunei). In Sabah found in Beaufort
Hill FR, and in Sarawak in Semengoh Arboretum, Kuching, 1st Div., and Lambir Hills, Miri
district, 4th Div.
Ecology. Mixed dipterocarp forests on deep sandy humult ultisol soils, to 300 m.
6. Parinari oblongifolia Hook. f.
Fig. 6.
(Latin, oblongus = rather long, folius = leaves; the leaf-shape)

l.c. 309; Ridley l.c. (1922) 335; Anderson l.c. 294; Cockburn l.c. 93; Prance l.c. (1989) 659;
Whitmore, Tantra & Sutisna l.c. 301. Type: Griffith, s.n., Malacca (K). Synonyms: Parinarium
borneense Merr. l.c. (1929) 93, Masamune l.c. 325; Parinari gigantea Kosterm., Reinwardtia 7, 2
(1965) 182; Prance l.c. (1989) 660; syn. nov.
Tree to 40 m tall, trunk with low thick buttresses to 2 m. Bark smooth, grey to pale brown,
very thin; inner bark reddish brown to brown, 612 mm thick. Sapwood white to pale
yellow; heartwood reddish, hard. Young branches minutely tomentellous, with numerous
pale, small, roundish lenticels. Stipules ovate to lanceolate, acute, 35 mm, pilose on
exterior, early caducous. Leaves coriaceous, elliptic to oblong, 1423 x 49 cm, glabrous
above, with stomatal crypts, filled with grey lanate pubescence beneath; rounded to
subcordate at base, shortly acuminate at apex, the tip 313 mm long; midrib plane above,
glabrous when mature except at base, prominent, glabrescent beneath; lateral veins 2335
pairs, erect, plane above, flattened and prominent beneath; intercostal veins prominulous,
parallel and more or less ladder-like beneath; petioles 917 mm long, thick-tomentellous
when young, glabrescent, eglandular or glandular. Inflorescences of large, spreading
terminal panicles, 1021 cm long, 712 cm broad, the rachis and branches yellow-grey
tomentellous; bracts and bracteoles ovate, c. 3 mm long, early caducous. Flowers with
178
receptacle campanulate, slightly gibbous, c. 3 mm long, densely grey-tomentose on exterior;

pedicels 13 mm long; calyx-lobes ovate, acute, 1.52 mm long, unequal, grey-tomentose;
petals white to bluish, lanceolate to spathulate; narrowed towards the base, c. 2 mm long,
glabrous; stamens 710, with tooth-like staminodes opposite; ovary pilose, style glabrous,
stigma truncate. Fruits ellipsoid, 59 x 34 cm; epicarp densely lenticellate; mesocarp 1.52
mm thick; endocarp hard, marbled, 713 mm thick, fibrous, densely lanate within.
Distribution. Sumatra, Peninsular Malaysia, Borneo (Sabah, Sarawak, Brunei, Kalimantan). In
Sabah found in Lahad Datu, Sandakan, Semporna, and Tawau districts; in Sarawak only
recorded from sterile ecological collections from Santubong to Miri.
Ecology. Lowland rainforest, on river banks or in valleys, to 450 m.
7. Parinari rigida Kosterm.

(Latin, rigidus = stiff; the thick coriaceous leaves)
Reinwardtia 7, 1 (1965) 53, 163; Prance l.c. 660. Type: Sinclair & Kiah SFN 40773, Malay Peninsula,
Trengganu (SING). Synonym: Parinari ashtonii Kosterm., Reinwardtia 7, 1 (1965) 53, Anderson l.c.
294.
Tree to 30 m tall. Bark smooth. Young branches tomentellous soon becoming glabrous,
obscurely lenticellate. Stipules caducous (not seen). Leaves rigidly coriaceous, elliptic to
oblong-ovate, 7.523 x 38 cm, those near to inflorescence much smaller than others,
broadest below mid point, glabrous and shiny above, sometimes slightly bullate, the lower
surface with stomatal crypts filled with pubescence, with 2 glandular areas at junction of
midrib and petiole below; rounded or subcordate at base, shortly and broadly acuminate at
apex, the tip 317 mm long; midrib plane or impressed for upper portion above, prominent
and appressed pilose beneath when young; lateral veins 1320 pairs, slightly impressed
above, prominent beneath, slightly curved at margins only; intercostal venation flattened or
rounded, parallel; petioles thick, 310 mm long, grey-pilose pubescent, rugose, with 2 small
glands on mid-point of upper side. Inflorescences of narrow terminal panicles to 13 cm
long, the rachis and branches tomentose; bracts and bracteoles lanceolate, to 2 mm long,
early caducous. Flowers with receptacle campanulate, slightly gibbous, c. 5 mm long,
densely villous-tomentose on exterior; pedicels c. 1 mm long; calyx-lobes elongate
triangular, 22.5 mm long; petals spathulate; stamens 68; ovary densely villous, style
equalling stamens, stigma capitate. Fruits irregularly ellipsoid, c. 5 x 4 cm, tapered towards
the base almost into a stipe; epicarp densely lenticellate; mesocarp thin, fleshy; endocarp
thick, woody, marbled, lanate within.
Distribution. Sumatra, Peninsular Malaysia, Borneo (known only from Lundu, Kuching
and Miri districts in Sarawak).
Ecology. Heath and swamp forests, lowlands to 1400 m.
179
CLETHRACEAE
A. Berhaman
Sandakan, Malaysia
Merrill, EB (1921) 460; Masamune, EPB (1942) 567; Sleumer, FM 1, 7 (1971) 139; Whitmore, TFM
2 (1973) 27; Anderson, CLTS (1980) 162; Whitmore, Tantra & Sutisna, CLK 1 (1989) 45.
A small monogeneric family in the Ericales of about 65 species distributed in the temperate,
tropical American and Asiatic-Malesian regions.
CLETHRA Gronov. ex L.
(Latinised old Greek word klethra = alder tree)
kolintuhan (Dusun Ranau, Sabah)
Gen. Pl. ed. 5 (1753) 188, Sp. Pl. (1753) 396; Merrill l.c. (1921) 460; Masamune l.c. 567; Sleumer,
Bot. Jahrb. 87 (1967) 85, l.c. (1971) 139; Whitmore l.c. 27; Anderson l.c. 162; Whitmore, Tantra &
Sutisna l.c. 45.
Trees or shrubs. Leaves simple, spiral, crowded towards the end of the shoots; margin
serrate with glandular teeth, more rarely entire; lateral veins and midrib sunken on upper
side, prominent on the lower side; stipules none. Inflorescence a solitary terminal raceme,
or (usually) a terminal raceme and several lower approximate racemes, from the axils of
reduced or caducous leaves; bracts mostly caducous during anthesis, rarely subpersistent.
Flowers bisexual; calyx-lobes 5(6), persistent, alternate with the petals; petals 5(6),
generally free; stamens 10(12) in 2 whorls of 5(6), filaments adnate to the corolla at base,
anthers dorsifixed, opening by apical valves; ovary superior, 3-celled, with axile
placentation; ovules many, small, anatropous; style simple, projecting out of the flower,
sometimes divided into three apical lobes, each lobe stigmatic at the top. Fruit a capsule, 3loculed, enclosed by the persistent calyx. Seeds many, small; endosperm fleshy; embryo
cylindrical.
Taxonomy. A monograph of the genus has been published by Sleumer l.c. (1967). There
are 10 species recorded for South East Asia. Only 3 species occur in Sabah and Sarawak;
they are closely related species differing mainly in the details of their hair and flower size.
Key to species
1.
Leaf lower surface with stellate hairs only on the midrib and veins, glabrous on the
intervenium. Calyx-lobes 1.52 mm long..............................................1. C. canescens
181
182
CLETHRACEAE (BERHAMAN)
Leaf lower surface with stellate hairs on the midrib and veins, and a fine pale tomentum
covering the entire surface of the intervenium. Calyx-lobes 2 mm or longer.................2
2.
Raceme slender (11.5 mm thick), axis covered by a mixture of large and tiny stellate
hairs (the hairs shorter than 0.5 mm). Calyx-lobes 23 mm long.......2. C. longispicata
Raceme more robust (1.52 mm thick), axis covered by a mixture of tufts of long hairs
(typically 0.51 mm long) and tiny stellate hairs. Calyx-lobes 34 mm long..................
.........................................................................................................3. C. pachyphylla
1. Clethra canescens Reinw. ex Blume

(Latin, canescens = turning grey; the pubescent leaves)
Bijdr. (1826) 863; Sleumer l.c. (1967) 85, l.c. (1971) 145; Anderson l.c. 162; Whitmore, Tantra &
Sutisna l.c. 46. Type: Reinwardt, s.n., 1821, Celebes, Minahasa, Mt. Klabat (L).
var. clementis (Merr.) Sleumer

l.c. (1967) 86, l.c. (1971) 145. Basionym: Clethra clementis Merr., Philip. J. Sc. Bot. 13 (1918) 104,
l.c. (1921) 460; Masamune l.c. 567; Anderson l.c. 162. Type: Clemens 11148, British North Borneo,
Mt. Kinabalu (PNH, destroyed). Synonym: Clethra canescens (non Reinw. ex Blume) Stapf, Trans.
Linn. Soc. Bot. 42 (1914) 105.
Small tree to 10 m. Bark smooth, pale brown; inner bark pale yellow. Sapwood yellowish.
Leaves chartaceous to thinly coriaceous, upper surface smooth and shiny, lower surface
with stellate hairs only on the midrib and veins, glabrous on the intervenium, oblong to
elliptic-obovate, 313(15) x 13(5.5) cm; base cuneate, apex acute to acuminate pointed;
lateral veins 1014 pairs, looping towards the margin; midrib sunken on upper side,
glabrous to sparsely covered with stellate hairs and rusty tomentum on lower side; stalk 1
1.5(2.2) cm long, glabrous to sparsely stellate-hairy. Inflorescence axis slender, 12 mm
thick, covered with a fine rusty tomentum of both smaller and larger stellate or fascicled
hairs, flowers densely arranged along the rachis; bracts subulate, hairy, c. 5 mm long.
Flowers white, scented; calyx-lobes 1.52 mm, covered with stellate and fascicled hairs;
petals white, spathulate, glabrous, fimbriate, 45 mm long; filaments glabrous, 33.5 mm
long; anthers obovate, 0.50.8 mm long; ovary densely covered with straight hairs; style
glabrous, 2.53.7 mm long. Fruit subglobose, 23 mm diameter. Seed irregularly ovoidangular, 0.61 mm across.
Distribution. Endemic to Borneo (Sabah, Sarawak and Kalimantan). In Sabah, var. clementis
is locally common on Mt. Kinabalu; in Sarawak it has been recorded from Mt. Dulit
(Richards 1772 & 1804) and the Baram River (Haviland 1828).
Ecology. In montane forest, rarely in lowland forest.
Taxanomy. Sleumer (1967 & 1971) recognises 5 varieties, viz. var. canescens, clementis,
ledermannii, luzonica, and novoguinensis, of which only var. clementis occurs in Sabah and
Sarawak.
2. Clethra longispicata J.J. Sm.

(Latin, longus = long, spicatus = spike; the inflorescence)
183
Bull. Jard. Bot. Btzg. 3, 4 (1922) 240; Sleumer l.c. (1967) 96, l.c. (1971) 148; Anderson l.c. 162;
Whitmore, Tantra & Sutisna l.c. 46. Type: Rachmat 553, C Celebes, Mt. Nanakan (BO, L). Synonym:
Clethra elongata J. J. Sm., Bull. Jard. Bot. Btzg. 3, 1 (1920) 398, non Rusby (1907).
Small tree to 10 m tall. Bark brown, shallowy fissured; inner bark yellow. Sapwood white.
Leaves chartaceous, upper surface smooth and shiny, lower surface with stellate hairs on
the midrib and veins, and a fine pale tomentum covering the entire surface of the intervenium,
oblong-lanceolate to elliptic-oblong, 713.5(15.5) x 24.5(6) cm; base cuneate, rarely
almost rounded, apex shortly acuminate; midrib sunken on upper side, covered with stellate
hairs on lower side; lateral veins 1214 pairs, looping towards the margin; stalk 1.53 cm
long, sparsely to densely covered with long simple and stellate hairs. Inflorescence axis
slender, 11.5 mm thick, laxly many-flowered, axis covered by a mixture of large and tiny
stellate hairs; bract subulate, caducous, 46 mm long. Flowers white, scented; calyx-lobes
narrowly ovate-subdeltoid, 23 mm long; petals white, broadly spathulate, glabrous, 23.5 x
11.5 mm; filaments white, glabrous, 11.5 mm long, anthers obcordate, c. 0.5 mm long;
ovary appressed hairy; style glabrous, 1.51.7 mm long. Fruit subglobose, 22.5 mm
diameter. Seed subglobose, c. 0.7 mm across.
Distribution. Borneo (Sabah, Sarawak and Kalimantan), Philippines, and C Celebes. Common
on the highlands in Sabah and Sarawak.
Ecology. In hill and submontane forests, also in secondary forest, at 5001500 m.
3. Clethra pachyphylla Merr.
Fig. 1.
(Greek, pakus = thick, phyllon = leaves)

l.c. (1918) 103, l.c. (1921) 460; Masamune l.c. 567; Sleumer l.c. (1967) 99, l.c. (1971) 150; Anderson
l.c. 162; Whitmore, Tantra & Sutisna l.c. 46. Type: Clemens 10692, British North Borneo, Mt.
Kinabalu, Paka Caves (PNH, UC).
Small tree to 10 m. Bark shallowly fissured, pale grey; inner bark yellowish. Sapwood
yellowish. Leaves coriaceous, upper surface smooth and shiny, lower surface with stellate
hairs on the midrib and veins, and a fine pale tomentum covering the entire surface of the
intervenium, oblong or obovate-oblong, 412(19) x (2) 35 cm; base cuneate, apex acute;
midrib sunken on upper side, densely covered with stellate and fascicled hairs on lower side;
lateral veins 912(14) pairs, looping towards the margin, covered with fascicled hairs; stalk
1.52.5 cm long, densely covered with long simple- and stellate-hairs. Inflorescence axis
robust, 1.52 mm thick, densely flowered, covered by a mixture of tufts of long hairs and
tiny stellate hairs; bracts subulate, caducous, 45(8)mm long. Flowers white, scented;
calyx oblong-obovate, 34(5) mm long; petals white to cream, glabrous, spathulate to
oblong, 3.55 mm long; filaments glabrous, 22.5 mm long, anthers subobovate, 0.81 mm
long; ovary densely covered with short straight hairs; style glabrous, 2.53 mm long. Fruit
depressed-globose, 2.53 mm diameter. Seed subtrigonous, 11.2 mm across.
184
CLETHRACEAE (BERHAMAN)
Distribution. Endemic to Borneo (Sabah and Sarawak). Common in the highlands in

Sabah and only known from Mt. Murut and Batu Lawi, in the Kelabit Highlands in
Sarawak.
Ecology. Hill and montane primary and secondary forests, at 8002500 m.
185
CONNARACEAE
Lesmy Tipot
Kepong, Malaysia
Hooker f., Fl. Brit. Ind. 2 (1879) 46; King, J. As. Soc. Beng. 66, 2 (1897) 1; Ridley, FMP 1 (1922)
544; Merrill, EB (1921) 291, PEB (1929) 94; Masamune, EPB (1942) 328; Leenhouts, FM 1, 5 (1958)
495; Kochummen, TFM 3 (1978) 47; Keng, OFMSP (1978) 211; Anderson, CLST (1980) 163;
Ashton, MNDTS 2 (1988) 223; Whitmore, Tantra & Sutisna, CLK 1 (1989) 49.
Mostly woody climbers or shrubs, rarely small trees. Leaves alternate or spirally arranged,
without stipules, pinnate with terminal leaflet (imparipinnate), rarely unifoliolate; base of
petioles and petiolules swollen; leaflets not always opposite, pinnately veined or 3-veined,
base often slightly peltate. Inflorescences paniculate, axillary or terminal, rarely on branches.
Flowers 45-merous, bisexual or unisexual; sepals free or joined at base; petals free;
stamens free or joined at base, in two whorls, inner (epipetalous) rudimentary; ovary
superior, usually 5-carpellate (in Connarus and Ellipanthus only 1-carpellate), ovules 2 in
each carpel, orthotropous or anatropous, placentation basal or axillar. Fruits follicular,
smooth, finely striate, or warty, reddish, stalked, with persistent calyx, splitting ventrally or
dorsally, rarely circumscise at the base or indehiscent. Seeds one per carpel, with an arillode
or sarcotesta, with or without endosperm; cotyledons thick, flat.
Distribution. A pantropical family with 16 genera and c. 300350 species, predominantly
African. In Sabah and Sarawak, represented by 6 genera with at least 27 species.
Ecology. Frequent in open areas, forest fringes and river banks in lowland rain forests.
Taxonomy. The Connaraceae was placed under the Dilleniales by Hutchinson (Fam. Fl. Pl.
1, ed. 2 (1959) 329). Keng (l.c.), however, included the Connaraceae in the Rosales,
considered a more primitive order than the Dilleniales. The Connaraceae was once thought
to be the link between the Rosales and Leguminosae but many recent workers believed an
affinity to the Sapindaceae is more natural. Some species of the Connaraceae are often
confused with those of Leguminosae, especially Derris. However, the absence of stipules in
the former distinguishes them from the latter.
Uses. Generally of no commercial value. Only the wood of Ellipanthus species is sometimes
used locally for house posts and bridges. Some species are said to have medicinal properties.
For instance, the roots of Agelaea macrophylla or akar malam, when boiled together with
roots of tapang (Koompasia malaccensis), are used traditionally to treat weakness in infants,
while young leaves of Rourea mimosoides are used for treating wounds and boils.
187
Key to genera
1.
Trees or treelets..........................................................................................................2
Woody climbers or shrubs............................................................................................3
2.
Leaves trifoliolate or pinnate........................................................1. Connarus (in part)

Leaves unifoliolate..................................................................................2. Ellipanthus
3.
Leaves trifoliolate, upper surface of leaflets with minute pits. Fruits usually warty.........
Agelaea Sol. ex Planch.
Linnaea 23 (1850) 437; Ridley l.c. 552; Merrill l.c. (1929) 95; Leenhouts l.c. 500;
Kochummen l.c. 48. Synonym: Cantanola Llanos, Mem. R. Ac. Cienc. Madr. 3, 2 (1859)
505, Masamune l.c. 328.
50 species, Africa, Madagascar and Malesia; 4 species in Sabah and Sarawak.

Woody climbers, rarely scrambling shrubs. Flowers bisexual, 5-merous; petals
much longer than sepals; stamens usually 10. Fruit papillose or rugulose to warty.
Seeds shiny black, partly covered with orange or yellow arillode.
Mainly in lowland primary and secondary forests. A. borneensis is widely distributed
in Sabah and Sarawak; A. insignis is very uncommon and in Sarawak is represented
by a few collections from Baram and Kuching areas but in Sabah is recorded only
from Tawau; A. macrophylla is endemic to Sarawak, whereas A. trinervis is endemic
to Sabah.
Leaves imparipinnate, upper surface of leaflets not pitted. Fruits smooth.....................4
4.
Leaflets with emarginate apex.......................................................................................

Roureopsis Planch.
l.c. 423; Masamune l.c. 331; Leenhouts l.c. 505; Kochummen l.c. 52. Synonym: Taeniochlaena
Hook. f. in Bentham & Hooker f., Gen. Pl. 1 (1862) 433, Masamune l.c. 332.
10 species, West Africa, Northern Burma, Southern China, Sumatra, Peninsular

Malaysia, Java, and Borneo; 2 species in Sabah and Sarawak, viz. R. acutipetala
and R. emarginata.
Woody climbers, sometimes scandent shrubs. Inflorescences axillary, racemose or
paniculate; bracts lanceolate, densely appressed hairy. Flowers bisexual, longstalked, usually 5-merous; petals linear; stamens usually 10. Fruits obliqueellipsoid, opening by a ventral slit, red, with persistent calyx. Seeds ellipsoid, hilum
partly or entirely covered by a yellow fleshy arilloid.
Apex of leaflets not emarginate....................................................................................5
5.
Fruits velvety hairy, pear-shaped. Leaflets usually more than 10 pairs...........................

Cnestis Juss.
Gen. (1789) 374; Ridley l.c. 553; Merrill l.c. (1929) 96; Masamune l.c. 329; Leenhouts l.c.
497; Kochummen l.c. 48.
About 40 species, predominantly tropical Africa and Madagascar. Two species in

Sabah and Sarawak.
Woody climbers or scandent shrubs. Inflorescenses axillary or in fascicles from
knobs on branches. Flowers 5-merous; stamens 10; carpels 5. Fruits velvety brown,
pear-shaped, stipitate. Seeds flattened bean-shaped, partially covered by a
188
CONNARACEAE (LESMY)
sarcotesta at the base. Found mainly in a primary and secondary mixed dipterocarp
forests to 500 m.
Fruits glabrous, finely striate, pod-like or ellipsoid to ovoid. Leaflets less than 10 pairs
....................................................................................................................................6
6.
Carpel only one per flower; calyx in fruit accrescent and hard. Fruit ellipsoid, base not
stipitate..........................................................................................................................
Rourea Aubl.
Hist. Pl. Guiane 1 (1775) 467, t. 187; Ridley l.c. 549; Merrill l.c. (1929) 95; Leenhouts l.c.
510; Kochummen l.c. 51. Synonym: Santaloides L. ex Kuntze, Rev. Gen. Pl. 1 (1891) 155
(as Santalodes), Merrill l.c. (1921) 291, Masamune l.c. 331.
About 100 species; Central and South America, Malesia, NE Australia and
Melanesia; 4 species in Sabah and Sarawak.
Woody climbers, shrubs, very rarely small trees. Leaves very variable in size and
shape. Inflorescences axillary. Flowers 5-merous; sepals ovate, imbricate, margin
ciliate; petals twice or more longer than sepals; stamens 10, joined at base. Fruits
usually 1 per flower, ellipsoid to ovoid, splitting ventrally. Seeds with the testa
partly or almost entirely fleshy or seeds enveloped by fleshy arillode.
Mainly found in low altitude, in primary and secondary mixed dipterocarp
rainforests, along forest edges, river banks, and roadsides.
Carpels more than one per flower. Calyx in fruit not accrescent. Fruit pod-like, base
narrowly stipitate.........................................................................1. Connarus (in part)
1. CONNARUS L.
(Greek, konaros, a plant name)
Sp. Pl. 2 (1753) 675, Gen. P1. ed. 5 (1754) 305; Bentham & Hooker f. l.c. (1862) 432; King l.c. (1897)
2; Ridley l.c. 544; Merrill l.c. (1921) 291, l.c. (1929) 94; Masamune l.c. 329; Leenhouts l.c. 525;
Kochummen l.c. 49; Anderson l.c. 163; Ashton l.c. 223.
Usually woody climbers, rarely shrubs or small trees. Leaves imparipinnate, sometimes
trifoliolate, rarely unifoliolate; leaflets almost always pellucid-glandular punctate. Inflorescences terminal or axillary rarely on the branches or stems. Flowers bisexual, fragrant, 5merous; sepals, petals, stamens punctate, seen as dark gland-dots in herbarium materials;
sepals thick and fleshy, not accrescent in fruit; petals nearly always hairy; stamens 10,
joined at base; ovary 1-carpellate, stigma capitate. Fruits pod-like, opening lengthwise, base
narrowed into a stipe, with small persistent calyx, apex short-beaked or curved. Seed 1,
shiny black, basal part partially enveloped by a fleshy, yellow arillode.
Distribution. About 100 species, pantropical. 13 species in Sabah and Sarawak, including
two unnamed taxa (indicated as Connarus sp. A and Connarus sp. B in the key given
below). Of these only one species, C. agamae, is a small tree.
189
Key to Connarus species

1.
Small trees.......................................................................................Connarus agamae

Climbers, scandent or erect shrubs...............................................................................2
2.
Inflorescences on stems or on leafless twigs.................................................................3

Inflorescences terminal or almost so on leafy shoots....................................................4
3.
Leaflets oblong-elliptic, apex shortly acuminate; all veins distinctly raised below, intercostal veins scalariform.................................................................................................
C. impressinervis B.C. Stone
Malays. For. 43, 2 (1980) 255.
Usually shrub, rarely small tree reaching 5 m tall and 5 cm diameter. Inflorescences on
short condensed panicles, peduncles stout. Fruits obliquely obovoid, glabrous,
stipitate, sparsely tomentose, slightly compressed, bright orange-red, drying brown,
apex rounded.
Primary mixed dipterocarp forest; collected several times from Bukit Raya, Kapit
District, Sarawak.
Leaflets narrowly lanceolate, apex distinctly emarginate; veins not raised below, intercostal veins not scalariform............................................................................................
Connarus sp. A.
A climber. Leaves trifoliolate, blade distinctly swollen at base, leatherly. Fruits
borne on branches, smooth, drying chestnut brown. Seed 1, arillode covering
mostly the lower part.
Mixed dipterocarp forest at 740 m. Collected once (S. 36403) from Bt. Goram,
Song, Kapit District, Sarawak.
4.
Leaflets hairy at least on the midrib below or at young stage........................................9
5.
Leaflet attachment distinctly peltate; lateral veins 1618 pairs, straight and close together; petiolule 2.54 cm long.....................................................................................
Connarus sp. B.
A climber. Leaflets 12 pairs, broadly ovate. Inflorescences terminal.
One collection (S. 44912) from kerangas forest in Bukit Sadok, Ulu Sekrang, Sri
Aman, Sarawak.
Leaflet attachment not peltate; lateral veins less than 10 pairs, usually curved; petiolules
much shorter................................................................................................................6
6.
Leaflets drying reddish brown. Fruits obovoid; pericarp coarse, thick and woody, c. 2
mm thick.......................................................................................................................
C. grandis Jack
Mal. Misc. 2, 7 (1822) 40; Masamune l.c. 329; Leenhouts l.c. 529. Synonym: Connarus
ellipticus King l.c. 7, Merrill l.c. (1929) 95.
Woody climber, rarely shrub. Branches glabrous. Leaflets 12 pairs, obovate to

lanceolate. Inflorescences usually densely fulvous-tomentose.
190
CONNARACEAE (LESMY)
Primary and secondary mixed dipterocarp forests, also in open areas and along
river banks. Distributed widely in Sabah and Sarawak, fairly common.
Leaflets not drying as above. Fruits ellipsoid or spindle-shaped; pericarp thin, not
coarse..........................................................................................................................7
7.
Leaflets drying pale green and purplish brown on midrib and lateral veins. Fruits not
stipitate..........................................................................................................................
C. lucens Schellenb.
Bot. Jahrb. 59 (1924) 36; Leenhouts l.c. 539.
Glabrous climber. Leaflets 34 pairs, obovate to oblong-lanceolate; lateral veins

drying purplish brown. Inflorescences broadly branched. Fruits trapezoid to
oblique-ellipsoid, slightly flattened, not stipitate, drying purplish brown.
Recorded from Saribas and Kapit, Sarawak.
Leaflets not drying as above. Fruit stipitate..................................................................8
8.
Fruits obliquely spindle-shaped, shortly stipitate. Reticulation of intercostal veins

conspicuous...................................................................................................................
C. monocarpus L.
Sp. Pl. (1753) 675; Leenhouts l.c. 538 (ssp. malayensis). Synonyms: C. falcatus Blume,
Mus. Bot. Lugd. Bat. 1 (1850) 266, Masamune l.c. 329; C. densiflorus Merr., Philip. J. Sc.
13 (1918) Bot. 70, Merrill l.c. (1921) 292, l.c. (1929) 95.
Woody climber. Leaflets 24 pairs. Inflorescences many-flowered. Sepals usually

distinctly keeled, pubescent outside, glabrous inside.
Uncommon; in Sabah once found along the seashore in Lahad Datu; in Sarawak,
collected from forest edges and river banks, and once on limestone.
Fruits ellipsoid, more or less bulging, stipe 0.51 cm long. Reticulation of intercostal
veins not conspicuous....................................................................................................
C. winkleri Schellenb.
l.c. (1924) 38; Masamune l.c. 330; Leenhouts l.c. 540. Synonym: C. pachyphyllus Merr. l.c.
(1918) 71, l.c. (1921) 292, Masamune l.c. 330.
Woody climber, to 20 m high. Leaflets 23 pairs. Inflorescences to 5 cm long,

laxly branched. Sepals ovate; petals ovate-lanceolate, thinly tomentose on both
surfaces.
Generally lowland forests. Few collections from Sabah, and only one from Sarawak.
9.
Leaflet attachment usually subpeltate, lateral veins 1018 pairs, straight to slightly
curved at margin............................................................................................................
C. euphlebius Merr.
J. Str. Br. R. As. Soc. 85 (1922) 200, l.c. (1929) 94; Masamune l.c. 329; Leenhouts l.c. 527.
Woody climber to 25 m high, sometimes a scandent shrub. Branchlets, petioles,

rachises and petiolules densely ferruginous hairy. Leaflets 14 pairs, petiolules 25
mm long. Inflorescences densely ferruginous-tomentose. Fruits obovoid and rather
flattened, shortly stipitate; pericarp densely minutely ferruginous-tomentose.
Generally lowland forests. Uncommon in Sabah and Sarawak, represented by only
three collections.
191
Leaflet attachment not subpeltate, lateral veins less than 10 pairs, curved and looped at
margin.......................................................................................................................10
10. Leaflets progressively increasing in size from basal pair upwards. Fruits glabrous.........
C. odoratus Hook. f.
Trans. Linn. Soc. 23 (1860) 72; Merrill l.c. (1921) 292, l.c. (1929) 95; Leenhouts l.c. 533.
Synonym: C. hebephyllus King l.c. (1897) 5, Merrill l.c. (1921) 292.
Climber or scandent shrub. Branches, leaves, petioles, rachis and petiolules densely
tomentose when young, later glabrescent. Inflorescences to 20 cm long, widely
branched and densely tomentose. Fruits oblique-ellipsoid, glabrous; pericarp thin,
minutely wrinkled outside, densely pubescent inside.
Lowland forests and thickets. Widespread in Sabah and Sarawak.
Leaflets equal in size. Fruits thinly or densely ferruginous-tomentose........................11
11. Leaflets densely ferruginous-pubescent beneath. Fruits not compressed..........................
C. villosus Jack
l.c. 38; Masamune l.c. 330; Leenhouts l.c 531. Synonym: C. plumoso-stellatus Merr. l.c.
(1918) 72, l.c. (1921) 292.
Woody climber or scandent shrub. Branches, leaves and inflorescences densely

pubescent. Leaflets 14 pairs. Inflorescences 1030 cm long, few-flowered. Fruits
ellipsoid, not stipitate; pericarp rather thin, outside densely orange-brown
pubescent, inside glabrous.
Primary mixed dipterocarp and kerangas forests at low altitudes. Found mainly in
the Kuching and Samarahan Divisions in Sarawak but only one record from Sabah.
Leaflets not so. Fruit compressed...............................................................................12
12. Fruits oblique-ellipsoid, shortly stipitate; hairs branched, usually stellate.......................
C. cullionensis Merr.
Philip. J. Sc. 4 (1909) Bot. 120; Leenhouts l.c. 531. Synonym: C. stellatus Merr. l.c. (1909) 119,
Masamune l.c. 330.
Woody climber or scandent shrub. Branchlets densely ferruginous-tomentose, later

glabrous. Leaflets 25 pairs, tomentose.
Usually occurs in open areas in the lowland mixed dipterocarp forests.
Fruits oblique-pyriform, stipitate, stipe to 1.5 cm long; hairs simple, not stellate............
C. semidecandrus Jack
l.c. 39; Leenhouts l.c. 534. Synonyms: C. mutabilis Blume l.c. 269; C. borneensis Merr. l.c.
(1918) 69, l.c. (1921) 292, Masamune l.c. 329; C. jackiana Schellenb. l.c. 40, Masamune l.c.
329.
Large woody climber or scandent shrub. Leaflets 13 pairs, sometimes 5 pairs.

Inflorescences terminal, to 35 cm long, broadly branched, many-flowered, minutely tomentose. Fruits compressed; pericarp thin, minutely ferruginous-pubescent,
glabrescent outside, inside densely pubescent.
Primary and secondary mixed dipterocarp forests especially in open areas along
forest edges and river banks, also in clearings along beaches and sometimes in
swampy habitats, on granite and limestone. Widely distributed in Sabah and
Sarawak.
192
193
194
CONNARACEAE (LESMY)
Connarus agamae Merr.
Fig. 1.
(Jos Agama, Deputy Conservator of Forests, British North Borneo, 18891982)

l.c. (1918) 68, l.c. (1921) 291, l.c. (1929) 94; Masamune l.c. 329; Leenhouts l.c. 527. Type: Agama
422, British North Borneo, Tawau (holotype K).
Small tree, rarely reaching 20 m high and 20 cm diameter; sometimes shrub, rarely climber.
Twigs thinly pubescent, glabrescent. Leaves with 12 pairs of leaflets, sometimes unifoliolate especially below the inflorescence; leaflets oblong, slightly oblique, 1120 x 410
cm, terminal ones ovate, thinly papery, glabrous above, drying reddish brown, densely
ferruginous-pilose beneath especially on midrib; base rounded to acute, apex blunt; lateral
veins 815 pairs, straight, strongly looped and closed toward the margin but not joining;
intercostal veins reticulate, inconspicuous above. Inflorescences densely ferruginouspubescent. Flowers: sepals ovate, brown-pilose on both sides; petals linear, punctate on
both sides; stamens all fertile, connate at base. Fruits obovate, 5 x 23.5 cm, beak acute at
4/5 of the height; pericarp glabrous, shiny and slightly wrinkled, woody, densely fulvoustomentose inside; stipe 11.5 cm long.
Distribution. Endemic to Borneo. In Sabah, recorded from Sandakan and Mostyn, but not
yet recorded from Sarawak.
Ecology. In primary mixed dipterocarp forest to 300 m.
2. ELLIPANTHUS Hook. f.
(Greek, ellipes = defective; anthos = flower;
referring to the defective or imperfect development of 5 of the 10 stamens)
l.c. (1862) 434, l.c. (1879) 55; King l.c. 8; Ridley l.c. 548; Merrill l.c. (1921) 291, l.c. (1929) 96;
Masamune l.c. 330; Leenhouts l.c. 520; Kochummen l.c. 50; Anderson l.c. 163; Ashton l.c. 224;
Whitmore, Tantra & Sutisna l.c. 49. Synonym: Pseudellipanthus Schellenb. in Mez, Bot. Arch. 1
(1922) 314, Masamune l.c. 331.
Shrubs or small trees reaching 25 m tall and 25 cm diameter. Twigs tomentose, at least when
young. Leaves alternate, unifoliolate, with a prominent joint at the base. Inflorescences
axillary, paniculate or clustered; bracts caducous, lanceolate, small. Flowers 45-merous,
protandrous, unisexual (and then plants dioecious) or bisexual; sepals densely hairy outside, valvate in bud; petals free, imbricate in bud; stamens twice as many as petals, connate
at base, episepalous ones well-developed, epipetalous ones rudimentary; ovary 1-carpellate,
flattened ovoid, style slender, stigmas disk-shaped to bilobed. Fruits densely tomentose,
apex shortly pointed, base stipitate; pericarp woody; persistent calyx not accrescent. Seed
one, ellipsoid, shiny black, covered by a yellowish orange arillode at base.
Distribution. About 10 species; Africa, Madagascar, Sri Lanka, continental SE Asia and
Malesia. 2 species in Sabah and Sarawak.
Ecology. Lowland mixed dipterocarp forest and mixed swamp forest.
195
Key to Ellipanthus species

Flowers predominantly 4-merous, unisexual. Leaf-blade usually glabrous above, densely
ferruginous-tomentose beneath, base rounded.................................................1. E. beccarii
Flowers predominantly 5-merous, usually bisexual. Leaf-blade usually tomentose above (at
least on the midrib), glabrous or thinly fulvous-tomentose beneath, base cuneate or subcordate.......................................................................................................2. E. tomentosus
1. Ellipanthus beccarii Pierre

Fl. Coch. 5 (1898) t. 378; Merrill l.c. (1921) 291; Leenhouts l.c. 524; Anderson l.c. 164; Ashton l.c.
225; Whitmore, Tantra & Sutisna l.c. 49. Type: Beccari PB 296, Sarawak, Matang (K).
Small dioecious tree or shrub, rarely reaching 10 m tall, 15 cm diameter. Leaves elliptic to
ovate, sometimes lanceolate, glabrous above, densely ferruginous-tomentose beneath, 715
x 35 cm; base rounded, peltate or not, apex acuminate; lateral veins 810 pairs, curved,
looped and joined; petiole jointed at leaf-base, c. 1 cm long. Flowers unisexual, 4-merous.
Fruits shortly stipitate, stipe to 1 cm long. Seed with a small cupular arilloid.
Key to varieties
Leaf-base not peltate.............................................................................................................
var. beccarii Leenh.
l.c. 524; Anderson l.c. 164; Whitmore, Tantra & Sutisna l.c. 49. Synonyms: E. mindanaensis (non
Merr.) Merr., J. Str. Br. R. As. Soc. 76 (1917) 84, l.c. (1921) 291, l.c. (1929) 96; Pseudellipanthus
beccarii (Pierre) Schellenb. l.c. (1922) 314; Dichapetalum tetramerum Ridl., Kew Bull. (1938)
234.
Known only in the Kuching area, Sarawak on leached yellow soils in mixed dipterocarp
forest, and Kalimantan.
Leaf-base peltate...................................................................................................................
var. peltatus (Schellenb.) Leenh.
l.c. 524; Anderson l.c. 164; Whitmore, Tantra & Sutisna l.c. 49. Basionym: Pseudellipanthus
peltatus Schellenb. l.c. (1922) 314.
Endemic to Borneo (Sabah, Sarawak, Brunei, Kalimantan). In Sabah uncommon; in

Sarawak frequent in mixed dipterocarp forest on fertile clay soils, particularly on basic
volcanic rocks.
Vernacular names. Sarawakkelana (Kenyah), merinang (Iban).
2. Ellipanthus tomentosus Kurz
Fig. 2.
(Latin, tomentosus = thickly and evenly covered with short matted hairs; the leaves)
J. As. Soc. Beng. 41, 2 (1872) 305; Hooker f. l.c. 56; Leenhouts l.c. 521; Kochummen l.c. 50;
Anderson l.c. 164; Ashton l.c. 225; Whitmore, Tantra & Sutisna l.c. 49. Type: Wallich Cat. 8551,
Lower Burma, Moulmain (holotype K).
196
CONNARACEAE (LESMY)
subsp. tomentosus var. luzoniensis (Vidal) Leenh.

l.c. 524; Whitmore, Tantra & Sutisna l.c. 49. Basionym: E. luzoniensis Vidal, Rev. Pl. Vasc. Filip.
(1886) 104. Synonyms: Connarus urdanetensis Elmer, Leafl. Philip. Bot. 7 (1915) 2594; E.
burebidensis Elmer l.c. 2596; E. vidalii Elmer l.c. 2596; E. longifolius Merr., Philip. J. Sc. 17 (1921)
Bot. 262; E. urdanetensis (Elmer) Merr., En. Philip. 2 (1923) 241; E. sarawakensis Schellenb., Pfl.
R. Heft 103 (1938) 185.
Shrub or tree to 20 m tall, 25 cm diameter. Bark smooth, greyish brown; inner bark
reddish brown. Sapwood pale yellow. Leaflets elliptic to lanceolate, 720 x 49 cm,
chartaceous, glabrous or thinly fulvous-tomentose below; base cuneate, sometimes rounded
or subcordate, apex acuminate; midrib tomentose above; lateral veins 57 pairs, looping
near margin; intercostal veins conspicuous below, very faint above; petiole jointed at base,
0.53 cm long. Flowers white, bisexual, 5-merous. Fruits stipitate, stipe c. 3.5 cm long,
apex pointed. Seeds c. 5 mm long, covered with faintly lobed arilloid.
Vernacular name. Sarawak-kelin (Melanau).
Distribution. Borneo, the Philippines and Central Celebes. In Sarawak uncommon, distributed
around Miri, Bintulu, Sibu, Sarikei and Kuching. In Sabah, recorded from Lahad Datu and
Tawau.
Ecology. Locally frequent in primary mixed dipterocarp forest and peat swamp forest.
Uses. The wood is considerably hard and durable, used for local construction works such as
bridges and house posts.
197
CORNACEAE
R.C.K. Chung
Kepong, Malaysia
Ridley, FMP 1 (1922) 889 (excluding Alangium, Aralidium and Nyssa); Merrill, EB (1921) 459, PEB
(1929) 233; Danser, Blumea 1 (1934) 46; Backer & Bakhuizen f., FJ 2 (1965) 158; Matthew, Blumea
23 (1976) 51, FM 1, 8 (1977) 85; Kochummen, TFM 3 (1978) 53; Anderson, CLTS (1980) 164;
Trees, shrubs, or rarely herbs. Leaves opposite, subopposite or alternate, simple, entire or
rarely serrate, pinnately veined, exstipulate, usually petiolate. Flowers regular (actinomorphic), 45-merous, bisexual or sometimes unisexual, arranged in axillary or terminal
cymes, panicles or heads; calyx-tube adnate to the ovary, 45-lobed or subtruncate, persistent in
fruit; petals 45, free, valvate or imbricate in buds; stamens short and equal in number with
the petals, 2-locular, dehiscing lengthwise; disc large, cushion-shaped at the top of the
ovary, generally persistent in fruit; ovary inferior, 14-locular, ovule 1 in each locule,
pendulous, with dorsal or ventral (in Curtisia and Mastixia) raphe, style simple or lobed.
Fruit usually a drupe, rarely a berry (in Aucuba and Griselinia). Seeds usually 12, rarely
4, with small elongate embryo embedded in copious oily endosperm.
Distribution. About 815 genera and 100 species; mainly in the temperate zones and in the
tropics (rare in Africa and South America). In Sabah and Sarawak, represented by the genus
Mastixia with 7 species.
Ecology. Insect-pollination is probably a general phenomenon in the family, except in
Griselinia litoralis which has been reported to be wind-pollinated.
Uses. In Sabah and Sarawak, very little is known about the utilization of members of the
family. However, the fruit of Chamaepericlymenum suecicum is eaten by the Eskimos. In
France, Cornus mas (Cornelian cherry) is grown as ornamentals, and the fruit is edible and
used as a source of raw material for making "Vin de Cornoulle". The Assegay tree or Cape
Lance wood (Curtisia faginea) produces durable, red to brown timber and used for making
furnitures in South Africa. Aucuba japonica is a popular evergreen garden shrub widely
cultivated in temperate and subtropical countries.
Taxonomy. The taxonomic and systematic status of the Cornaceae remains controversial
(Matthew l.c. 1976, l.c. 1977; Mabberley, The Plant Book (1987) 146; and Brummitt,
Vascular Plant Families & Genera (1992) 543). In the Malesian region, Ridley (l.c.) and
Keng (OFMSP, 1969) included Alangium, Aralidium, Mastixia and Nyssa in the Cornaceae,
whereas van Steenis (Checklist Gen. Names Mal. Bot. (1987) 50) placed Alangium in its
own family (Alangiaceae), Aralidium in the Araliaceae, and Nyssa in the Nyssaceae, leaving
Mastixia as the sole representative of the family in the Malesian region.
199
MASTIXIA Blume
(Greek , mastix = whip; the whiplike apex of the petals)
Bijdr. (1825) 654, Mus. Bot. Lugd. Bat. 1 (1850) 256; Bentham & Hooker f., Gen. Pl. (1887) 950;
Harms in Engl. & Prantl, Pfl. Fam. 3 (1898) 262; Wangerin in Engl., Pfl. Reich. 4, 229, 41 (1910) 19;
Ridley l.c. 889; Merrill l.c. (1921) 459, l.c. (1929) 233; Danser l.c. 47; Backer & Bakhuizen f. l.c. 159;
Matthew l.c. (1976) 51, l.c. (1977) 85; Kochummen l.c. 53; Whitmore, Tantra & Sutisna l.c. 49; Ng,
MFFSS 1 (1991) 47.
Trees, usually without buttresses. Bark grey to grey-brown, smooth with horizontal rings,
rarely cracked to shallowly fissured, often exuding white resin when bruished; inner bark
orange-yellow, gritty, granular, with strong smell of sugarcane water. Sapwood soft,
yellowish white or white. Leaves alternate, subopposite or opposite, margin entire; midrib
sunken on adaxial surface, prominent below; lateral veins usually distinct on the abaxial
surface. Inflorescences terminal or sometimes axillary cymose-panicles. Flowers bisexual,
in triads, sessile, subtended by tiny, persistent bracts; calyx-tube obconical or barrel-shaped
or cup-shaped, lobes spreading or not, broader than long or sometimes appear as minute
sharp tips, persistent in fruit; petals thick, concave, valvate in bud, inflexed and strongly
connate at the upper parts; stamens attached below the disc, abutting on and alternating with
disc-lobes, opposite the calyx-lobes, filaments subulate, flattened and tapered toward its
upper part, anthers cordate, dorsifixed, introrse-latrorse; ovary turbinate, one-locular,
surmounted by a fleshy, lobed and grooved disc which is sometimes persistent in fruit; style
very short, stout, ribbed; stigma punctiform, sometimes bifid or 45-lobed, reflexed, some
persistent in fruit. Fruit a drupe, ovoid, ellipsoid or oblong, surmounted by calyx-tube and
crowned by the persistent disc (the exposed part of the fruit); pericarp (exocarp and
mesocarp) thin or thick, turning to dark purple or blue when ripe; fruit-wall formed by
calyx-tube and pericarp; endocarp stony. Seed 1, ovoid or ellipsoid, with membranous testa;
endosperm large and U-shaped in transverse section; embryo small and straight, cotyledons
thin and foliaceous; germination epigeal.
Distribution. About 13 species; from the western Ghats and Sri Lanka, NE India, Bhutan,
Burma, Thailand, Indo-China, S Yunnan, Hainan, through Malesia to New Britain and the
Solomon Islands. 7 species (including 2 endemic) are found in Sabah and Sarawak.
Ecology. Found mainly in valleys, slopes or ridges of primary and secondary mixed dipterocarp
forests, often in moist habitats, from sea-level to 2200 m.
Uses. Although the trees may reach a considerable size, the scattered occurrence does not
contribute to their general use as timber. In addition the timber has little commercial value
and therefore only used for packing cases and temporary constructions.
Taxonomy. Mastixia is strictly a SE Asian genus. It was included in the Cornaceae by
Bentham & Hooker f. (l.c.) and Hutchinson (Genera of Flowering Plants 2 (1967) 42),
whereas Harms (l.c.) and Wangerin (l.c.) included it in a distinct subfamily. Wangerin
distinguished two subgenera, viz. Tetramastixia and Pentamastixia. Matthew (1976 &
1977), on the other hand, established two subgenera, Manglesia (2 species) and Mastixia
(11 species). He recognised two series, the Oppositae and Alternae within the subgenus
Mastixia based on a single character, viz. whether the first branches of the inflorescence are
opposite (or subopposite) or alternate. In the absence of flower and fruit and due to the
200
CORNACEAE (CHUNG)
presence of resin in the bole and on the cut ends of logs, the genus can be easily confused
with some species of the Dipterocarpaceae that have smooth or cracked bark, such as Vatica
species. However, in Mastixia the inner bark is thick and gritty with a strong sugarcane
smell and the wood is soft. Foresters sometimes confuse this genus with those of the
Lauraceae (medang) because of the gritty inner bark and strong aromatic smell; however
medang has no resin.
Key to Mastixia species

1.
Leaf surface velvety hairy below, if glabrescent then midrib hairy................................2

Leaf surface glabrous...................................................................................................3
2.
Leaves alternate. Twigs and stalks distinctly grooved.......................4. M. macrocarpa

Leaves opposite. Twigs and stalks not so............................................7. M. trichotoma
3.
Leaves glaucous below...............................................................................3. M. glauca

Leaves not glaucous below...........................................................................................4
4.
Leaf-margin wavy when dry; stalks 23 cm long...............................2. M. eugenioides

Leaf-margin not wavy when dry; stalks shorter than 2 cm............................................5
5.
Leaves thickly leathery; margin curled inwards. Twigs dark brown to black when dry
...........................................................................................................5. M. pentandra
Leaves thinly leathery; margin not curled inwards. Twigs grey or grey-brown when dry
....................................................................................................................................6
6.
Leaf lateral veins looping near margin. Flower buds up to 3 mm diameter; petals
densely hairy on the abaxial surface......................................................1. M. cuspidata
Leaf lateral veins not looping near margin. Flower buds up to 2 mm diameter; petals
glabrous on the abaxial surface...............................................................6. M. rostrata
1. Mastixia cuspidata Blume

(Latin, cuspidatus = sharp-pointed; the leaf apex)
l.c. (1850) 256; Miquel, Fl. Ind. Bat. I, 1 (1855) 772; Ridley l.c. 891; Merrill l.c. (1921) 459; Danser
l.c. 55 (excl. var. margarethae); Matthew l.c. (1976) 79, l.c. (1977) 95; Kochummen l.c. 54;
Whitmore, Tantra & Sutisna l.c 49. Type: Korthals, s.n., Sumatra, West Coast (lectotype L;
isolectotype U). Synonyms: Mastixia pentandra Blume var. cuspidata (Blume) Miq. l.c. (1858) 1095;
Mastixia bracteata Clarke in Hooker f., Fl. Brit. Ind. 2 (1879) 746.
Tree to 40 m tall, and 40 cm diameter. Bark greyish to chocolate-brown, smooth to

shallowly fissured; inner bark yellowish to brownish, mottled. Sapwood yellowish to
brownish. Twigs subglabrous, grey-brown. Leaves alternate, subopposite or opposite, stalk
slender, 0.51.5 cm long; blades narrowly obovate, elliptic or oblong, 3.59 x 1.53.5 cm,
thinly leathery, glabrous; base cuneate, apex cuspidate with a tip 0.51(1.5) cm long,
oblique; lateral veins (4)5 pairs, curving near the margin and joining with next one to form
looped intramarginal veins, sunken above; intercostal veins faint to inconspicuous below,
201
inconspicuous above. Inflorescences to 4 cm long, subglabrous to puberulous. Flowers 5merous, green to yellow, buds to 3 mm diameter; calyx-tube densely silky-hairy, with 5
lobes, broader than long, subglabrous; petals 1.52 x 0.51 mm, densely silky-hairy on
abaxial surface; stamens 5, filament 13.2 mm long; disc yellowish. Fruits oblong, 1.53 x
0.51.3 cm; fruit-wall thin; persistent disc well-exposed; persistent calyx-lobes
inconspicuous. Seeds ellipsoid, 1.52.5 x 0.41 cm.
Vernacular name. Sarawakbiansu gunong (Iban).
Distribution. Sumatra, Peninsular Malaysia and throughout Borneo. Uncommon in Sabah
and Sarawak. In Sabah, recorded from Sipitang, Sandakan, Lahad Datu, and Tawau. In
Sarawak, known from Mt. Matang (1st Div.), Sri Aman (2nd Div.), Mt. Dulit near Long
Kapa (4th Div.), and Belaga (7th Div.).
Ecology. In primary and secondary mixed dipterocarp forests, to 900 m. Flowering in
FebruaryMarch and JulyOctober; and fruiting in January, March, JulyAugust and
October.
Sterile specimens of small-leaved M. cuspidata are difficult to distinguish from M. rostrata
ssp. caudatifolia.
2. Mastixia eugenioides Matthew

(leaves resembling that of Eugenia)
l.c. (1976) 73, l.c. (1977) 93; Whitmore, Tantra & Sutisna l.c. 50. Type: Beccari PB 2033, Sarawak,
Mt. Matang (holotype FI; isotype L).
Tree to 30 m tall and 30 cm diameter. Bark greyish to yellowish brown, smooth; inner bark
yellowish brown. Twigs glabrous. Leaves opposite, stalk green to greenish yellow, stout,
1.53 cm long; blades elliptic to oblong-elliptic, 415 x 25.5 cm, thickly leathery,
glabrous; base cuneate, margin wavy after drying, apex acuminate to caudate; lateral veins
57 pairs, flattened above; intermediate lateral veins and intercostal veins prominent below.
Inflorescences to 8 cm long, glabrous. Flowers 4-merous, buds about 1.52 mm diameter;
calyx-tube glabrous, with 4 lobes, broader than long, glabrous; petals 4, 11.5 x 0.50.8
mm, glabrous on abaxial surface; stamens 4, filament 0.61 mm. Fruits green turning to
purple when ripe, ovoid (young stage) to oblong, 22.5 x 11.5 cm; fruit-wall thin;
persistent disc not well-exposed; persistent calyx-lobes inconspicuous. Seed ellipsoid, 1.9
2.4 x 0.81.1 cm.
Vernacular name. Sarawaksempetan (Kenyah).
Distribution. Endemic to Borneo (Sarawak, Sabah, Brunei, and Kalimantan). In Sarawak,
recorded from Bukit Senyandang Lingga (2nd Div.), Lambir Hills National Park; Similajau
FR (4th Div.), Lawas (5th Div.), and Kapit (7th Div.). In Sabah, known from Tenom,
Sandakan and Tawau.
Ecology. In primary mixed dipterocarp forests, from lowlands up to 1200 m. Flowering in
JulyAugust, fruiting in SeptemberDecember.
202
CORNACEAE (CHUNG)
3. Mastixia glauca Matthew

(Greek, glaukos = bluish-grey; the lower surface of the leaves)
l.c. (1976) 76, l.c. (1977) 95; Whitmore, Tantra & Sutisna l.c. 50. Type: Hj. Bujang S. 13481,
Sarawak, 1st Div., Kuching, G. Santubong East (holotype L; isotypes K, SAR).
Tree to 15 m tall, 25 cm diameter. Twigs yellowish, glabrous. Leaves alternate, stalk stout,
23.5 cm long; blades obovate, 716 x 48.5 cm, thickly leathery, glaucous and waxy
below, glabrous; base obtuse, margin wavy when dry, apex apiculate; lateral veins 45
pairs with intermediate lateral veins faint to inconspicuous; intercostal veins incon-spicuous.
Inflorescences subglabrous to puberulous. Flowers 4(5)-merous, greenish yellow; buds 2.8
3.2 mm diameter; calyx-tube puberulous, with 4(5) lobes, as long as wide, puberulous;
petals 4(5), 1.62.1 x 0.61.2 mm, hairy on abaxial surface; stamens 45, filament 0.91.2
mm long. Fruits & seeds unknown.
Distribution. Endemic to Sarawak, and so far only known from Mt. Santubong.
Ecology. In mixed dipterocarp forest at about 90 m. Flowering in AprilMay.
4. Mastixia macrocarpa Matthew

(Greek, macros = large, carpos = fruit)
l.c. (1976) 75, l.c. (1977) 94; Whitmore, Tantra & Sutisna l.c. 50. Type: Benang S. 24745, Sarawak,
Miri, Bakam Road (holotype A; isotypes BO, K, KEP, L, MEL, NY, SAN, SAR, SING).
Tree to 20 m tall, 20 cm diameter. Twigs covered with sticky resin when fresh, rusty brown,
rough-hairy and distinctly grooved. Leaves alternate, stalk stout, 46 cm long, grooved,
hairy like the twigs; blades elliptic-oblong to oblong, 1326 x 5.513 cm, thinly leathery,
villous below; base cuneate to asymmetric, apex acute to acuminate; lateral veins 710
pairs, flatten above, prominent below; intercostal veins prominent, scalariform or
reticulate. Inflorescences to 9 cm long, densely hairy. Flowers 5-merous, densely hairy,
buds c. 4 mm diameter; calyx-tube densely hairy, with 5 lobes, broader than the length,
villous; petals 5, 1.82.3 x 0.81.2 mm, velvety on abaxial surface; stamen 5, filament 11.5
mm long. Fruits pale green, oblong-ovoid, c. 4 x 2 cm; fruit-wall thin; persistent disc not
well-exposed; persistent calyx-lobes prominent, 45 mm long. Seeds ellipsoid, c. 3.8 x 1.8
cm.
Distribution. Borneo, Philippines (Luzon). In Sarawak known from Miri, Bakam Road; not
yet recorded from Sabah, Brunei, and Kalimantan.
Ecology. In mixed dipterocarp forest, at about 75 m. Flowering in October, fruiting in June.
203
5. Mastixia pentandra Blume

(Greek, penta = five, -andrus = male; with 5 stamens)
l.c. (1825) 654; Danser l.c. 49; Backer & Bakhuizen f. l.c. 159; Matthew l.c. (1976) 80, l.c. (1977) 95;
Whitmore, Tantra & Sutisna l.c. 50. Type: Blume, s.n. (= Leiden no. 901, 169375), Java (lectotype L;
isolectotypes NY, W).
subsp. scortechinii (King) Matthew

l.c. (1976) 80, l.c. (1977) 54. Basionym: Mastixia scortechinii King, J. As. Soc. Beng. 71, 1 (1902)
72. Type: Scortechini 1971, Perak (lectotype K; isolectotypes BM, CAL, G, L, P). Synonyms:
Mastixia megacarpa Ridl. l.c. 891; Mastixia parvifolia Hallier f., Beih. Bot. Centralbl. 34, 2 (1916) 41,
Merrill l.c. (1921) 459.
Tree to 40 m tall, 80 cm diameter; buttresses short. Bark grey-brown, smooth to shallowly

fissured with horizontal rings and lenticels in rows; inner bark yellowish brown to dark
yellow, with strong smell of sugarcane. Sapwood pale yellow to brownish. Twigs glabrous,
dark brown to black. Leaves alternate, spiral or subopposite, stalk stout, 12 cm long;
blades obovate to oblong or elliptic, 512 x 2.55 cm, thickly leathery, glabrous; base
cuneate, margin curled inwards, apex acute or acuminate but sometimes caudate; lateral
veins 46 pairs, flattened above; intercostal veins and reticulations faint to inconspicuous
on both surfaces. Inflorescences to 8 cm long, puberulous to villous. Flowers (4)5merous, greenish or yellowish white, buds c. 2 mm diameter; calyx-tube puberulous, with
(4)5 lobes, as long as wide; petals (4)5, 1.21.5 x 0.81 mm, hairy on the abaxial surface;
stamens (4)5, filament 0.50.7 mm long. Fruits green, ripening purple to bluish black,
ovoid to oblong, 1.63.5 x 0.81.2 cm; fruit-wall thick; persistent disc well-exposed and
sometimes bulging; persistent calyx-lobes inconspicuous. Seeds ovoid, 0.91.2 x 0.50.8
cm.
Vernacular names. Sabah and Sarawakkaju wulu, medang surungan (Malay).
Distribution. Thailand, Sumatra, Peninsular Malaysia, Borneo (Sabah and Kalimantan), and
Celebes. In Sabah recorded only from Mt. Kinabalu and Mesilau Hill.
Ecology. Uncommon, distributed from lowland to submontane forests to 1500 m. Flowering
and fruiting in JanuaryDecember
Taxonomy. In Matthew's treatment, Mastixia pentandra was segregated into six subspecies,
viz. pentandra, moluccana, chinensis, cambodiana, philippinensis and scortechinii. Of
these, only M. pentandra subsp. scortechinii is known from Sabah.
6. Mastixia rostrata Blume
Fig. 1.
(Latin, rostratus = with a beak, narrowed into a slender tip or point; the leaf apex)
l.c. (1850) 258; Danser l.c. 52; Backer & Bakhuizen f. l.c. 159; Matthew l.c. (1976) 73, l.c. (1977) 94;
Anderson l.c. 164; Whitmore, Tantra & Sutisna l.c. 50. Type: Blume, s.n. (= Leiden no. 901, 169
384), Java (lectotype L).
204
CORNACEAE (CHUNG)
subsp. caudatifolia (Merr.) Matthew

l.c. (1976) 74, l.c. (1977) 94. Basionym: Mastixia caudatifolia Merr. l.c. (1929) 233. Type: Elmer
21584, British North Borneo, Tawau, Elphinstone Prov. (holotype UC; isotypes A, BM, BO, GH,
HBG, K, L, NY, P, SING, U, UC, US). Synonyms: M. margarethae Wangerin, Fedde Rep. 4 (1907)
335; M. cuspidata Blume var. margarathae (Wangerin) Hallier f. l.c. 41.
Tree to 30 m tall and 50 cm in diameter. Bark greyish to chocolate-brown, smooth to

occasionally shallowly fissured; inner bark yellowish to pale orange-yellow, fibrous, soft.
Sapwood yellowish. Twigs glabrous, grey or grey-brown. Leaves alternate, subopposite or
opposite, stalk slender, to 1(1.5) cm long; blades elliptic-oblong to elliptic, 48(10) x 25
cm, papery to thinly leathery, glabrous; base cuneate, margin not curled inward, apex
caudate with the tip to 1.5 cm long; lateral veins 46 pairs, prominent below, sunken above,
not looping toward leaf margin; intercostal veins faint to inconspicuous below.
Inflorescences to 6 cm long, subglabrous. Flowers 4-merous, green-yellow, buds to 1 mm
diameter; calyx-tube glabrous, with 4 lobes, broader than long, glabrous; petals 4, 1.11.3 x
0.70.9 mm, glabrous on the abaxial surface; stamens 4, filament 11.3 mm long; disc
yellowish. Fruits ovoid to oblong, 1.52.2 x 0.51 cm; fruit-wall thick; persistent disc not
well-exposed; persistent calyx-lobes inconspicuous. Seeds ellipsoid, 1.32 x 0.60.8 cm.
Vernacular name. Sarawakpatoli entelit (Iban).
Distribution. Sumatra, Peninsular Malaysia (new record), and Borneo. Widely distributed
in Sabah and the W and C parts of Sarawak.
Ecology. In primary mixed dipterocarp forest on clay-rich soils, from lowland to 1600 m.
Flowering in June to October, fruiting in August to March.
Taxonomy. In Matthew's treatment, Mastixia rostrata was segregated into two subspecies,
viz. rostrata and caudatifolia. Of these, only M. rostrata subsp. caudatifolia is known from
Sabah and Sarawak.
7. Mastixia trichotoma Blume

(Greek, tri = three, tomos = part; the 3-branched inflorescence)
l.c. (1825) 655, King l.c. 72; Wangerin l.c. (1910) 24; Merrill l.c. (1921) 459; Danser l.c. 57; Matthew
l.c. (1976) 68, l.c. (1977) 92; Kochummen l.c. 54; Anderson l.c. 164; Whitmore, Tantra & Sutisna l.c.
50. Type: Blume, s.n., West Java, Mt. Salak (lectotype L; isolectotypes BM, W). Synonyms: Mastixia
laxa Blume l.c. (1850) 257 (including var. angustifolia); Mastixia acuminatissima Blume l.c. (1850)
258; Mastixia caesia Blume l.c. (1850) 258; Mastixia kimanilla Blume l.c. (1850) 258.
Tree to 40 m tall and 50 cm diameter. Bark yellowish grey to grey-brown, smooth to

shallowly fissured; inner bark yellowish brown to pale brown. Sapwood pale white. Twigs
not distinctly grooved, yellowish brown to pale brown, puberulous to woolly. Leaves
opposite, stalk stout or slender, 13.5 cm; blades ovate, elliptic, lanceolate to oblong, 524 x
2.512 cm, thinly to thickly leathery, subglabrous to velvety hairy below; base acute,
cuneate, obtuse or attenuate, apex acute to acuminate; lateral veins 515 pairs, sunken
above, prominent below; intercostal veins reticulate, faintly visible to prominent below.
Inflorescences to 15 cm long, puberulous to woolly. Flowers 45-merous, green to
yellowish green; buds 12.5 mm diameter; calyx-tube puberulous to villous, with 4 or 5
lobes, as long as wide, puberulous to villous; petals 4 or 5, puberulous to villous on the
abaxial surface; stamens 4 or 5. Fruits ovoid to ellipsoid, 1.53 x 0.61.5 cm, fruit-wall
205
thin; persistent disc well-exposed and bulging or not; persistent calyx-lobes inconspicuous
to slightly prominent. Seeds ovoid to ellipsoid, 1.32.9 x 0.41.4 cm.
Key to varieties
1.
Inflorescence villous to woolly. Lateral veins looping to form marginal veins;

intercostal veins distinct and prominent below.............................................................2
Inflorescence subglabrous to puberulous. Lateral veins not looping; intercostal veins
not prominent below....................................................................................................3
2.
Twigs woolly. Leaves villous to woolly, thickly leathery. Flowers 4-merous...................

var. maingayi (Clarke) Danser
l.c. 63; Matthew l.c. (1976) 70, l.c. (1977) 93; Kochummen l.c. 54. Basionym: M. maingayi
Clarke l.c. 746. Type: Maingay 2680 = Kew Distr. 711, Singapore (holotype K; isotypes
BM, GH, L). Synonyms: M. maingayi Clarke var. subtomentosa King l.c. 75; M. propinqua
Ridl., J. Fed. Mal. St. Mus. 4,1 (1909) 25.
Twigs woolly by short yellowish brown hairs. Leaf-stalk stout, 23.5 cm, woolly
brown-hairy. Leaves 8.522 x 3.510.5 cm, thickly leathery, abaxial surface woolly
brown-hairy, adaxial surface glabrous except the midrib and veins; lateral veins 5
6 pairs, prominently sunken above and raised below, forming marginal loops
below; intercostal veins prominently sunken above and raised below.
Inflorescences villous to woolly. Flowers 4-merous. Fruits with well-exposed
persistent disc; persistent calyx-lobes 1.52 mm long.
Sumatra, Peninsular Malaysia, Borneo. In Sabah recorded from Lahad Datu,
Ranau, Sipitang, and Tawau districts. In Sarawak, recorded from only Kapit (3rd
Div.) and Ulu Lawas (5th Div.). In primary and secondary mixed dipterocarp
forests, to 1500 m.
Twigs subglabrous to velutinous. Leaves glabrous to subglabrous. Flowers 5-merous.....
var. korthalsiana (Wangerin) Danser
l.c. 63; Matthew l.c. (1976) 70, l.c. (1977) 93. Basionym: M. korthalsiana Wangerin l.c.
(1907) 335. Type: Korthals, s.n., E Borneo, Bandjermasin, G. Sakoembang (L).
Twigs subglabrous to velutinous. Leaf-stalks slender, 1.52 cm. Leaves 713 x 35

cm, glabrous to subglabrous; lateral veins 56 pairs, forming looped marginal veins
below; intercostal veins prominent below. Inflorescences velutinous to woolly.
Flowers 5-merous. Fruits with well-exposed persistent disc; persistent calyx-lobes
to 1 mm long.
Sumatra and Borneo (Sabah and Kalimantan). In Sabah, recorded from Kota Belud,
Lahad Datu, Sandakan, Sipitang, and Tawau districts. No record from Sarawak. In
primary mixed dipterocarp forest, from low altitude to 1100 m.
206
207
3. Leaves 924 x 412 cm, margin not wavy after drying, under surface not pale yellow
when dry. Twigs and leaves (sub)glabrous. Fruits with persistent disc well-exposed and
bulging..........................................................................................................................
var. rhynchocarpa Danser.
l.c. 64; Matthew l.c. (1976) 71, l.c. (1977) 93. Type: Endert 4769, E Borneo, East Coast,
near Long Hoet, 150 m (lectotype L; isolectotypes A, BO, K). Synonyms: Mastixia
trichotoma Blume var. benculuana Danser l.c. 64; Mastixia trichotoma Blume var.
simalurana Danser l.c. 65.

Twigs subglabrous. Leaf-stalks stout, 1.52 cm long. Leaves 924 x 412 cm,
thickly leathery, subglabrous, under surface not pale yellow when dry; lateral
veins 515 pairs, not looping near the margin; intercostal veins faintly visible
below. Inflorescences subglabrous to puberulous. Flowers 4-merous. Fruits
crowned by bulging, well-exposed persistent disc; persistent calyx-lobes to 0.5 mm
long.
Sumatra, Java, Borneo, Celebes, and the Moluccas. In Sabah widely distributed. In
Sarawak in Ulu Mujong (3rd Div.), Lawas (5th Div.), and Kapit (7th Div.). In
primary mixed dipterocarp forests, to 1800 m.
Leaves 7.515 x 2.56 cm, margin wavy after drying, under surface pale yellow when
dry. Twigs and leaves covered with powdery tomentum. Fruits not so............................
var. clarkeana (King) Danser
l.c. 62; Matthew l.c. (1976) 72, l.c. (1977) 93; Kochummen l.c. 54. Basionym: Mastixia
clarkeana King l.c. 72. Type: Scortechini 869, Perak (lectotype K; isolectotypes CAL, G, L,
P). Synonyms: Mastixia clarkeana King var. macrophylla King l.c. 72; Mastixia
korthalsiana Wangerin var. macrophylla Wangerin l.c. (1907) 336; Vitex premnoides
Elmer l.c. 2874; Mastixia premnoides (Elmer) Hallier f. l.c. 42; Mastixia trichotoma
Blume var. tenuis Danser l.c. 61.
Twigs covered with pale brown powdery tomentum. Leaf-stalks slender, 11.5 cm
long. Leaves elliptic, 7.515 x 2.56 cm, under surface pale yellow on drying;
lateral veins 57 pairs, not looping near the margin; intercostal veins faintly
visible below. Inflorescences subglabrous to puberulous. Flowers 4-merous. Fruits
without well-exposed persistent disc; persistent calyx-lobes prominent, 0.51 mm
long.
Pattani (Peninsular Thailand), Sumatra, Peninsular Malaysia, Borneo, and Mindanao
(Philippines). Uncommon in Borneo, recorded only from Kuching and Mt.
Penrissen (1st. Div.) in Sarawak. No record from Sabah. In primary mixed
dipterocarp forest, from low altitude to 1100 m.
Vernacular names. Sabahbantis (Dusun Kinabatangan), medang kanigara (Malay).
Sarawakitan beruang (Kelabit), medang kanigara (Malay), priabu (Murut).
Taxonomy. Of the five varieties recognised by Matthew l.c., four occur in Sabah and
Sarawak.
208
DATISCACEAE
E.J.F. Campbell-Gasis
c/o Forest Research Centre,
Sandakan, Malaysia
Merrill, EB (1921) 414; Masamune, EPB (1942) 507; van Steenis, FM I, 4 (1953) 382; Browne,
FTSB (1955) 82; Backer & Bakhuizen f., FJ 1 (1963) 313; Smythies, CST (1965) 43; Burgess, TS
(1966) 86; Whitmore, TFM 2 (1973) 29; Cockburn, TS 1 (1976) 73; Anderson, CLTS (1980) 165;
Ashton, MNDTS 2 (1988) 234; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 52.
Trees, shrubs or herbs. Leaves spiral, simple, without stipules; blade palmately veined,
domatia/glands present. Flowers in axillary and/or terminal elongated spikes or panicles,
unisexual, regular, parts not overlapping; male: sepals 49, free and unequal or connate in a
lobed tube; petals 49 or 0, free; stamens 49 on the sepals, filaments often long, anthers
basifixed, 2-celled, longitudinally dehiscent; female: sepals 38, often joined above the
ovary or free; petals 0; staminodes 0; ovary inferior, 1-celled with 38 parietal placentas
and numerous ovules; styles 38 opposite calyx-lobes, thick, free, mostly adnate to the
margin of the calyx, simple or bifid, stigma club-like or capitate. Fruits capsular, wall
papery, opening at apex with slits and/or splitting laterally. Seeds very numerous, minute;
endoperm none; embryo straight, cylindric.
Distribution. 3 genera, with 4 species: Datisca (2 species, extra-Malesian), Octomeles (1
species, Indomalesian) and Tetrameles (1 species, Indomalesian). Only Octomeles is found
Taxonomy. The systematic position of the Datiscaceae remains controversial. Several
authors (e.g., Whitmore l.c.; Mabberley, PB (1987) 172) maintain the Datiscaceae as most
closely related to the Begoniaceae and Cucurbitaceae, in the Cucurbitales. Mabberley l.c and
others (e.g., Willis, DFPF (1973) 1140) suggest that both Octomeles and Tetrameles should
be taken out of the Datiscaceae and put in the Tetramelaceae (Warb.) Airy-Shaw.
OCTOMELES Miq.
(Greek, octo = eight, melos = parts, limbs; the 8-merous flowers)
Fl. Ind. Bat., Suppl. (1861) 133, 336; van Steenis l.c. 382; Burgess l.c. 86; Cockburn l.c. 73; Anderson
l.c. 165; Wong, DMT (1984) 25; Ashton l.c. 234; Whitmore, Tantra & Sutisna l.c. 53.
Evergreen trees with thick twigs sharply 3-angular at apex. Leaves roundish cordate, 57(
9)-veined; stalk long, 5-angled; lower surface with domatial glands. Flowers sessile, green,
58-merous, dioecious, solitary; male: calyx-tube short with free triangular lobes; petals
209
triangular; stamens strongly incurved in bud, filaments thick, anthers large, kidney-shaped,
curved; female: sepals joined to form a tube; ovary apex cup-shaped by the thick calyx-tube;
styles fleshy, inserted in the calyx-rim opposite the free triangular calyx-lobes, stigma
capitate. Fruit a barrel-shaped capsule crowned with persistent styles, the styles and
exocarp drop away irregularly leaving a pale membranous endocarp behind; endocarp
splitting from the top downwards, the segments spreading stellately, persistent. Seeds
spindle-shaped, with narrower tail and thickened head.
Octomeles sumatrana Miq.
Fig. 1.
(of Sumatra)
l.c. 133, 336; van Steenis l.c. 382; Browne l.c. 82; Smythies l.c. 43; Cockburn l.c. 73; Anderson l.c.
165; Ashton l.c. 234; Whitmore, Tantra & Sutisna l.c. 53. Type: Teijsmann, s.n., Sumatra, Prov.
Palembang (holotype L; isotype K).
Tree to 70 m tall and 3 m diameter; bole columnar; buttresses steep, to 10 m high, spreading
to 5 m away from the tree. Branches radial, ascending, pagoda-like; crown monopodial.
Bark pale brownish cream or grey, thin, hooped (branch-scars), dot-like lenticels visible;
inner bark pinkish cream, soft, fibrous, thick. Sapwood cream, exudate slight, clear. Twigs
stout, glabrous, with prominent large round leaf-scars subtended by 5 almost papery sharp
ridges. Leaves broadly ovate to round, 1233 x 1629 cm, drying rather papery, with
scattered white hairs below; base heart-shaped, often deeply to 16 cm, margin toothed or
wavy, apex acute to acuminate; midrib raised above and below; lateral veins 710 pairs, the
basal 12 pairs originating from the stalk insertion, giving a somewhat 3-veined or
palmately veined appearance, mostly parallel and looping near margin, raised above and
below; intercostal veins net-like and flat; midrib and lateral veins dark red on both surfaces;
obscure dark domatia present at apex, at the junctions of the midrib and lateral veins and
alongside smaller veins (obvious in dry leaves); stalk glabrous, 35 mm thick and 632 cm
long, smooth on upper side, 5-ridged on lower side, grooved near base for 12 cm. Male
flowers in spikes 2060 cm long, c. (5)8 mm across and 7 mm long; sepals connate for
most of the length, lepidote, lobes free, triangular, 1.52 mm; petals triangular, alternate
with calyx-lobes, c. 3 mm long; stamens 410 mm long, anthers 23 mm. Female flowers
in shorter spikes 835 cm long, c. 5 mm across and 510 mm long; calyx-tube 24 mm
high; ovary 12 mm long, styles fleshy, triangular, 23 mm long, stigma thick, 12 mm.
Fruits 812 mm long, with pale brown endocarp. Seeds thicker in the middle, c. 0.8 mm
long, numerous.
Vernacular names. Sabahbinuang (Malay). Sarawakbenuang (Iban), binong (Bidayuh),
lemeng (Berawan, Punan, Tutoh).
Distribution. Sumatra, Borneo, Philippines, Celebes and New Guinea. In Sabah, common
on the east coast and interior. In Sarawak, in lowlands and hills in all districts.
Ecology. Often associated with riverine and alluvial deposits but also found in primary and
secondary mixed dipterocarp forests on well-drained fertile friable soils below 600 m on the
east coast of Sabah and to 800 m on the west coast and in Sarawak. It is more typically
210
211
found growing in disturbed habitats such as logged-over forest and after cultivation where it
can grow extremely fast, often in gregarious even-aged stands. Flowering and fruiting from
November to April in Sabah and Sarawak. Flowers probably wind-pollinated and the seeds
probably wind-dispersed.
Silviculture. Binuang is one of the fastest growing trees and a strong light-demander. It
takes only about 4 months from sowing to planting out in the field. Seed viability, however,
soon decreases after collection. Its ability to grow on low hilly sites and in temporarily
flooded areas, quick crown development, and self-pruning abilities, make it a suitable
plantation tree. For optimal growth, however, the seedlings/saplings would have to be
planted at sufficiently wide spacings to accommodate the huge crowns of the mature trees.
Timber strength & wood anatomy. Binuang timber is a soft, light hardwood of the obeche
(Triplochiton scleroxylon Sterculiaceae) type with long fibres and a low density. The
sapwood is almost white but of drab appearance and the heartwood is pale brown or pinkish
brown. The wood has an interlocked grain and the texture is coarse (Burgess l.c., Wong
l.c.). The pores are rather large, oval, mostly open, evenly distributed, with broader rays
distinct, almost white and the cut end surfaces are dull (Browne l.c.). The dry weight is 270
465 kg/m3 air-dried. The wood seasons slowly and is not durable. There is a tendency for
both brittle-heart and fungal stain to develop, and damage by pinhole-borer beetles and
powder-post beetles is sometimes present but treatment with preservatives gives moderate
resistance (Burgess l.c., Wong l.c.).
Uses. The timber works easily with machine and hand tools but the timber can crumble or
give a woolly finish if the tool cutters are not kept sharp. It also stains and polishes
satisfactorily. Burgess l.c. and Wong l.c. suggest that the wood can be used for cores, backs
to plywood, and is a useful utility timber for packing cases, concrete shuttering, matchboxes
and other temporary purposes. It is a potentially important tree for wood chip and pulping
and as a shade tree or for stabilising river banks but its potential is not yet exploited in
Sabah and Sarawak.
Notes. The seedlings and saplings have usually larger leaves with a cordate base and a few
coarse teeth, a distinctly opposite and decussate branching, and characteristically angled
twigs which make them easy to identify. Some young Macaranga species (Euphorbiaceae)
have similar leaves but these can be distinguished easily as the twigs of this genus do not
have papery ridges. Older trees can sometimes be confused with Neolamarckia cadamba
(Roxb.) Bosser (Rubiaceae), but Octomeles sumatrana has radial branching in distinct tiers
due to its episodic growth pattern whereas N. cadamba produces radial branches continuously
up the stem. The leaves of O. sumatrana are always dotted with holes due to insect damage
unlike the leaves of N. cadamba. It has been noted by van Steenis l.c. as being a preferred
tree for nesting bees like the other lofty tree species, Koompassia excelsa (Becc.) Taub.
(Leguminosae).
212
GOODENIACEAE
K.M. Wong
Forestry Department, Sabah
Merrill, EB (1921) 586; Masamune, EPB (1942) 724; Leenhouts, FM 1, 5 (1957) 335, FM 1, 5 (1958)
567, FM 1, 6 (1972) 949, FM 1, 7 (1976) 827, FM 1, 9 (1982) 566.
Small trees, shrubs or herbs, or scrambling-climbing plants. Leaves spirally arranged or

opposite, or whorled, simple, pinnately veined; leaf axils often with hair tufts; stipules none.
Inflorescences cymose, bracteate. Flowers 5-merous, bisexual, protandrous (male parts
maturing first); calyx tubular with distinct lobes; corolla members fused into one or two
broad lip-like structures in the open flower, the lobes with thin membranous margins;
stamens 5, free, borne on the calyx, filaments linear, anthers basifixed and introrse, 2-celled;
disc none; ovary 2-celled or imperfectly 1-celled, style cylindrical, stigma surround-ed by a
cup-shaped structure (indusium) with ciliate margin; ovules 1many, placentation axile or
basal. Fruits capsular or drupaceous. Seeds 1many, with endosperm.
Distribution. 16 genera, about 430 species, mostly Australian. In Malesia, 5 genera with 11
species. In Sabah and Sarawak, 5 species of Scaevola are documented.
Ecology. The family seems confined to or adapted to poor soils. In Malesia its species occur
in rather open sites in beach or coastal areas, or on ultramafic soils and in low-stature
montane forests.
Taxonomy. The Goodeniaceae are closely related to Lobelia and related genera in the
Campanulaceae.
SCAEVOLA L.
(the name given to Gajus Mucius around 507 B.C., who burnt his own hand in
fearlessness, to which is likened the white, 5-lobed corolla which turns brown)
Mant. (1771) 145; Merrill l.c. 586; Masamune l.c. 724; Leenhouts l.c. (1957) 339, l.c. (1958) 567, l.c.
(1972) 951. Synonym: Temminckia de Vriese, Ned. Kruidk. Arch. 1, 2 (1851) 141.
Shrubs or small trees, or scrambling-climbing plants. Leaves spirally arranged or opposite,

or whorled, entire or crenate to dentate, the margins and tip provided with glands; leafstalks with broad insertion on the shoots. Flowers in axillary, bracteate cymes; calyx-tube
fused to the ovary; corolla members all fused to form a broad lip-like structure in the open
flower, the free lobes with membranous margins and fimbriate at their base; ovary inferior
213
or semi-inferior, 12-celled; ovules 1 per ovary cell. Fruits drupaceous, with a hard stone.
Seeds 1 or 2.
Distribution. About 130 species, predominantly Australian, one species reaching to Taiwan
and Hainan in south China, a few in east Malesia (New Guinea), and two widely distributed, chiefly littoral species (S. sericea in the Indo-Pacific region, S. plumieri in the IndoAtlantic region). In Sabah and Sarawak, 5 species.
Key to Scaevola species

1.
Leaves in whorls of four.......................................................................5. S. verticillata

Leaves spirally arranged..............................................................................................2
2.
Leaves sessile to subsessile, the blades decurrent more or less all the way to the nodes
or the stalks exceedingly short and inconspicuous........................................................3
Leaves distinctly stalked, decurrent basal part of the blades very narrow and inconspicuous...........................................................................................................................4
3.
Plants of sandy, rocky or cliff sites beside the sea or the coast. Leaves exceeding 10 cm
long, with long-decurrent bases. Cymes (2)35(6)-times branched, often 610 cm
long. Calyx-lobes linear to lanceolate, 23 mm long (sometimes elongating to 57 mm
long); corolla with a fused part 1520 mm long and free lobes 510 mm long. Fruits
broad-obovoid to globose, 812 mm long, strongly ridged..........................4. S. sericea
Plants of high elevation on mountains (known only on Mt Kinabalu, Sabah). Leaves 1
7 cm long, more or less abruptly sessile. Cymes 12-times branched, not exceeding 3
cm long. Calyx-lobes triangular, less than 1 mm long; corolla with a fused part 46 mm
long and free lobes 46 mm long. Fruits narrow-obovoid, 3.55 mm long, slightly
ridged .........................................................................................................1. S. chanii
4.
Plants of ultramafic substrates (in Borneo known only in Sabah). Leaves 815 cm long,
distinctly crenate-dentate on the margins. Cymes 36-times branched, 59 cm long, the
first pair of bracts conspicuously larger and leaf-like, peduncle 23.5 cm long. Flowers
in the forks of cyme branches, stalked......................................2. S. micrantha
Plants so far known only on limestone (in Sarawak). Leaves not exceeding 8 cm long,
only minutely and inconspicuously crenate on the margins. Cymes 12-times branched, not exceeding 2.5 cm long, the first pair of bracts tiny triangular structures,
peduncle to 1 cm long only. Flowers in the forks of cyme branches, sessile................
..............................................................................................................3. S. muluensis
1. Scaevola chanii Wong
Fig. 1.
(C.L. Chan, orchidologist and botanical illustrator)

Sandakania 3 (1993) 12. Type: Chan, Jamal & Wong WKM 2360, Sabah, Mt. Kinabalu, summit trail,
2900 m alt. (holotype SAN; isotypes K, KEP, L, SAR, SNP).
Shrub or treelet, to 23 m high. Leaves spirally arranged, obovate, 17 x 0.63 cm, margin
minutely crenate-dentate, coriaceous; glabrous except for small tufts of silvery white hairs
214
215
in the leaf axils, and tiny silvery round scales all over the young leaves; lateral veins 48
pairs, inconspicuous; stalks very short, only 12 mm long at most, inconspicuous. Inflorescences 1.52.5 cm long, branched only 12-times, peduncle 0.72 cm long; first pair of
bracts triangular-linear, small; glabrous except for tufts of pale hairs in the bract axils.
Flowers sessile; calyx cup obovoid, glabrous, the lobes triangular, less than 1 mm long, pale
pilose on the margins; corolla sparsely pale hairy on the outside, the fused part 46 mm
long, the free lobes 46 mm long, pale pilose inside; stamens with filaments c. 3 mm long,
anthers 0.71 mm long; style 68 mm long; indusium around the stigma glabrous on the
surface, densely pale hairy on the margin. Fruits obovoid, 3.55 mm long, slightly ridged.
Distribution. Recorded only from Mt. Kinabalu, Sabah. So far endemic.
Ecology. Upper montane forest on ultramafic soil, at 25003000 m.
2. Scaevola micrantha C. Presl

(Greek, micros = small, anthos = flower)
Rel. Haenk. 12 (1835) 58; Leenhouts l.c. (1957) 342. Type: Haenke 124, Philippines (W). Synonyms:
Temminckia micrantha (C. Presl) de Vriese l.c. (1851) 145; S. pedunculata Merr., Philip. J. Sc., Bot. 5
(1910) 251, incl. var. mollis, l.c. (1921) 586, Masamune l c. 724; S. merrillii Elmer, Leafl. Philip. Bot.
4 (1912) 1491.
Shrub or small tree to 10 m tall, 5 cm diameter. Bark dark grey to greenish grey, smooth to
slightly flaky. Leaves spirally arranged, obovate to oblanceolate, 815 x 25.5 cm, margin
crenate to distinctly dentate, chartaceous to thinly coriaceous; subglabrous (with scattered
hairs on the stalk, leaf base and midrib on the lower side) to scantily short-hairy to velvety
on the lower side, the leaf axils with tufts of silvery white hairs; lateral veins 611 pairs,
inconspicuous; stalks 417 mm long, distinct. Inflorescences 59 cm long, branched 36times, peduncle 23.5 cm long; first pair of bracts ovate and leaf-like, much larger than
subsequent bracts; glabrous to pale velvety all over. Flowers on stalks 0.51.5 mm long;
calyx-cup obovoid, glabrous to densely pale pilose, the lobes triangular, c. 0.5 mm long,
short-hairy on the margins to pale pilose all over; corolla scantily to densely pale pilose all
over the outside, the fused part 57 mm long, the free lobes 45 mm long, pale pilose
inside; stamens with filaments c. 4 mm long, anthers c. 1.5 mm long; style 67 mm long;
indusium around the stigma scantily hairy to pale pilose on the surface, densely pale-hairy
on the margin. Fruits obovoid, 56 mm long, slightly ridged.
Distribution. Borneo and the Philippines, including Palawan Is. In Borneo, known only
from Sabah (Kinabalu-Ranau area, Telupid, Beluran, Mt. Tawai and Mt. Danum). The
Celebes collection (Lam 3266) from Talaud Is. attributed to this species by Leenhouts l.c.
(1957) appears to be a different species; it has entirely glabrous leaves, 1-mm-long calyx
teeth, linear and longer (68 mm) inflorescence bracts, and only 23-times branched
inflorescences.
Ecology. Recorded only from ultramafic substrates, from the lowlands to submontane forest
at c. 1500 m.
216
GOODENIACEAE (WONG)
3. Scaevola muluensis Wong

(of Mt. Mulu, Sarawak)
Sandakania 3 (1993) 15. Type: Argent & Jermy 1011, Sarawak, 4th Division, Gunong Api (holotype
SAN; isotypes E, L, SAR).
Shrub or treelet to c. 1 m high. Leaves spirally arranged, obovate, 46.5 x 23 cm; margin
only minutely and inconspicuously crenate; thinly coriaceous; glabrous except for scattered
short pale hairs on the stalk and tufts of short pale hairs in the leaf axils; lateral veins 56
pairs, slightly elevated on both surfaces when dry; stalks 513 mm long, distinct.
Inflorescences 12 cm long, branched only 12-times, peduncle to 1 cm long only; first
pair of bracts tiny triangular structures; glabrous except for scanty pale hairs in the bract
axils. Flowers sessile; calyx-cup obovoid, glabrous, the lobes triangular, c. 0.5 mm long,
glabrous to sparsely short-hairy on the margins; corolla sparsely pale-hairy on the outside,
the fused part 66.5 mm long, the free lobes 4.55 mm long, pale pilose inside; stamens
with filaments c. 4 mm long, anthers c. 1 mm long; style 66.5 mm long; indusium around
the stigma with pale long hairs all over, densely pale hairy on the margin. Fruits obovoid,
45 mm long, slightly ridged, ripening black.
Distribution. Known from two collections (S.30907 and Argent & Jermy 1011) between
1000 m and 1600 m on the limestone of Gunong (Mt.) Api in the Gunong Mulu National
Park in Sarawak's 4th Division.
Ecology. Probably restricted to limestone, found on exposed ridges and cliff faces.
4. Scaevola sericea Vahl

(Latin, sericeus = silky with long hairs; the leaves)
Symb. Bot. 2 (1791) 37; Leenhouts, l.c. (1957) 339. Type: G. Forster, s.n., Pacific, Niue Is. (C).
Synonyms: Lobelia plumieri (non L.) Burm. f., Fl. Ind. (1768) 189; S. koenigii Vahl, Symb. Bot. 3
(1794) 36; S. taccada Roxb., Hort. Beng. (1814) 15, Leenhouts l.c. (1972) 951; S. plumieri (non Vahl)
Blume, Bijdr. (1826) 730; S. leschenaultii DC., Prod. 7 (1839) 506; S. macrocalyx de Vriese l.c.
(1850) 138; S. piliplena Miq., Fl. Ind. Bat. 2 (1857) 581; S. frutescens (non Lobelia frutescens Mill.)
Krause, Pfl. R. Heft 54 (1912) 125, Merrill l.c. (1921) 586, Masamune l.c. 724.
Shrub or small tree to 10 m tall and 10 cm diameter. Bark yellowish green to grey, smooth.
Leaves spirally arranged, obovate, 1226 x 510 cm; base decurrent to the node, margin
crenate to subentire, apex obtuse, rounded; thickly succulent to thinly coriaceous; glabrous
to scantily hairy to tomentose on both sides, the leaf axils with dense tufts of pale longhairs; lateral veins 812 pairs, slightly elevated on upper side; stalks nil or very short,
hardly 12 mm long. Inflorescences (4)610 cm long, branched (2)35(6)-times,
peduncle c. 1 cm long; first pair of bracts linear-triangular; glabrous to densely short-hairy
all over. Flowers sessile on the slender cyme branches; calyx cup obovoid-ellipsoid,
glabrous to scantily to densely pale hairy, the lobes linear to lanceolate, 23 mm, elongating to
57 mm long, scantily to densely short-hairy all over; corolla scantily to densely pale-hairy
all over the outside, the fused part 1520 mm long, the free lobes 58(10) mm long, pale
pilose inside; stamens with filaments 1015 mm long, anthers 23 mm long; style 1520
mm long, indusium around the stigma scantily to densely hairy all over, densely pale-hairy
on the margin. Fruits broadly obovoid to subglobose, 812 x 1015 mm, conspicuously
ribbed, fleshy, stone to 8 mm across.
217
Vernacular name. Sarawakbutun laut (Lundu Malay).

Distribution. Madagascar, SE Asia, throughout Malesia, tropical Australia, Micronesia,
Melanesia, to Hawaii. In Borneo common along the sea and coast in Sabah, Sarawak,
Ecology. Common on sea-shores and sandy sites behind the shore, on rocky cliffs and even
some sandstone hills on the coast (e.g., Trig Hill in Sandakan). Flowers and fruits all year
round. The stones, with a corky outer layer, are buoyant and adapted to dispersal by sea
currents. The seeds remain viable after long periods in sea water, but will germinate only
with fresh water as after being washed ashore on a rainy day (Lesko & Walker, Ecology 50
(1969) 730).
Uses. The pith ("taccada") is used for making Malayan rice-paper and crafting artificial
objects of decoration, such as flowers (Burkill l.c.). Mamit (S. 35139, Sarawak, 1st Division,
Lundu) records that the liquid extracted from the fruit can be applied directly to cure sore
eyes.
5. Scaevola verticillata Leenh.

(Latin, verticillatus = whorled; the leaves)
Blumea 12 (1964) 317, f. 1, l.c. (1972) 951. Type: Meijer SAN 22818, Sabah, Ranau, Mt.
Tambuyokon (holotype L; isotypes K, SAN, SAR).
Shrub or treelet. Leaves in whorls of four, obovate, 25.5 x 0.52 cm; margin minutely
crenate and recurved strongly; thickly succulent to coriaceous; densely tomentose but later
glabrescent on upper side, densely woolly tomentose on lower side and stalk, leaf axils with
tufts of pale long-hair; lateral veins 812 pairs, inconspicuous to slightly elevated on upper
side; stalks very short, 36 mm long. Inflorescences 1.52 cm long, branched only 24times, peduncle to 0.71.2 cm long only, the branches very short; first pair of bracts slightly
larger and broader than subsequent bracts; densely hairy all over. Flowers sessile; calyx-cup
obovoid, scantily to densely short-hairy all over, the lobes of different sizes and shapes (one
ovate and c. 2.5 mm long, one triangular and c. 1 mm long, three rounded and c. 0.5 mm
long), scantily short-hairy on the surface, densely long-hairy on the margin; corolla densely
woolly tomentose on the outside, the fused part 56 mm long, the free lobes 56 mm long,
pale pilose inside; stamens with filaments 4.55 mm long, anthers c. 1 mm long; style 6.57
mm long; indusium around the stigma with pale long hairs around the base, densely palehairy on the margin. Fruits obovoid-ellipsoid, 33.5 mm long, slightly ridged, glabrescent
except for the hairy persistent calyx-lobes.
Distribution. Recorded only from Mt. Tambuyukon near Mt. Kinabalu, Sabah. Probably
endemic.
Ecology. Found in stunted subalpine vegetation at around 2500 m, on ultramafic substrate.
218
HYPERICACEAE
K.M. Wong
Sandakan, Malaysia
Merrill, EB (1921) 392 (as part of Guttiferae); Masamune, EPB (1942) 477, 482 (as part of
Guttiferae); Browne, FTSB (1955) 195; Smythies, CST (1965) 68; Burgess, TBS (1966) 311;
Kochummen, TFM 2 (1973) 248; Robson, FM 1, 8 (1974) 1; Shea, TS 1 (1976) 142; Anderson, CLTS
(1980) 217; Corner, WSTM 1 (1988) 364; Ashton, MNDTS 2 (1988) 295; Whitmore, Tantra &
Sutisna, CLK 1 (1989) 181.
Trees, shrubs or herbs. Leaves simple, opposite, entire, without stipules, often with translucent or dark glandular dots. Inflorescences terminal and sometimes also axillary, rarely
axillary only, cymose, thyrsoid or paniculate, 1many-flowered. Flowers bisexual, radially
symmetrical, in some species with short-styled and long-styled forms; sepals 5, free or
partially fused, imbricate, often with linear glands, persistent in the fruit; petals 5, free,
imbricate, sometimes with scale-like nectar-bearing appendages at the base, glabrous;
stamens 5many, fused and grouped into 35 fascicles, anthers dorsifixed, staminodal
fascicles sometimes present; ovary superior, 35-celled or just 1-celled, styles (2)35, free
or partially fused; ovules 2many; placentas 25, parietal. Fruits capsular, splitting septicidally or loculicidally. Seeds 1many, unwinged or winged; endosperm none.
Distribution. Seven genera and about 550 species, cosmopolitan except for the Arctic
regions. In Malesia, including Borneo, only two genera (Cratoxylum and Hypericum) occur.
In Sabah and Sarawak, 6 species of Cratoxylum (shrubs and trees) and 2 species of
Hypericum (herbs) are found.
Ecology. In Sabah and Sarawak, species of Cratoxylum are mainly lowland plants; a few
species occur also in lower montane areas. In general these are all fast-growing trees
commonly found in forest fringes, gaps and disturbed habitats. The light seeds are winddispersed. Hypericum petiolulatum is a widespread species which in Borneo occurs only on
Mount Kinabalu. H. japonicum occurs in lowland marshy places and rice fields.
Uses. Cratoxylum timber is distinguished into two types, one heavier and classified as
derum in the trade, the other lighter and known as geronggang. C. cochinchinense, C.
formosum and C. maingayi produce derum timber, which is moderately hard and moderately
heavy to heavy; the timber is sometimes known as serungan batu (Sabah) or entemu
(Sarawak). C. arborescens and C. glaucum produce geronggang timber, a light hardwood
sometimes called serungan in Sabah. Derum is suitable for heavy and medium construction
under cover, for tool handles and for wooden pallets. Geronggang has been found suitable
for veneers and plywood, flooring, interior works and light to medium construction (Wong
219
(1982), DMT). Geronggang logs float in water. Browne l.c. and Burgess l.c. review the
wood characteristics.
Taxonomy. Three tribes are recognised within the family: Vismieae (3 genera, Africa and
America), Cratoxyleae (3 genera, Madagascar, Indo-Malesia, E Asia, NE America), and
Hypericeae (Hypericum, mostly montane tropical and temperate areas). Phytochemically,
the Hypericaceae are closely related to the Guttiferae; there are also many basic morphological
and anatomical similarities between the two families.
Key to genera
Herbs. Bark without resinous sap. Flowers yellow to orange; petals without scale-like
appendages. Fruits septicidally dehiscent (splitting along the internal partitions). Seeds
without wings.......................................................................................................................
Hypericum L.
Gen. Pl. ed. 5 (1754) 341; Robson l.c. 14.
About 400 species. Most tropical and temperate regions, but notably absent from the
Amazon basin and uncommon in Australasia.
Small trees, shrubs or herbs; in Borneo the two species are herbs. H. petiolulatum
Hook. f. & Thoms. ex Dyer occurs from Nepal through Indo-China to Sumatra and
Borneo; in Borneo it is known only from Mount Kinabalu. H. japonicum has
naturalised in marshy places and rice fields in Borneo; it is distributed from Japan and
Korea through China, Sri Lanka to Malesia, Australasia and Hawaii.
Shrubs or trees. Bark exuding yellow resinous sap which dries black. Flowers white to
reddish pink; petals sometimes with basal scale-like appendages. Fruits loculicidally
dehiscent (splitting between the internal partitions). Seeds winged....................Cratoxylum
CRATOXYLUM Blume
(Greek, kratos = strong, xylon = wood; referring to the hard and durable timber)
derum, geronggang (Malay)
Verh. Bat. Gen. 9 (1823) 174; Merrill l.c. 392; Masamune l.c. 477; Browne l.c. 195; Smythies l.c. 68;
Burgess l.c. 311; Gogelein, Blumea 15 (1967) 453; Kochummen l.c. 248; Robson l.c. 4; Shea l.c. 144;
Anderson l.c. 217; Corner l.c. 364; Ashton l.c. 295; Whitmore, Tantra & Sutisna l.c. 181.
Shrubs or small to medium-sized trees. Bark exuding a yellow resinous sap which dries
black. Leaves with translucent glandular dots. Flowers basically 5-merous; sepals 5,
coriaceous, with longitudinal glandular lines or dots, persistent and often elongating in fruit;
petals 5, white or pink to crimson, with longitudinal glandular lines or dots, in some species
with an inner basal nectariferous scale-like appendage; stamen fascicles 3, the two larger
attached to sepals, the smaller third attached to a petal, each with many stamens, filaments
white to crimson, anthers white to crimson; (sterile) staminodal fascicles 3, short and
alternating with the stamen fascicles; ovary incompletely 3-celled; styles 3, free, linear;
220
HYPERICACEAE (WONG)
stigmas small, knob-like; ovules 4many, attached to the basal part of the placenta,
ascending. Fruits ellipsoid, coriaceous to woody, 3-valved, splitting loculicidally. Seeds
4many on each placenta, linear to ovoid, winged all round or only on one side.
Distribution. 6 species, from India through S China to Malesia (absent in the Moluccas and
New Guinea); all present in Borneo.
Ecology. Rare in primary forest, and more common in gaps, forest fringes and disturbed
habitats in the lowlands. C. arborescens and C. formosum can occur on well-drained as well
as swampy substrates. C. glaucum is found mostly in peat swamps, freshwater swamps, and
wet heath forests on white sands. C. maingayi is restricted to limestone. The species are
deciduous or semi-deciduous except for the evergreen C. arborescens and C. glaucum. C.
formosum and C. maingayi flower on the bare branches; the other species bear
inflorescences with the expanded new foliage or on mature leafy branches.
Key to Cratoxylum species

1.
Leaf lateral veins inconspicuous, rather straight from midrib to very near the margin
where a smooth-running marginal vein occurs. Seeds with wing all round (section
Isopterygium)..............................................................................................................2
Leaf lateral veins distinct and prominent at least on lower leaf surface, arching at least
23 mm away from the margin, not joining at all, or joining to form conspicuous veinloops. Seeds with wing on one side only.......................................................................3
2.
Leaves not glaucous beneath, apex with at least a short tip, acuminate or caudate; leafstalks at least 4 mm long. Seeds 1018 in each fruit cell....................1. C. arborescens
Leaves glaucous beneath, apex rounded, blunt or notched; leaf-stalks 13 mm long.
Seeds 48 per fruit cell............................................................................4. C. glaucum
3.
Leaf lateral veins not or only weakly joining to form vein-loops. Inflorescences developing
on leafy branches, either terminal panicles or short cymes of a few flowers each at the
ends of shoots or in leaf axils. Petals crimson to dark red or orange, without nectary
scale (section Cratoxylum)...........................................................................................4
Leaf lateral veins joining strongly to form conspicuous vein-loops towards the margin.
Inflorescences developing on leafless parts of branches, of single axillary flowers or on
short axillary shoots and so appearing clustered. Petals white to pale lilac, with a basal
nectary-bearing scale (section Tridesmos)....................................................................5
4.
Lateral veins on lower leaf-surface distinctly raised and rather coarse. Inflorescence a
terminal panicle of many flowers. Persistent sepals less than half the capsule length
.........................................................................................................6. C. sumatranum
Lateral veins on lower leaf-surface slightly raised and rather fine. Inflorescence a short
1few-flowered cyme, terminal or axillary. Persistent sepals more than half the capsule
length...........................................................................................2. C. cochinchinense
221
222
HYPERICACEAE (WONG)
5. Mature leaves typically at least 6 cm long, lateral veins more than 7 pairs, stalks 610
mm long. Persistent sepals to only a third the capsule length................3. C. formosum
Mature leaves typically not exceeding 4.5 cm long, lateral veins only 56 pairs, stalks
to only 5 mm long. Persistent sepals nearly half the capsule length........5. C. maingayi
1. Cratoxylum arborescens (Vahl) Blume

(Latin, arbor = tree; the habit)
Mus. Bot. Lugd. Bat. 2 (1852) 17; Merrill l.c. 392; Masamune l.c. 477; Browne l.c. 195; Smythies l.c.
68; Kochummen l.c. 249; Shea l.c. 144; Anderson l.c. 217; Corner l.c. 365; Ashton l.c. 296;
Whitmore, Tantra & Sutisna l.c. 181. Basionym: Hypericum arborescens Vahl, Symb. Bot. 2 (1791)
86, t. 43. Type: Konig, s.n., 1778, Malacca (C).
Tree, sometimes a shrub, to 60 m tall and 120 cm diameter; buttresses if present to 1 m high.
Bark grey to dark brown or reddish brown, smooth to fissured-cracking and papery scaly;
inner bark pink to pale brown, finely laminated. Sapwood pale yellow. Leaves elliptic to
obovate, 514 x 26.5 cm, coriaceous; base cuneate, apex cuspidate to slightly caudate;
midrib flat to sunken on upper side; lateral veins 3050 pairs, fine, indistinct, running
straight to margin where there is a clear marginal vein; stalk 610 mm. Flowers arranged
in a panicle; petals pink to crimson, with punctate glands, basal scale small. Fruits 79 x
34 mm, persistent sepals to half or more than half the capsule length. Seeds 1018 per
locule, each with a wing all round.
Vernacular names. Sabahgeronggang, serungan (general). Sarawakdat (Kayan, Punan
Tutoh), di'it (Melanau Oya), geronggang (Bidayuh), idat (Melanau Oya), kata mudung
(Kenyah), labakan (Kelabit), manat (Melanau Rejang), mertilan (Iban), serungan labakan
(Murut, Kenyah), tat (Kenyah). Bruneigeroking (Belait), geronggang (Belait, Dusun,
Iban), labakan (Murut), madak (Tutong). Browne l.c. coins the name geronggang gajah.
Distribution. S Burma, Sumatra, Peninsular Malaysia, Borneo. In Sabah and Sarawak in all
districts. Also in Brunei and Kalimantan.
Ecology. Lowlands principally, also up to lower montane forest at 1400 m. Primary and
secondary forests, including kerangas (heath) and peat swamp forest, sometimes gregarious,
and may form thickets in exploited forest areas.
Uses. The species provides much of the geronggang timber in Sabah and Sarawak.
2. Cratoxylum cochinchinense (Lour.) Blume
Fig. 1.
(of Indo-China)
l.c. (1852) 17; Masamune l.c. 477; Shea l.c. 145; Kochummen l.c. 251; Anderson l.c. 217; Corner l.c.
365; Ashton l.c. 299; Whitmore, Tantra & Sutisna l.c. 181. Basionym: Hypericum cochinchinense
Lour., Fl. Cochin. (1790) 472. Type: Loureiro, s.n., "in sylvis Cochinchinae" (BM). Synonyms: C.
polyanthum Korth., Verh. Nat. Gesch. Bot. (1842) 175, t. 36; C. myrtifolium Blume l.c. (1852) 17; C.
ligustrinum (Spach) Blume l.c. (1852) 16.
223
Tree, sometimes a shrub, to 30 m tall and 65 cm diameter; trunk sometimes with small
buttresses and sometimes spiny. Bark pale brown to reddish brown, smooth to flaky or
papery; inner bark greenish yellow to pink, thin. Sapwood white to yellowish. Leaves
elliptic, 3.511 x 14 cm, glaucous beneath, chartaceous to thinly coriaceous; base cuneate,
apex acute; midrib sunken on upper side; lateral veins 912 pairs, fine, distinct, not joining
to form a marginal vein; stalks 23(10) mm. Flowers 1several in short axillary cymes;
petals crimson, with linear glands, without basal scale. Fruits 812 x 45 mm, persistent
sepals more than half the capsule length. Seeds 58 per locule, each with a wing only on
one side.
Vernacular names. Sabahselangan biabas (Suluk). Bruneimachit laling (Belait),
mertilan (Iban), serungan (Brunei Malay), taikakang (Kedayan).
Distribution. Burma, Indo-China, South China, Sumatra, Peninsular Malaysia, Borneo, and
the Philippines. In Sabah and Sarawak in all districts. Also in Brunei and Kalimantan.
Ecology. Lowland and hill forest, in primary or secondary forest including kerangas (heath)
forest.
3. Cratoxylum formosum (Jack) Dyer

(Latin, formosus = beautiful; the flowers)
in Hooker f., Fl. Br. Ind. 1 (1874) 258; Merrill l.c. 392; Masamune l.c. 477; Shea l.c. 146; Browne l.c.
197; Anderson l.c. 217; Corner l.c. 365, Pl. 80; Kochummen l.c. 251; Ashton l.c. 299; Whitmore,
Tantra & Sutisna l.c. 181. Basionym: Elodea formosa Jack, Mal. Misc. 2 (1822) 24. Type: Jack, s.n.,
Sumatra (K).
Tree, sometimes a shrub, to 20 m tall and 30 cm diameter, trunk sometimes spiny. Bark
grey to red-brown or black, smooth to fissured or scaly; inner bark yellowish. Sapwood pale
yellowish. Leaves broadly elliptic, 415 x 2.58 cm, glaucous beneath, chartaceous to
coriaceous; base cuneate, apex rounded to acute or shortly tipped; midrib sunken on upper
side; lateral veins 816 pairs, coarse, raised, distinct, arching and forming vein-loops
towards the margin; stalks 510 mm long. Flowers single, axillary, opposite; petals white to
pale lilac, with punctate glands, basal scale large. Fruits 1016 x 46 mm, persistent
sepals to only a third of the capsule length. Seeds 717 per locule, each with a wing on one
side only.
Vernacular names. Sabahgeronggang, serungan (generally). Sarawakdat tetong (Punan
Tutoh), entemu (Iban), kajo jelan (Berawan), melan (Kenyah), mirinos (Bidayoh Sadok),
nyalin bahe (Kayan), patok tilan (Iban), raja tugag (Nidayoh Padawan), sidodot (Bidayoh
Bau).
Distribution. Indo-China, S Andaman Is., Sumatra, Peninsular Malaysia, Java, Borneo,
Philippines, Celebes. In Sabah and Sarawak in all districts. Also in Brunei and Kalimantan.
Ecology. Principally lowlands, in primary or secondary forests, including peat swamps.
Recorded on sea shores, kerangas (heath forest) on white sand, sandstone, alluvium,
limestone and ultramafic soils. Trees of this species can be fully deciduous and the
inflorescences develop on bare parts of branches before and as a new flush of leaves
develops. Ashton (BRUN 5554) noted this species as flowering gregariously in riverbank
forest in Brunei.
224
HYPERICACEAE (WONG)
4. Cratoxylum glaucum Korth.

(Latin, glaucus = pale blue-green; the lower leaf surface)
l.c. 176; Merrill l.c. 392; Masamune l.c. 477; Anderson l.c. 217; Ashton l.c. 298; Whitmore, Tantra &
Sutisna l.c. 181. Type: Muller L 904, Karrau, Doeson (lectotype BM; isolectotype W). Synonym: C.
procerum Diels, Bot. Jahrb. (1926) 311.
Small tree or shrub, to 10 m tall, rarely to 25 m tall and 45 cm diameter. Bark reddish
brown, flaky. Leaves broadly elliptic, 25 x 1.53 cm, glaucous beneath, coriaceous; base
cuneate, apex rounded, blunt or notched; midrib flat to sunken on upper side; lateral veins
1014 pairs, fine, indistinct, running straight toward the margin where there is a clear
marginal vein; stalks 13 mm long. Flowers in a panicle or thyrse; petals crimson, with
punctate glands, basal scale small. Fruits 710 x 34 mm, persistent sepals to half the
capsule length. Seeds 48 per locule, each with a wing all round.
Vernacular names. Sarawakgeronggang (Iban), geronggang lompong (Iban), kirap
(Bidayoh), pidang (Kenyah), timau (Iban). Bruneigeronggang timau (Iban). Browne l.c.
coins the name geronggang puteh.
Distribution. Sumatra (Lingga, Bangka and Billiton), Peninsular Malaysia (Johore),
Karimata and Natuna Islands, Borneo (W and SE). In Sarawak in coastal areas throughout
the state. Also in Brunei and Kalimantan.
Ecology. Found in lowland primary and secondary kerangas (heath) forest on white sand, in
peat swamps (especially those dominated by the dipterocarps Dryobalanops rappa and
Shorea albida), and also freshwater swamps. Also documented at 1100 m in mossy
kerangas forest on sandstone ridges.
5. Cratoxylum maingayi Dyer

(A.C. Maingay, 18361869, botanist of the East India Company)
l.c. 258; Browne l.c. 197; Anderson l.c. 217; Corner l.c. 367; Kochummen l.c. 251; Ashton l.c. 300;
Whitmore, Tantra & Sutisna l.c. 181. Type: Maingay, s.n., Penang (holotype BM; isotype L).
Synonym: C. cochinchinense var. calcareum Ridl., Kew Bull. (1938) 115.
Tree or shrub, to 10 m tall and 10 cm diameter, often crooked. Bark brown, smooth to
fissured or flaky. Leaves broadly elliptic, 1.54.5(5.5) x 0.83 cm, coriaceous; base
cuneate, apex rounded, notched or obtuse with a short tip; midrib flat to sunken on upper
side; lateral veins 56 pairs, raised, distinct, arching and forming vein-loops towards the
margin; stalks to 5 mm long. Flowers single, axillary, opposite; petals pale pink, with
punctate glands, basal scale large. Fruits 615 x 35.5 mm, persistent sepals nearly half
the capsule length. Seeds 56 per locule, each with a wing on one side only.
Vernacular names. Sarawakgerunggang (Iban), patok tilan (Iban).
225
Distribution. Indo-China, Sumatra, Peninsular Malaysia, Borneo. In Sarawak, documented

from the 1st Div. near Kuching only; also reported in Kalimantan.
Ecology. Lowlands, known only from limestone substrates. Trees of this species are often
bare of leaves for a short time, when the inflorescences develop on bare parts of branches.
6. Cratoxylum sumatranum (Jack) Blume

(of Sumatra)
l.c. (1852) 16; Kochummen l.c. 251; Shea l.c. 146; Anderson l.c. 218; Ashton l.c. 301; Whitmore,
Tantra & Sutisna l.c. 181. Basionym: Elodea sumatrana Jack l.c. 22. Type: Jack, s.n., Sumatra, Tello
Dalam (K). Synonyms: C. hypericum (Blume) Merr. l.c. 392; C. celebicum Blume l.c. (1852) 16.
Tree or shrub, to 35 m tall and 60 cm diameter, trunk sometimes with buttresses to 1 m high.
Bark grey to pale or dark brown, cracking-fissured to scaly; inner bark pale yellow to
reddish, soft. Sapwood pale yellow to white. Leaves ovate-elliptic, 4.517 x 1.54.5 cm,
chartaceous to thinly coriaceous; base cuneate, apex acute; midrib flat to sunken on upper
side; lateral veins 922 pairs, coarse, raised, distinct, not joining to form marginal vein;
stalks 23 mm long. Flowers in a panicle; petals crimson, with linear glands, basal scale
absent. Fruits 710 x 35 mm, persistent sepals less than half the capsule length. Seed
310 per locule, each with a wing on one side only.
Vernacular names. Sabahserungan (Malay). Bruneiserungan mampat (Malay).
Distribution. Southeast Asia, Sumatra, Java, Borneo, Philippines, Celebes, Lesser Sunda Is.
In Sabah and Sarawak in all districts. Also in Brunei and Kalimantan.
Ecology. Lowlands, in primary and secondary forest, documented on sandstone, white sand
in kerangas (heath) forest, and also on ultramafic soils. Rarely found up to 1300 m.
Taxonomy. The Bornean taxon is subsp. sumatranum; subsp. blancoi (Blume) Gogelein is
restricted to the Philippines.
226
ILLICIACEAE
Richard M.K. Saunders
Department of Ecology & Biodiversity,
University of Hong Kong
A.C. Smith, Sargentia 7 (1947) 8; Hutchinson, Fam. Fl. Pl. 1, 2nd ed. (1959) 125; Gen. Fl. Pl. 1 (1964)
57; Ng, TFM 2 (1973) 253.
The family contains only one genus, Illicium, which has been the source of considerable
discussion regarding its evolutionary relationhips. Its affinities with the Magnoliales have
long been recognised, and this has been reflected historically by its classification in both the
families Magnoliaceae (e.g., Bentham & Hooker, Gen. Pl. 1 (1862) 16) and Winteraceae (e.g.,
Ridley, FMP 1 (1922) 18). The most recent comprehensive treatment of the genus is the
monograph by A.C. Smith (Sargentia 7 (1947) 1), who proposed that it should be isolated as
the family Illiciaceae on the basis of various morphological and anatomical criteria,
discussed in detail by Bailey & Nast (J. Arn. Arb. 26 (1945) 37; l.c. 29 (1948) 77), and Keng
(Bot. Bull. Acad. Sin. 6 (1965) 61). The Illiciaceae bears the closest relationship to the
Schisandraceae, a small family of scrambling and twining woody vines. The isolated
evolutionary position of these two families has been recognised more recently by their
classification as the sole members of the order Illiciales (e.g., Takhtajan, Bot. Rev. 46 (1980)
225; Cronquist, Integr. Syst. Fl. Pl. 1981); it is generally agreed, however, that the Illiciales are
derived from the Magnoliales, and are probably from the same stock as the Winteraceae.
ILLICIUM L.
(Latin, illicere = an attractant, probably in reference to the presence of aromatic oils)
Syst. Nat. ed. 10 (1759) 1050; Stapf, Trans. Linn. Soc. London Ser. 2, 4 (1894) 128; Ridley, FMP 1
(1922) 18 (under Winteraceae); A.C. Smith l.c. 10; Ng, l.c. 253.
Shrubs or small to medium-sized trees; evergreen, glabrous, aromatic with scattered ethereal
oil cells. Leaves simple, entire, alternate often clustered to give appearance of whorls of
36 at distal nodes; exstipulate; petioles with groove on adaxial surface; lamina papery to
leathery, venation pinnate; base generally attenuate, decurrent, apex generally acuminate.
Flowers solitary or in clusters of 23, mostly axillary, sometimes borne on the stem;
bisexual, regular; perianth not differentiated into sepals and petals, parts numerous (7 33),
free, overlapping, spirally arranged, white, cream, pink, red or purplish; androecium of (4)
numerous (up to c. 50) stamens, spirally arranged in one to several series, filaments short,
thick; anthers basifixed, introrse-lateral, dehiscing by longitudinal slits; gynoecium of
(5)715(21), free, superior carpels, arranged in a single whorl, obliquely attached to
receptacle; carpel unilocular, with single, near-basal, anatropous, bitegmic (with 2 layers of
integument), crassinucellar (with a thick nucellus) ovule, stigma dry, nonpapillate, decurrent.
227
228
ILLICIACEAE (SAUNDERS)
Fruits of single-seeded follicles, star-shaped, green (ripening red), splitting along ventral
edge of each segment when ripe. Seeds solitary in each segment, glossy, brown, with
copious oily endosperm.
Distribution. A medium-sized genus, with a disjunct distribution in Southeastern North
America (5 species) and eastern Asia (c. 35 species). In Asia the distribution extends from
southern Japan to northern Sumatra and Borneo, and from Assam to the Philippines, and the
centre of diversity lies in northern Burma and southern China (Yunnan, Guangxi and
Guangdong). In Borneo, it is restricted to the cooler montane regions, where two species, I.
kinabaluense and I. stapfii, are known.
Ecology. Vegetative growth is markedly discontinuous, with periods of dormancy of
vegetative buds alternating with active growth (Ng l.c.). The resumption of growth involves
the rapid elongation of a bud to produce a stem several centimetres long, which bears small
caducous leaves; apical "pseudowhorls" of leaves are then produced, consisting of alternately arranged normal leaves that are tightly clustered together. Studies in the reproductive
biology of North American species have shown that they are pollinated by a wide variety of
small insects, but primarily Diptera, and that a gametophytic self-incompatibility system
operates (Thien et al., Amer. J. Bot. 70 (1983) 719; White & Thien, J. Elisha Mitchell Sci.
Soc. 101 (1985) 15). A system of ballistic seed dispersal (autochory) is apparent in the
genus (Roberts & Haynes, Pl. Syst. Evol. 143 (1983) 227), although its efficacy has not
been demonstrated in the Bornean species.
Uses. The fruit of I. verum Hook. f. from China and Indo-China is the source of the spice
Chinese Star Anise, used for flavouring food and liqueurs. Although this species does not
occur in Borneo, the spice has been imported extensively from China and is traded in
Malaysia as bunga lawang or adas china (Burkill, EPMP 2 (1966) 1244). The fruits of the
Japanese species I. anisatum L. (syn. I. religiosum Sieb. & Zucc.) are poisonous, although
small quantities can be used for flavouring, and are sometimes retailed in SE Asia (Burkill
l.c.). Other Illicium species have various reported medicinal properties, often as stomachics,
carminatives, stimulants or vermifuges (Perry, MPESE (1980) 180). The timber is of very
limited value due to the small size of the trees.
Taxonomy. The last comprehensive revision of the genus was the monograph by A.C.
Smith (l.c.), in which 42 species were recognised. These species were classified into two
sections, viz. sect. Badiana Spach (which includes the type species and should therefore
bear the autonym sect. Illicium), and sect. Cymbostemon (Spach) A.C. Sm.; the Bornean
species belong to the latter section. Smith recognised three species in Borneo (I. kinabaluense,
I. stapfii and I. cauliflorum); the acceptance of these species was based on the examination
of only 11 herbarium sheets, however, and evidently requires re-evaluation. Only two
species are accepted in the present treatment.
Key to Illicium species

Leaves (5)67.5(11) x (1.5)23(4) cm. Stamens 78. Carpels 8.........1. I. kinabaluense
Leaves (6)1011.5(16.5) x (2)45(8.5) cm. Stamens 914 (20). Carpels 813.................
...........................................................................................................................2. I. stapfii
229
1. Illicium kinabaluense A.C. Sm.

l.c. 61. Type: J. & M.S. Clemens 50154, British North Borneo, Mt. Kinabalu, Penibukan (holotype A;
isotype L).
Small tree, to 15 m tall, 45 cm diameter. Leaves clustered in "pseudowhorls" of up to 6

leaves; lamina elliptic, c. (5)67.5(11) x (1.5)23(4) cm; base attenuate, margins slightly
revolute, apex (short-)acuminate; midrib slightly impressed above and prominent below;
lateral veins 58 per side; petioles 917 mm long, grooved on adaxial surface. Flowers
axillary or subterminal, generally solitary; pedicels 1020 mm long at anthesis; perianth
parts 810, pink, red or purplish, outermost ones 4.56 x 33.5 mm, largest 47 x 35 mm,
innermost 3.55 x 2 mm; androecium of 78 stamens, uniseriate, stamens 1.72.5 mm long;
gynoecium of 8 carpels, carpels 1.62 mm long.
Vernacular name. Sabahlongugan (Dusun).
Distribution. Restricted to Mt. Kinabalu, Sabah.
Ecology. Primary forests at 12002000 m.
2. Illicium stapfii Merr.
Fig. 1.
(Otto Stapf, 18571933, botanist at Kew in charge of "Indian" collections)

Philip. J. Sci. Bot. 13 (1918) 67, EB (1921) 252; Masamune, EPB (1942) 278; A.C. Smith l.c. 65;
Anderson, CLTS (1980) 346. Type: M.S. Clemens 10995, British North Borneo, Mt. Kinabalu, Marai
Parai Spur (PNH). Synonyms: Illicium sp. Stapf l.c. 128; I. cauliflorum Merr., Sarawak Mus. J. 3
(1928) 522, Masamune l.c. 277.
Medium-sized tree, to 25 m tall, 80 cm diameter. Leaves clustered in "pseudowhorls" of up

to 6 leaves; lamina elliptic, (6)1011.5(16.5) x (2)45(8.5) cm; base obtuse to attenuate,
margins markedly revolute, apex (short-) acuminate, acute or obtuse; midrib markedly
impressed above and prominent below; lateral veins 512 per side; petioles 1035 mm long,
grooved on adaxial surface. Flowers axillary or subterminal, occasionally borne on the bare
branches, solitary; pedicels up to 40 mm long at anthesis; perianth parts 915, pink, red or
purplish, outermost ones 58.5 x 45 mm, largest 59.5 x 3.56 mm, innermost (3.5)77.5
x 2.54.5 mm; androecium of 914(20) stamens, uniseriate, stamens 23.5 mm long;
gynoecium of 813 carpels, carpels 25 mm long.
Vernacular names. Sarawakala (Iban), bowlong (Kenyah).
Distribution. Endemic to Borneo. Montane regions, from Mt. Kinabalu (Sabah) in the north
to the Linau-Balui Plateau (Sarawak) in the south; also reported from northern Kalimantan.
Ecology. Primary forests at 8002000 m.
Taxonomy. Merrill (l.c. 1918) and A.C. Smith (l.c.) recognised I. cauliflorum as a distinct
species largely on the basis of its "cauliflorous" habit and greater number of carpels and
230
ILLICIACEAE (SAUNDERS)
stamens. Only the holotype was available to these authors for examination (E. Mjberg 114,
Mt. Murud, Sarawak (UC)). This specimen is recorded to have flowered at a height of only
1 m, and would more correctly be referred to as "ramiflorous". The study of numerous
collections of I. stapfii now available indicates that no clear distinction can be drawn
between ramiflory and the formation of flowers on young growth, suggesting that it is a
poor taxonomic character for the genus. The greater number of carpels and stamens in I.
cauliflorum is furthermore of little significance, since these characters are considerably
more variable in I. stapfii than observed by Smith (l.c.). There do not appear to be any
convincing criteria for retaining I. cauliflorum as a distinct species.
231
JUGLANDACEAE
c/o Forest Research Centre,
Sandakan, Malaysia
Lesch. ex Blume, Bijdr. 10 (1825) 528; Blume, Fl. Jav. Jugl. (1829) 5; Merrill, EB (1921) 210, Enum.
Philip. Pl. 2 (1923) 23; Ridley, FMP 3 (1924) 368; Masamune, EPB (1942) 231; Dilmy, Rimba
Indonesia 4 (1955) 29; Jacobs, FM 1, 6 (1960) 143; Backer & Bakhuizen f., FJ 2 (1965) 158; Burgess,
TBS (1966) 321; Manning, Bull. Torrey Bot. Club 93 (1966) 34; Whitmore, TFM 1 (1972) 233;
Cockburn, TFS 1 (1976) 151; Anderson, CLTS (1980) 220; Ashton, MNDTS 2 (1988) 303; Corner,
WSTM 1 (1988) 367; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 185.
Trees with aromatic tissues; leaves, young twigs and reproductive parts mostly lepidote with
golden yellow glandular scales. Stipules none. Leaves spirally arranged, paripinnate;
leaflet blade often slightly asymmetric. Inflorescence a panicle of catkins, axillary or
occasionally terminal, with flowers of one sex or both sexes in the same inflorescence; male
catkins usually pendulous; female catkins erect at least initially. Flowers unisexual, in axils
of 3-lobed bracts; perianth 4-lobed; male: perianth often reduced or irregular, stamens 413
(in Asiatic species) and sessile or on short filaments, anthers longitudinally dehiscing;
female: perianth lobes in 2 whorls, partly connate with the 2-carpellate ovary; carpels 1locular with an incomplete transverse septum, ovule 1; style 1, short or absent; stigmas 2(
4) in Engelhardia, persistent, papillose, or 4, sessile. Fruit a drupe or in Engelhardia a
small nut, attached to an enlarged wing-like 34-lobed bract. Seed 1, without endosperm;
cotyledons often much contorted.
Distribution. 7 genera and about 60 species, chiefly in north temperate regions in both the
Old and New World, with a few species in the tropics. In Sabah and Sarawak, represented
by one genus, Engelhardia, with 9 species.
Uses. Walnut (Juglans) and hickory (Carya) are important north temperate timber genera.
The family is also well known as the source of edible nuts, walnuts (Juglans) and pecan
(Carya). In Sabah and Sarawak, the timber of Engelhardia is too rare to be of much
commercial importance (see under genus).
Taxonomy. The family seems most closely allied to the Myricaceae within the Amentiferae.
ENGELHARDIA Lesch. ex Blume

(N. Engelhardt, 17611831, a Governor in Java)
sansanglang (Iban, Sarawak), tansanglang (Malay, Sarawak)
l.c. (1825) 528, l.c. (1829) 5, "Engelhardtia"; Merrill l.c. (1921) 210, l.c. (1923) 23; Ridley l.c. 368;
Masamune l.c. 231; Dilmy l.c. 29; Jacobs l.c. 143; Backer & Bakhuizen f. l.c. 158; Burgess l.c. 321;
233
Manning l.c. 34; Whitmore l.c. 233; Cockburn l.c. 151; Anderson l.c. 220; Ashton l.c. 303; Corner l.c.
367; Whitmore, Tantra & Sutisna l.c.185; Campbell-Gasis, Sandakania 3 (1993) 1. Synonyms:
"Engelhardtia" C. DC., Ann. Sc. Nat. 4, 18 (1862) 35; Pterilema Reinw., Syll. 2 (1826) 13;
Oreomunnea Oerst., Vid. Medd. Nat. For. Kjbenh. (1856) 52.
Small to medium-sized evergreen trees to 35 m (exceptionally to 50 m) tall, to 150 cm

diameter; bole columnar at first but later often of poor form with short flutes; buttresses thin,
small, but sometimes large, steep, thick and branching; crown generally oblong to
hemispherical, rather obtuse, with somewhat twisted branches. Bark from pale grey through
brown-grey to red-brown, smooth to fissured, peeling off in small, rectangular, papery
flakes, often purplish; inner bark from yellow to pinkish to brown, fibrous. Sapwood white
to yellow, soft to hard. Young twigs, leaves and inflorescences covered in golden or orange
scurfy scales and pale yellow and orange hairs (visible with a X10 hand lens). Twigs often
with conspicuous leaf-scars and cream to orange lenticels. Leaves often with a small
protuberance at the apex of the rachis; leaflets 27 pairs, alternate to subopposite, sessile or
stalked; base often unequal; veins often faintly looping near margin; above glabrous to
scaly, below variously scaly and hairy, vein axils often with tufted domatia (visible with a
X10 hand lens). Inflorescences so far known with flowers of only one sex. Flowers 4merous, tiny, often yellow or green, perianth inconspicuous; male: stamens 413, attached
to the fused perianth-bract structure; female: perianth partly connate with the ovary and
fused to a small one-sided 3-lobed bract, ovule erect and conical with a broad base. Fruits
attached to a 34-lobed bract; bract membranous, dry; the abaxial and lateral wings with
midribs and reticulate venation, the abaxial wing the largest, a smaller adaxial bract often
present; nut small, attached to base of bract; pericarp set with scales or hairs, or both.
Distribution. 12 species, 3 of which are found in the Americas (Mexico, Costa Rica and
Guatemala), the other 9 in the Old World from the western Himalayas to south China and
extending southwards across Malesia to New Guinea. All 9 species are found in Sabah and
Sarawak.
Ecology. Occurring as scattered individuals in lowland mixed dipterocarp forest below 2000
m, on leached acid yellow soils, but becoming common though never dominant at higher
elevations (up to 2700 m) in oak-laurel forest on more fertile soils and on ultramafic and
basaltic soils. Often shortly deciduous and then flowering. The flowers in catkins suggest
wind pollination but the male catkins of E. roxburghiana have been noted to have a slight
fragrance and may therefore be insect-pollinated also. The fruits are at least partially
dispersed by wind. The seedlings often have a terminal leaflet which, as the trees become
older, is reduced until all that is left in the adult is a small terminal protuberance. Saplings
and pole-sized trees tend to have leaves with more pairs of leaflets, which are usually
thinner, longer acuminate, sometimes serrate and often more densely hairy than those of
adult trees. Sterile trees may be mistaken for sapindaceous species because of the similarity
in their pinnate leaves.
Uses. The wood is locally used for timber but is of inferior quality, being not durable and
only of use under cover. The wood is rather soft to moderately hard, light to moderately
heavy, with a straight grain, shallowly interlocked, or slightly wavy. The texture is
moderately fine to slightly coarse and even. The timber is easy to work. The bark, containing
tannin-like compounds, is occasionally used for fish-poison in Sumatra.
234
JUGLANDACEAE (CAMPBELL-GASIS)
Key to Engelhardia species

(based on shoot and leaflet characters)
1.
Leaflet margins distinctly toothed in some part............................................................2

Leaflet margins entire or wavy, not toothed..............................................................3
2.
Distal leaflets often twice as large as the lower ones; stalk nil to 1 mm, rarely more.
Leaflet base acute or wedge-shaped; intercostal veins and areoles mostly invisible on
the upper surface in dried specimens.........................................................8. E. serrata
Distal leaflets sometimes larger than lower ones but not up to twice as large; stalk
longer, to 4 mm. Leaflet base rounded to subcordate; intercostal veins and areoles
distinct on the upper surface in dried specimens......................................1. E. apoensis
3.
Leaflets coriaceous, elliptic-obovate (or if lanceolate-falcate then with veins impressed

on the upper side), and not hairy on the lower side.......................................................4
Leaflets chartaceous, or lanceolate-falcate, or with hairy veins on the lower side (the
veins not impressed on the upper side).........................................................................6
4.
Leaflet intercostal venation coarse, the areolation strongly prominent under X10
magnification..................................................................................3. E. kinabaluensis
Leaflet intercostal venation finer, the areolation only faintly visible under X10
magnification..............................................................................................................5
5.
Shoot-apices and leaf rachises brown scurfy hairy.......................................6. E. rigida

Shoot-apices and leaf rachises not scurfy hairy.................................5. E. mersingensis
6.
Leaflets not decreasing markedly in size from apex to base of leaf rachis; veins on
lower side hairy.........................................................................................9. E. spicata
Leaflets decreasing in size by one third to half from apex of leaf rachis; veins on lower
side minutely scaly to subglabrous or apparently glabrous............................................7
7.
Leaflets elliptic, papery, base only slightly unequal, margin not thickened; stalk almost
nil or only to 1 mm long.....................................................................2. E. danumensis
Leaflets ovate to lanceolate or falcate, leathery, base very unequal, margin thickened;
stalk 110 mm long.....................................................................................................8
8.
Leaflets to 11 cm long, usually 45 pairs per leaf; stalk 14 mm long............................

.....................................................................................................4. E. mendalomensis
Leaflets usually larger, to 16 cm long, usually 23 pairs per leaf; stalk longer, to 10
mm.................................................................................................7. E. roxburghiana
Key to Engelhardia species

(based on fruit characters)
1.
Nut 614 mm long with a conspicuously narrowed apex..........................1. E. apoensis

Nut shorter, to 6 mm long, apex rounded.....................................................................2
235
236
2. Adaxial lobe of bract not developed, the nut fully exposed...........................................3

Adaxial lobe of bract well-developed, hiding most of the nut.......................................4
3.
Nut-stalk 48 mm long. Nut covered with scales only. Perianth-lobes broadly ovate
.......................................................................................................7. E. roxburghiana
Nut-stalk shorter, to 3 mm long. Nut hairy. Perianth-lobes linear.....5. E. mersingensis
4.
Perianth of inconspicuous tiny lobes adnate to the style..............................9. E. spicata

Perianth of ovate to linear lobes at the apex of the nut..................................................5
5.
Nut set with stiff, loose, irritant long hairs.................................................8. E. serrata

Nut indumentum not so...............................................................................................6
6.
Adaxial lobe of bract cleft in the middle.......................................................................7

Adaxial lobe of bract not cleft in the middle.................................................................8
7.
Adaxial lobe of bract only shallowly cleft. Perianth-lobes broadly ovate....2. E.

danumensis
Adaxial lobe of bract deeply cleft to the base. Perianth-lobes linear.......3. E.
kinabaluensis
8.
Adaxial lobe of bract orbicular....................................................................6. E. rigida

Adaxial lobe of bract subtruncate, the margin sometimes wavy.....4. E. mendalomensis
1. Engelhardia apoensis Elmer ex Nagel

(of Mount Apo in the Philippines)
Bot Jahrb. 50 (1914) 477; Elmer, Leafl. Philip. Bot. 7 (1915) 2693; Merrill l.c. (1923) 23; Jacobs l.c.
151; Burgess l.c. 321; Manning l.c. 48; Whitmore l.c. 234; Cockburn l.c. 152; Ashton l.c. 304; Corner
l.c. 368; Whitmore, Tantra & Sutisna l.c. 185. Type: Elmer 11744, Philippines, Mindanao, Mount
Apo (holotype NY; isotypes BM, BO, GH, L, P).
Tree to 50 m tall and 150 cm diameter; bole often straight; buttresses to 4 m tall and 1.5 m
out; crown spreading to dense. Bark grey-brown to reddish brown; inner bark pink with
white wedges to purplish and reddish brown, soft. Sapwood pale yellow to white. Twigs,
leaves and inflorescences set with golden to orange scales and pale yellow hairs. Leaf
rachis (8)1323(32) cm long, 3.511 cm to first leaflet, thickened at base, densely to
occasionally covered in scales and occasional hairs; leaflets (3)45(7) pairs, alternate,
ovate to lanceolate, (5)913(16) x (2.2)34(6) cm, gradually decreasing a little in size
towards base, drying orange-brown to dark chocolate-brown below, thinly leathery, upper
side with occasional scales and hairs, lower side with occasional to dense scales, only
midrib and lamina hairy; base rounded to subcordate, unequal, margin toothed irregularly,
at least in lower half, often inrolling, apex acute; tufted domatia occasional to frequent in
vein axils with midrib; lateral veins 916(18) pairs, above and below visible; intercostal
veins net- to ladder-like, above visible, below sometimes invisible; petiolule (1)23(4)
mm long, with dense to occasional scales and hairs. Inflorescences axillary, scaly and hairy;
male catkins 920 cm long, peduncle densely scaly and hairy; stamens 9, anthers scaly and
hairy; fruiting catkins (8)1219(30) cm long, peduncle and rachis scaly and hairy. Fruits
sessile to 3-mm-stalked; bracts with hairs and scales, abaxial wings 5575 x 1215 mm,
lateral wings 2835 x 8 mm, adaxial lobes entire to shallowly 4-lobed, hiding top of nut;
nuts ovoid, 614 x 56 mm, densely covered in long, soft hairs, without scales; perianthlobes adnate to the style, tiny, inconspicuous, not hiding the stigmas.
237
Distribution. Peninsular Malaysia, Borneo, and the Philippines. In Sabah, an uncommon

tree known on Mount Kinabalu only (e.g., SAN 27565, SAN 22408); uncommon also in
Sarawak, known only on Mount Dulit (Richards 1521). Also in E Kalimantan.
Ecology. Primary mixed dipterocarp forest from 700 to 1300 m, on fertile and ultramafic
soils.
2. Engelhardia danumensis Campbell-Gasis
Fig. 1.
(of the Danum Valley area in Sabah)

Sandakania 3 (1993) 6. Type: Cockburn SAN 85066, Sabah, Lahad Datu, Ulu Segama Forest Reserve,
Ulu Sungai Segama (holotype SAN; isotypes A, BO, FHO, K, KEP, L, SAR, SING).
Tree to 35 m tall and 60 cm diameter; buttresses thin, plank-like, c. 1 m tall and 2 m out.
Bark dark chocolate-brown; inner bark orange-brown, cut wood and bark smelling like
coconut water. Sapwood white, hard. Twigs and leaves covered with golden scales but
without hairs. Leaf rachis 410 cm long, 2.84.2 cm to first leaflet, blackish, not thickened
at base, scales occasional; leaflets 23 pairs, alternate to subopposite, elliptic, 2.47 x 1.5
1.5 cm, decreasing to about half the size towards base, drying dark brown to blackish
below, thin, papery; upper side glabrous, lower side with occasional to scattered scales all
over; base acute to wedge-shaped, subequal, margin entire, undulate, marginal vein not
thickened, apex acute to shortly acuminate, acumen to 5 mm long; tufted domatia occasional
in vein axils with the midrib; lateral veins 812 pairs, raised on both sides, conspicuous;
intercostal veins net-like, obscure on both sides, areoles obscure; petiolule absent to 1 mm
long, glabrous or with occasional scales. Inflorescences axillary, flowers not known;
fruiting catkins 1418 cm long, peduncle and rachis slender, angular, with golden scales and
hairs. Fruits sessile to 1-mm-stalked; bracts covered in scales and a few hairs, abaxial wings
1930 x 69 mm, lateral wings 811 x 34mm, adaxial lobe shallowly cleft in the middle,
often hiding the top of the nut; nuts globose c. 3 mm across, covered in golden scales and
short golden hairs; perianth-lobes on the nut apex ovate, free from style; not hiding the
stigmas.
Distribution. Endemic to Sabah. Uncommon, and known only from the type collection.
Vegetatively it approaches E. rigida but the leaflets almost blackish on drying, have a thin
and papery texture, the lower leaflets decrease in size by half, and the lamina is set with
scales but never hairs. Also, the nut in this species differs in being set with both scales and
short hairs.
3. Engelhardia kinabaluensis Campbell-Gasis

(of Mount Kinabalu in Sabah)
l.c. 7. Type: Gibot SAN 66824, Sabah, Ranau, Mamut Copper Mine (holotype SAN).
Tree to 24 m tall and 35 cm diameter. Bark grey to blackish; inner bark yellow to pale
white, exudate yellowish. Sapwood cream. Twigs, leaves and inflorescences set with golden
238
scales but without hairs. Leaf rachis 3.78 cm long, black, 1.53.5 cm to first leaflet,
thickened at base, occasionally with scales; leaflets 23 pairs, subopposite, elliptic to
obovate or falcate, 1.56.2 x 0.72.8 cm, not decreasing in size towards base, drying greybrown to orange-brown below, leathery, upper side glabrous, lower side glabrous to
sparsely scaly all over; base acute to round, very unequal, margin entire and wavy with
thickened marginal vein, apex acute to shortly acuminate, acumen to 2 mm long; tufted
domatia absent to rare in vein axils with midrib; lateral veins 57 pairs, flat to occasionally
impressed above, often obscure; intercostal veins net-like, coarse, areoles visible and
strongly prominent under X10 magnification; petiolule 23 mm, glabrous or occasionally
with scales. Inflorescences axillary; flowers not known; fruiting catkins 710 cm long, the
rachis scaly and hairy. Fruit stalk 12 mm long; bract occasionally scaly, hairs absent,
abaxial wings 2024 x 56 mm, lateral wings 812 x 23 mm, adaxial lobes deeply cleft in
the middle, not hiding the top of the nut; nuts transversely ellipsoid, 23 x 34 mm, with
golden scales and short, golden hairs; perianth-lobes on the nut apex linear, free from style,
not hiding the stigmas.
Distribution. Uncommon. In Sabah only known on or near Mount Kinabalu (at the Mamut
Copper Mine and in the Mesilau Area of Kundasang; e.g., SAN 74274, SAN 111603, besides the
type).
Ecology. Lowland mixed dipterocarp forest and submontane forest from 200 m to 1500 m.
Vegetatively, it approaches E. rigida but the leaflets with their thickly, leathery texture, the
cover of scales on the leaflets and nuts, and the lack of hairs on the leaflets distinguish it
from that species.
4. Engelhardia mendalomensis Campbell-Gasis

(of Mendalom Forest Reserve in Sabah)
l.c. 7. Type: Fidilis SAN 116714, Sabah, Tenom District, Mendalom Forest Reserve (holotype SAN).
Tree to 35 m tall and 80 cm diameter; buttresses to 3 m high and 4 m out. Bark pale browngrey; inner bark pale yellow to orange-brown. Sapwood cream-white to yellowish. Twigs
and leaves covered in golden scales, without hairs. Leaf rachis 614 cm long, 2.26 cm to
first leaflet, blackish, thickened at base, occasionally with scales and scant hairs; leaflets
(3)45 pairs, subopposite, ovate to lanceolate or falcate, 411 x 24 cm, often decreasing
to about two-thirds to half the size towards base, drying dark brown to blackish below,
thinly leathery, occasionally with scales but no hairs all over on both sides; base acute or
rarely round, very unequal, margin entire, slightly wavy, with thickened marginal vein, apex
acute to long-acuminate, acumen to 10 mm long; tufted domatia occasional to absent in vein
axils with midrib; lateral veins 711 pairs, above raised but rather obscure, below raised,
conspicuous; intercostal veins net-like, areoles on both sides obscure; petiolules 14 mm
long with scattered scales. Inflorescences axillary, flowers not known; fruiting catkins
1330 cm, peduncle and rachis quite stout, angular, drying blackish, scaly and hairy. Fruits
sessile to 1-mm-stalked; bracts with scales and hairs, abaxial wing 2150 x 810 mm,
lateral wings 1118 x 46 mm, adaxial lobes subtruncate (the margin sometimes wavy),
often hiding the top of the nut; nuts transversely ellipsoid, 34 x 45 mm, set with golden
scales and scattered, short golden hairs; perianth-lobes on the nut apex ovate, free from
style, not hiding the stigmas.
239
Distribution. Uncommon. In Sabah, only known from the type collection.

Ecology. Primary mixed dipterocarp forest from sea-level to 550 m.
Vegetatively it approaches E. roxburghiana but differs in having four to five pairs of leaflets
drying dark brown to blackish, with fewer pairs of veins and being less densely set with
scales. Also, the sessile to 1-mm-pedicellate nuts set with both hairs and scales distinguish it
from that species.
5. Engelhardia mersingensis Campbell-Gasis

(of Mount Mersing in Sarawak)
l.c. 8. Type: Ashton S. 16724, Sarawak, Bukit Mersing, Ulu Anap (holotype SAR; isotypes A, BO, K,
KEP, L, MEL, SAN, SING).
Tree to 30 m tall and 70 cm diameter; buttresses tall, sinuate and branching. Bark pale
orange-brown, thin, powdery. Twigs, leaves and inflorescences set with golden and orange
scales, but without hairs. Leaf rachis 4.514 cm long, 2.54.5 cm to first leaflet, black,
thickened at base, occasionally with scales, without hairs; leaflets 34 pairs, alternate to
opposite, elliptic to obovate, 4.511 x 2.54.8 cm, drying orange-brown to blackish below,
chartaceous, upper side glabrous, lower side occasionally scaly all over; base acute to
round, subequal to very unequal, margin entire, slightly wavy, with thickened marginal vein,
apex acute to acuminate, acumen to 9 mm long; tufted domatia occasionally in vein axils
with midrib; lateral veins 712 pairs, impressed or keeled above, below raised; intercostal
veins net-like, areoles only slightly visible under X10 magnification; petiolules 12 mm
long, with scattered scales. Inflorescences terminal or axillary, flowers not known; fruiting
catkins 9.520 cm long, peduncle and rachis quite stout, angular, drying dark brown or
blackish, scaly and hairy. Fruits sessile to 0.5-mm-stalked; bracts glabrous to scaly, without
hairs, abaxial wings 2031 x 710 mm, lateral wings 916 x 46 mm, adaxial lobes not
developed and not hiding the nut; nuts transversely ellipsoid, 3 x 46 mm, with scales and a
few short hairs present; perianth-lobes on nut apex linear, free from style, not hiding the
stigmas.
Distribution. Uncommon. In Sarawak known only from the type collection. Also in East
Kalimantan (on Gunung Beratus).
Ecology. Primary mixed dipterocarp to submontane forest on basaltic soils, at 6001000 m.
Vegetatively, this species approaches E. rigida but the larger leaflets set with scales but not
hairs, fewer pairs of lateral veins, and a sessile nut with scales and short hairs distinguish it
from that species.
6. Engelhardia rigida Lesch. ex Blume

(Latin, rigidus = stiff; the leaflets)
l.c. (1825) 528, l.c. (1829) 13, t. 3; Jacobs l.c. 148; Backer & Bakhuizen f. l.c. 158; Burgess l.c. 321;
Manning l.c. 40, 45; Cockburn l.c. 152; Anderson l.c. 220; Ashton l.c. 304; Whitmore, Tantra &
Sutisna l.c. 185. Type: Blume 1958, Java (holotype L; isotypes K, P). Synonyms: E. subsimplicifolia
Merr., Govt. Lab. Publ. Philip. 34 (1906) 6, l.c. (1923) 24; E. rigida var. subsimplicifolia (Merr.)
Manning l.c. 40; E. lepidota Schltr., Bot Jahrb. 50 (1913) 66; E. zambalensis Elmer l.c. 3195.
240
Tree to 50 m tall, and 90 cm in diameter; buttresses to 3 m high and 2 m out. Bark pale
brown-grey to reddish brown, soft; inner bark white, orange-yellow to brownish, soft.
Sapwood white. Twigs, leaves and inflorescences set with golden to orange scales and pale
yellow hairs. Leaf rachis (1)26(21) cm long, 1.52.5 cm to first leaflet, blackish,
thickened at base, sometimes densely scaly and hairy; leaflets typically 23 pairs, opposite,
ovate to elliptic, 1.57.1(16.5) x 1.44(7.5) cm, often drying orange-brown to brown
below, thickly leathery, often somewhat bullate; base acute to wedge-shaped, subequal,
margin entire, with thickened marginal vein, sometimes inrolled, apex shortly acuminate,
acumen 12 mm long; tufted domatia occasional to frequent in vein axils with midrib;
midrib on upper and lower sides glabrous to densely scaly and hairy; lateral veins 69 pairs,
distinct on both sides; intercostal veins ladder-like, indistinct on both sides; petiolules rarely
absent to 3(5) mm long, with scant to dense scales and sometimes hairs. Inflorescences
axillary on leafy and older leafless twigs, scaly and hairy; male catkins 1.56 cm long,
perianth-lobes narrowly triangular to completely reduced, stamens (3)46(7) with anthers
equal or unequal, 0.51 mm long, more or less white hairy; fruiting catkins (6)1015(22)
cm long, peduncle and rachis slender, round, more or less densely scaly and hairy. Fruits
0.51-mm-stalked; bracts scaly and hairy, abaxial wings 2060 x 38 mm, lateral wings
1214 x 34 mm, adaxial lobes orbicular or absent so the top of the nut is sometimes
exposed; nuts globose, c. 3.5 x 4 mm, densely covered by long, soft hairs, but without
scales; perianth-lobes on the nut apex ovate, free from style, not hiding the stigmas.
Distribution. Sumatra, Java, Borneo, the Philippines, Celebes, Moluccas, New Guinea. In
Sabah uncommon, known only on Mount Kinabalu; in Sarawak uncommon in lowlands and
mountains. Also in Brunei and Kalimantan.
Ecology. Primary mixed dipterocarp forest from sea-level to 2000(2500)m on both leached
yellow, sandy and ultramafic soils.
Uses. The wood is used for canoes and general building work in Sumatra.
Taxonomy. Manning (l.c.) distinguished two varieties of this species in Borneo, var.
subsimplicifolia and var. rigida. Var. subsimplicifolia was said to differ from the other in
having fewer pairs of leaflets and shorter fruiting catkins and fruits. The type specimen of E.
rigida var. rigida unfortunately does not have any fruits but otherwise looks like var.
subsimplicifolia with one extra pair of leaflets. The other specimens of E. rigida var. rigida
named by Manning had all the characters which Manning ascribed to var. subsimplicifolia,
including fewer leaflets, shorter fruiting catkins and smaller fruits. It is felt that E. rigida, at
least in Borneo, cannot be separated into these two varieties.
7. Engelhardia roxburghiana Wall.

(W. Roxburgh, 17511815, a Scottish botanist)
Pl. As. Rar. 2 (1831) 85, t. 199; Jacobs l.c. 154; Burgess l.c. 321; Manning l.c. 38, 48; Whitmore l.c.
235; Cockburn l.c. 154; Anderson l.c. 220; Ashton l.c. 305; Corner l.c. 369; Whitmore, Tantra &
Sutisna l.c. 185. Type: Wallich Cat. 4942, Malaya (holotype CAL). Synonyms: Juglans pterococca
Roxb., Hort. Beng. (1814) 68; E. chrysolepis Hance, Ann. Sc. Nat. 4, 15 (1861) 227; E. wallichiana
Lindl. ex Wall., Cat. (1831/32) no. 4942; E. pterococca (Roxb.) Kuntze, Rev. Gen. Pl. 2 (1891) 637;
241
E. spicata var. formosana Hay, Fl. Mont. Form. 6 (1908) 199; E. formosana Hay, Ic. Pl. Form. 6
(1916) 61; E. fenzelii Merr., Lingn. Sc. J. 7 (1931) 300.
Tree to 35 m tall and 70 cm diameter; buttresses to 2 m high, thin, branching; crown dense
or spreading. Bark fawn to dark chocolate-brown or black; inner bark yellow to reddish
brown, soft, moist; exudate clear, white. Sapwood white to pale yellow. Twigs often with
crusts of white exudate, together with leaves and inflorescences set with golden to orange
scales and pale yellow to orange hairs. Leaf rachis (5.5)614(17) cm long, 3.49.5 cm to
first leaflet, thickened at base, scantily to densely scaly and hairy; leaflets 23(4) pairs,
alternate to subopposite, ovate to lanceolate, (5)1016(23) x (1.4)3.55(8) cm, gradually
decreasing in size towards base, drying greenish to greyish brown or brown below,
leathery; base acute to wedge-shaped, very unequal, often somewhat decurrent, margin
entire, slightly wavy, with thickened marginal veins, sometimes inrolled, apex longacuminate, acumen 1017 mm long; tufted domatia absent to rare in the vein axils with the
midrib; midrib and veins above with occasional to scattered scales, below with occasional to
dense scales but without hairs; lateral veins 812 pairs, above raised to flat, below raised;
intercostal veins ladder-like, distinct above, indistinct below, areoles distinct above and
below; petiolules (1)610(12) mm long thickened at base, sparsely to densely scaly and
hairy. Inflorescences terminal on lateral twigs, paniculate; male catkins 810 per
inflorescence, 910 cm long, peduncle slender, scaly but without hairs; perianth-lobes 11.5
mm across; stamens 812, inserted (2)3 at the base of each perianth-lobe, anthers equal, c.
0.5 mm long, filaments equal to unequal, 0.30.5 mm long, glabrous, faintly sweet scented;
fruiting catkins 1017(23) cm long, peduncle and rachis slender, scaly and densely hairy.
Fruit stalk 45(8) mm long; bracts scaly but without hairs, abaxial wings 2845(55) x
58 mm, lateral wings 21 x 56 mm, adaxial lobes not developed so nut is visible; nuts
globose, 45 x 45 mm, with scales but without hairs; perianth-lobes at the nut apex ovate,
enclosing the 4 sessile stigmas.
Distribution. India to Indo-China, Sumatra, Peninsular Malaysia, and Borneo. In Sabah
uncommon, only known from the Crocker Range (SAN 44313); in Sarawak scattered
throughout mixed dipterocarp and submontane forests (S. 28588, S. 32811, Nooteboom &
Chai 2153), locally abundant as on the summit ridges of the basaltic Bt. Mersing and the
Tau Range, where it grows together with E. mersingensis. Also in Brunei at Ulu Mendamit.
Ecology. Primary mixed dipterocarp forest in hilly country, to 1750 m, on clay-rich, relatively
fertile soils.
8. Engelhardia serrata Blume

(Latin, serratus = toothed like a saw; the leaflet margin)
l.c. (1829) 14, t. 4; Jacobs l.c. 150; Backer & Bakhuizen f. l.c. 158; Burgess l.c. 321; Manning l.c. 40,
45; Whitmore l.c. 236; Cockburn l.c. 152; Anderson l.c. 220; Ashton l.c. 305; Corner l.c. 370;
Whitmore, Tantra & Sutisna l.c. 185. Type: Blume 2221, Java, Mt. Salak (holotype L). Synonyms: E.
palembanica Miq., Fl. Ned. Ind., Suppl. (1861) 346, 139; E. parvifolia C. DC. l.c. 34; E. nudiflora
Hook. f., Fl. Br. Ind. 5 (1888) 596; E. serrata Blume var. nudiflora (Hook. f.) Manning l.c. 46; E.
permicrophylla Elmer, Leafl. Philip. Bot. 9 (1934) 3194.
242
Tree to 30 (45) m tall, 120 cm diameter; buttresses to 34 m high and 3 m out. Bark light
to dark grey to reddish brown; inner bark from pinkish brown to orange- and reddish brown,
soft. Sapwood yellowish to white. Twigs, leaves and inflorescences set with golden and
orange scales and pale yellow hairs. Leaf rachis (2.6)518(24) cm long, 1.23.1 cm to
first leaflet, slightly thickened at base, densely to scantily covered with scales and hairs;
leaflets (2)36(9) pairs, alternate to subopposite, oblong to lanceolate in young trees to
obovate in older trees, (1)215(20.5) x (0.5)14(8) cm, often decreasing to about half
the size towards base, drying from ochre-brown to dark brown below, chartaceous in young
trees to leathery in older trees, upper side with only the midrib and veins sparsely to densely
scaly and hairy, lower side with scattered to dense scales and sparse hairs all over; base
acute to wedge-shaped, rarely rounded, subequal, margin regularly to irregularly toothed at
least in upper third, apex short- to long-acuminate, acumen 15 mm long; tufted domatia
frequent in vein axils with midrib; lateral veins 415 pairs, fewer in older trees, above
distinct or not, raised to flat; intercostal veins net-like, above and below distinct or not;
petiolules absent to 1(3) mm long, occasionally with dense scales and hairs. Inflorescences axillary on leafy or older leafless twigs, scaly and hairy; male catkins 23 per
inflorescence, 313 cm long, perianth-lobes more or less completely reduced, stamens (3)
57, anthers sometimes unequal and 0.51 mm long, filaments sessile to 0.3 mm long and
sparsely hirsute; fruiting catkins 914(30) cm long, peduncle and rachis stout, angular,
densely covered with scales and hairs. Fruits sessile to 1-mm-stalked; bracts glabrous or
scaly and hairy; abaxial wings 1640 x 611 mm, lateral wings 612 x 36 mm, adaxial
lobe frilled to shallowly 23-lobed, often hiding the top of the nut; nuts globose, c. 3 mm
across, sometimes with a few scales, densely covered in stiff, long hairs; perianth-lobes on
nut apex ovate, free from style, not hiding the stigmas.
Vernacular names. Sabahpusing-pusing (Malay). Sarawakentalun (Iban), momon
(Malay), owl (Murut), tepanga (Kenyah).
Distribution. Burma, Thailand, Indo-China, Sumatra, Peninsular Malaysia, Java, Borneo,
Philippines, and the Moluccas. In Sabah, not uncommon in the lowlands but also on the
Crocker Range. In Sarawak, scattered in lowland and submontane forests. Also in Brunei
and Kalimantan.
Ecology. Primary mixed dipterocarp forest, on sandy or clayey soils from sea-level to 2200
m; rarely in secondary forest.
Taxonomy. Manning l.c. recognised four varieties, viz. var. serrata, parvifolia (C. DC.)
Manning, nudiflora (Hook. f.) Manning, and cambodica Manning, of which the first three
occur in Sabah and Sarawak. These taxa seem to share the characters described above, and
with limited material available for study, it is impossible to say whether the varietal status
proposed by Manning is truly justified.
9. Engelhardia spicata Lesch. ex Blume

(Latin, spicatus = bearing a spike; the inflorescence)
243
l.c. (1825) 528; Merrill l.c. (1921) 210, l.c. (1923) 24; Ridley l.c. 368; Masamune l.c. 231; Jacobs l.c.
151; Backer & Bakhuizen f. l.c. 158; Burgess l.c. 321; Manning l.c. 41; Whitmore l.c. 236; Cockburn
l.c. 152; Corner l.c. 370; Whitmore, Tantra & Sutisna l.c. 186. Type: Blume, s.n., Java (holotype L;
isotypes K, NY, P). Synonym: E. philippinensis C. DC. l.c. 34, t. 2, f. 15.
var. aceriflora (Reinw.) Koords. & Valeton

Bijdr. Booms. Java 5 (1900) 167. Basionym: Pterilema aceriflorum Reinw., Syll. Pl. Nov. Soc.
Rolisb. 2 (1826) 13. Type: Reinwardt, s.n., Java (holotype L). Synonyms: E. aceriflora (Reinw.)
Blume l.c. (1829) 11, t. 2 and 5B; E. pterococca Roxb. ex Kuntze var. aceriflora (Reinw.) Kuntze l.c.
637.
Tree to 40 m tall, 280 cm diameter; buttresses sometimes present, to 3 m tall; crown dense,
rounded. Bark light brown-grey. Twigs with pale golden scales and hairs. Leaf rachis
(5.5)1030 cm long, to 5.5 cm to first leaflet, slightly thickened at base, brown or blackish,
glabrous to scaly and hairy; leaflets (2)35(7) pairs, subopposite to opposite, oblong to
lanceolate, (6)416(30) x (3)1.56(8) cm, not decreasing in size towards base,
leathery; upper and lower sides glabrous to sparsely scaly to very hirsute, or both scaly and
hairy, especially on the midrib and veins on the lower side; base rounded to subcordate,
very unequal, margin entire, wavy, apex shortly acuminate, tip obtuse to acute; tufted
domatia frequent in the vein axils with the midrib and often also in the axils of veins which
join near the margin, visible to the unaided eye; lateral veins 1118 pairs, above distinct,
raised, below often indistinct; intercostal veins indistinct above and below; petiolules absent
to 3 mm long. Inflorescences paniculate, axillary on leafy or older leafless twigs; male
(4)917 cm long; perianth-lobes present or reduced, very irregular, lobes to 2 mm long;
stamens 614, anthers equal or unequal, (sub)sessile, hirsute; fruiting catkins
(8)1930(40) cm long, peduncle and rachis angular, glabrous or scaly. Fruits sessile to
1-mm-stalked; abaxial wings 2060 x 715 mm, lateral wings 2025 x 58 mm, wings
often very irregular, adaxial lobes frilled to shallowly 45-lobed, often hiding the top of the
nut; nuts globose to transversely ellipsoid, 36 x 38 mm, with long, soft, dense hairs,
without scales; perianth-lobes tiny, inconspicuous, adnate to style, not hiding the stigmas.
Distribution. India to Indo-China, Sumatra, Peninsular Malaysia, Java, Borneo, the Philippines,
and Lesser Sunda Islands. In Sabah, uncommon on Mount Kinabalu (e.g., SAN 42865, SAN
61785); not reported from Sarawak or Brunei. Also in East Kalimantan.
Ecology. Primary forest, more common on mountains, from sea-level to 2500 m.
Taxonomy. Koorders & Valeton l.c. and Manning l.c. recognised three varieties, namely
var. acerifolia (Reinw.) Koord. & Valeton, colebrookeana (Lindl. ex Wall.) Kuntze, and
spicata, of which only var. acerifolia is so far found in Sabah.
244
MONIMIACEAE
P. C. Yii
Kuching, Malaysia
and
Lesmy Tipot
Kepong, Malaysia
Hooker f., Fl. Br. Ind. 5 (1890) 114; Merrill, EB (1921) 271, PEB (1929) 77; Ridley, FMP 3 (1924)
73; Masamune, EPB (1942) 305; Backer & Bakhuizen f., FJ 1 (1963) 116; Anderson, CLTS (1980)
253; Philipson, Blumea 28 (1982) 77, Blumea 30 (1985) 389, FM 1, 10 (1986) 255; Ng, TFM 4 (1989)
261; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 235.
Trees or shrubs, rarely woody climbers. Leaves simple, decussate or rarely alternate or in
whorls of three, usually with round oil-cells in the lamina, bearing simple or stellate hairs
or glabrous. Stipules absent. Inflorescences cymes, solitary or fascicled, terminal or axillary,
rarely on the trunk. Flowers unisexual or bisexual, regular; receptacles usually welldeveloped or rarely reduced, round, urceolate or bell-shaped; tepals inconspicuous, rarely
developing as distinct sepals and petals, decussate, in whorls or spirals. Male flowers with
few to many stamens arranged in whorls, spirally or irregularly, filaments strap-shaped,
anthers 24-loculed, opening through slits or valves. Female flowers with or without
staminodes; carpels few to many, sessile or stalked (stipitate), free or immersed in the
receptacles; ovule solitary, anatropous, erect or pendulous, with a thick nucellus
(crassinucellar), bitegmic or unitegmic. Fruits achenes or drupes, rarely plumose, usually
enclosed by the persistent receptacle, stalked or sessile, set free by splitting of the
receptacles. Seed one; endosperm copious, oily; embryo straight; cotyledons appressed or
divergent.
Distribution. About 33 genera and 320 species, mostly in the warmer parts of the southern
hemisphere (Malesia, Australia, SW Pacific, islands in the western Indian Ocean and S
America). In Sabah and Sarawak, represented by 2 genera with 3 species.
Taxonomy. The two genera (Kibara and Matthaea) occurring in Sabah and Sarawak are
included by Philipson (l.c. (1986) 261) in the subfamily Mollinedieae and characterised by
the male flowers having globose or flask-shaped receptacles and by drupaceous fruits borne
in heads. For a detailed account of the systematic position of the family see Philipson's
account (l.c. 1986).
245
Key to genera
Leaves ovate to broadly elliptic; lateral veins diminished and not joined toward the margin.
Stamens 69, arranged in 2 series, anthers opening by a single longitudinal slit. Fruit-stalk
shorter than 8 mm................................................................................................1. Kibara
Leaves narrowly lanceolate to lanceolate-oblong, lateral veins looped and joined at the
margin. Stamens usually 4, free, anthers opening by two longitudinal slits. Fruit-stalk longer
than 8 mm.............................................................................................2. Matthaea
1. KIBARA Endl.
( a Sundanese plant name)
Gen. Pl. (1837) 314; Hooker f. l.c. 114; Merrill l.c. (1921) 271, l.c. (1929) 77; Ridley l.c. 73;
Masamune l.c. 305; Backer & Bakhuizen f. l.c. 117; Anderson l.c. 253; Philipson l.c. (1985) 389, l.c.
(1986) 287; Ng l.c. 261; Whitmore, Tantra & Sutisna l.c. 235. Synonyms: Brongniartia Blume, Bijdr.
9 (1825) 455 (non Knuth); Sciadicarpus Hassk., Flora 25, 3 (1842) 20; Sarcodiscus Griff., Not. Pl. As.
4 (1854) 380.
Trees or shrubs with aromatic smell. Terminal vegetative buds conical or pyramidal,
surrounded by scale leaves. Leaves simple, decussate; blades entire or toothed, usually with
short, tufted, soft, brown hairs, gradually becoming glabrous; lateral veins arched and
diminishing toward the margins. Inflorescences terminal or axillary cymes (racemose in the
non-Bornean K. streimannii), paniculate or fasciculate; pedicels thickening distally into flat
or cup-shaped receptacles. Male flowers usually smaller than the females, with a minute
opening surrounded by 24 decussate pairs of tepals; stamens 69, arranged in 2 series,
with an outer series of 4(5) larger stamens, and an inner series of 4 smaller, often infertile
stamens, anthers opening by a single slit, with a filament or subsessile. Female flowers with
a minute opening surrounded by 5 decussate pairs of tepals, the inner pairs thickened and
grandular; carpels many, free on the inside of the receptacles; style very short; ovule 1.
Fruit a sessile or short-stalked drupe enclosed by enlarged receptacle. Seeds coniform;
seed-coat membranous; embryo small.
Distribution. About 40 species, in Peninsular Thailand, Nicobar Islands, Malesia and
Queensland (Australia). Only 2 species occur in Sabah and Sarawak.
Ecology. Mostly understorey shrubs and small trees in rain forest from sea-level to 2800 m;
occasionally on limestone hills, sandy and coral beaches.
Key to Kibara species

Leaves broadly ovate to elliptic-oblong, base rounded or subcordate, margins entire or
toothed, apex distinctly acuminate ................................................................1. K. coriacea
Leaves narrowly to broadly elliptic, base narrowly to broadly cuneate, margins entire, apex
obtuse...............................................................................................................2. K. obtusa
1. Kibara coriacea (Blume) Tulasne

Arch. Mus. Hist. Nat. Paris 7 (1855) 404; Hooker f. l.c. 114; Ridley l.c. 75; Merrill l.c. (1929) 77;
Masamune l.c. 305; Backer & Bakhuizen f. l.c. 117; Philipson l.c. (1985) 406, l.c. (1986) 298; Ng l.c.
246
MONIMIACEAE (YII & LESMY)
261; Whitmore, Tantra & Sutisna l.c. 235. Basionym: Brongniartia coriacea Blume l.c. (1825) 436.
Type: Blume, s.n., Java (L). Synonyms: K. blumei Steud., Nomencl. Bot. (1840) 846; Sciadicarpus
brongniartii Hassk. l.c. 20; Sarcodiscus chloranthiformis Griff. l.c. 350; K. chartacea Blume, Mus.
Bot. Lugd. Bat. 2 (1856) 89; K. cuspidata Blume l.c. (1856) 89, Merrill l.c. (1921) 271, Masamune l.c.
305; K. tomentosa and macrophylla J.R. Perkins, Bot. Jahrb. 25 (1898) 571; K. trichantha J.R.
Perkins l.c. (1898) 572; K. serrulata J.R. Perkins l.c. (1898) 575; K. angustifolia J.R. Perkins l.c.
(1898) 577; K. motleyi J.R. Perkins, Bot. Jahrb. 45 (1911) 424, Merrill l.c. (1921) 272, Masamune l.c.
305; K. grandifolia Merr., Philip. Govt. Lab. Bur. Bull. 29 (1905) 15; K. ellipsoidea Merr., Philip. J.
Sc. 1 (1906) Suppl. 56; K. mollis Merr., Philip. J. Sc. 3 (1908) Bot. 225.
Shrub or tree, to 15 m tall, 15 cm diameter. Bark smooth, pale grey; inner bark pale yellow.
Twigs slightly pubescent, gradually become glabrous. Leaves broadly ovate to ellipticoblong, 626.6 x 418.5 cm, leathery or papery, glabrous or sparsely to rather densely hairy
beneath; base cuneate, rounded or subcordate, margin entire or toothed toward the apex,
apex distinctly acuminate; midrib and lateral veins prominent beneath; lateral veins arched,
ascending and joining near the margins; stalk 525 mm long, slightly channelled above,
pubescent or glabrous. Inflorescence a terminal or axillary, simple 3-flowered or compound
cyme, with the male flowers at the proximal and the female flowers at the distal parts. Male
flowers rounded, 1.52 mm across, hairy; tepals 68 with rounded apex; stamens 4 in outer
series and 4 smaller ones in the inner series, filaments strap-shaped. Female flowers larger
than male flowers, rounded, about 35 mm across; tepals about 6 with swollen pendulous
glands within the minute opening. Fruit an ovoid drupe, c. 2 x 1.5 cm, ripening deep blue,
purple or black, on short swollen orange stalk; in clusters of 313. Seed coat membranous,
orange when dried.
Vernacular names. Sabahambibiliw, labak (Dusun).
Distribution. Throughout Malesia; in Sabah and Sarawak widespread.
Ecology. Lowland rain forests including swamp, coral beach, limestone hill, mixed dipterocarp
forests, and lower montane forests from sea-level to 1600 m.
Uses. The fruit is said to be edible and the leaves are used as flavouring in meat dishes.
2. Kibara obtusa Blume
Fig. 1.
(Latin, obtusus = blunt or rounded; the leaf apex)

l.c. (1856) 89; Philipson l.c. (1985) 409, l.c. (1986) 300. Type: Blume, s.n., Celebes (holotype L).
Synonym: K. depauperata Merr., Philip. Govt. Lab. Bur. Bull. 35 (1906) 13.
Tree to 20 m tall, 20 cm diameter. Bark scaly to shallowly fissured, pale yellow; inner bark
dull orange. Young twigs with short stiff-hairs. Leaves narrowly to broadly elliptic, 716.5
x 3.310 cm; base cuneate, margin entire, apex obtuse or rounded; midrib and lateral veins
prominent beneath, glabrous or with sparse stiff-hairs; stalk 1018 mm long, pubescent or
glabrous. Inflorescence a terminal or axillary, simple or compound cyme, to about 70 mm
247
long, with a pair of small bracteoles. Male flowers obovoid, c. 2 mm across; tepals 4,
minute; stamens usually 4 in the outer series and 2 smaller, infertile ones in the inner series.
Female flowers larger than male flowers, globose, about 2.53 mm across; tepals 4, apex
obtuse, with 4 swollen glands projecting among the carpels; carpels about 13, hairy, with
blunt stigmas. Fruit an ovoid drupe, 1724 x 1012 mm, ripening black, seated on a short
orange stalk. Seed coat membranous, orange in colour when dried.
Distribution. Borneo, Philippines, Celebes, and W New Guinea. In Sabah, uncommon and
only known from 4 collections, 3 from Lahad Datu (SAN 29844, SAN 31104, SAN 33382)
and one from Semporna (SAN 46055). Not yet reported in Sarawak.
Ecology. Primary rain forest from sea-level to 700 m.
2. MATTHAEA Blume
(Matteo de S. Guiseppe, 16171691, an Italian Missionary and Botanist in India)
l.c. (1856) 89; Hooker f. l.c. 115; Merrill l.c. (1921) 272; Ridley l.c. 73; Masamune l.c. 305; Anderson
l.c. 254; Philipson l.c. (1982) 77, l.c. (1986) 319; Ng l.c. 263.
Trees or shrubs. Leaves simple, opposite, entire or sub-serrate, leathery. Inflorescences

axillary, rarely terminal cymes, much shorter than leaves. Male receptacle subglobose;
tepals 4; stamens 4, free, filaments short, anthers opening by 2 longitudinal slits. Female
receptacle depressed globose; tepals 4, without apical pore or gap but upper half abscissing
as a calyptra at anthesis to reveal numerous carpels. Fruits long-stalked drupes, fleshy; one
or more on the enlarged receptacle.
Distribution. 6 species, all Malesian: Sumatra, Peninsular Malaysia, Anambas Island, Borneo,
Celebes, Philippines, and N Moluccas. Only 1 species has been recorded in Sarawak.
Ecology. Understorey trees in lowland and submontane forests to about 1700 m.
Matthaea sancta Blume
Fig. 2.
(Latin, sanctus = holy; alluding to the religious work of Matteo de S. Guiseppe)

l.c. (1856) 90; Hooker f. l.c. 115; Merrill l.c. (1921) 272; Ridley l.c. 73; Masamune l.c. 305; Anderson
l.c. 264; Philipson l.c. (1982) 82, l.c. (1986) 323; Ng l.c. 263. Type: Blume, s.n., Sumatra and
Borneo (holotype L). Synonyms: M. latifolia J.R. Perkins l.c. (1898) 563; M. calophylla J.R. Perkins
l.c. (1898) 563, Merrill l.c. (1921) 272, Masamune l.c. 305; M. ellipsoidea Merr. ex J.R. Perkins l.c.
(1911) 423.
Shrub or small tree, to 10 m tall, 10 cm diameter. Twigs green, glabrous. Leaves papery,
glabrous, lanceolate-oblong to oblong, 15.529 x 5.59.5 cm; base broadly cuneate, truncate or
rounded, margins entire or slightly toothed at the upper part, apex acuminate; lateral veins
arched, ascending and looping far from the margins, impressed on the upper surface,
prominently raised beneath; stalk 1.53 cm long, glabrous. Inflorescences axillary cymes.
Male flowers depressed globose, 23 mm across, opening by a small gap between a pair of
248
249
250
lip-like perianth-lobes; stamens 4. Female flowers similar to males but much larger, c. 10
mm across, on 14 pedicels arising in clusters of 35 from short peduncles; without any
apical pore or gap but the upper half abscissing as a calyptra at anthesis to reveal numerous
sessile carpels which are closely packed on a dish-like receptacle. Fruit a drupe, ellipsoid,
c. 2.5 x 1.5 cm, with thin bony endocarp, ripening blue or purple, attached in clusters of up
to 18 to an enlarged orange receptacle. Seed coat light brown when dried.
Distribution. Sumatra, Peninsular Malaysia, Anambas Is., Borneo, Philippines, and Celebes.
Uncommon in Sabah but widespread in Sarawak.
Ecology. Lowland rain forests, including limestone hill and mixed dipterocarp forests on
clay-rich soils, and lower montane forest, from sea-level to 1400 m.
251
NYSSACEAE
P. C. Yii
Kuching, Sarawak
Wangerin in Engler, Pfl. R. 41, 2 (1910) 1; Wasscher, Blumea 1 (1935) 343, FM 1, 4 (1948) 29;
Masamune, EPB (1942) 517; Hutchinson, Fam. Fl. Pl. 1 (1959) 175, Gen. Fl. Pl. 2 (1967) 50; Eyde, J.
Arn. Arb. 44, 1 (1963); Backer & Bakhuizen f., FJ 2 (1965) 161; Kochummen, TFM 1 (1972) 346;
Cockburn, TS 2 (1980) 61; Mabberley, PB (1987) 404; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990)
277.
Trees or shrubs. Leaves simple, spirally arranged, without stipules. Flowers more or less
regular, male or hermaphrodite, solitary or more often in axillary heads or condensed racemes;
calyx 5-toothed or absent; corolla composed of 5(10) imbricate petals; nectary disc
present; stamens (8)10(15), usually in 2 whorls, the outer whorl opposite the petals in
male flowers or as many as and alternate with the petals in hermaphrodite flowers, anthers
dorsifixed, dehiscing lengthwise; ovary inferior, 12-loculed, with 12 basally united styles,
ovule 1, pendulous, anatropous. Fruit a drupe. Seeds with scanty, oily endosperm; embryo
straight.
Distribution. 3 genera, with 810 species, distributed in N America, Mexico, E and SE
Asia. In Sabah and Sarawak, represented by 1 genus (Nyssa) with 1 species (N. javanica).
Taxonomy. The family is closely related to the Cornaceae with which it has been frequently
included in previous publications (Ridley, FMP 1 (1922) 889; Keng, OFMSP (1978) 219;
Brummitt, Vasc. Pl. Fam. & Gen (1992) 543). It can be distinguished from the Cornaceae by
the following characters: male flowers with tepaloid sepals and petals, stamens 810
arranged in two whorls; hermaphrodite flowers solitary or in heads, sepals and petals well
developed; style 1, rarely 2, each with 1 stigma; fruit a drupe.
NYSSA L.
(a legendary town in India where Bacchus,
the Greek God of Wine, was brought up by nymphs)
Sp. Pl. (1753) 1058; Ridley l.c. 895 (under Cornaceae); Wasscher l.c. (1935) 343, l.c. (1948) 29;
Masamune l.c. 517; Backer & Bakhuizen f. l.c. 161; Kochummen l.c. 346; Cockburn l.c. 61;
Whitmore, Tantra & Sutisna l.c. 277. Synonyms: Agathisanthes and Ceratostachys Blume, Bijdr.
(1825) 644; Agathidanthes Hassk., Cat. Hort. Bog. (1844) 254; Daphniphyllopsis Kurz, J. As. Soc.
Beng. 44, 2 (1875) 201.
253
254
NYSSACEAE (YII)
Dioecious trees or shrubs. Leaves simple, spirally arranged, without stipules. Flowers male
or hermaphrodite, often in heads, in the axils of a bract and with 2 bracteoles. Male flowers
in axillary heads or short racemes; calyx campanulate, rim smooth or 5-toothed; petals 5,
imbricate in bud, alternate with the calyx-lobes; stamens 816, in 2 alternating whorls;
anthers dorsifixed, opening lengthwise; disc pulvinate; ovary and style rudimentary.
Hermaphrodite flowers in 210-flowered, axillary, stalked heads; calyx entire or 5-toothed;
petals 58; stamens of the inner whorl partly sterile; ovary 1-locular, connate with the calyx,
style with 2 appressed, curving, often torulose branches, stigmatose inside. Fruits
drupaceous, ovoid to ellipsoid-oblong.
Distribution. About 6 species, 4 in the Atlantic N America, 1 in China, and 1 (N. javanica)
widespread from India to W Malesia.
Nyssa javanica (Blume) Wangerin
Fig. 1.
(of Java)
in Engler l.c. 15; Wasscher l.c. (1935) 344, l.c. (1948) 29; Masamune l.c. 517; Kochummen l.c. 346;
Cockburn l.c. 61; Whitmore, Tantra & Sutisna l.c. 277. Basionym: Agathisanthes javanica Blume l.c.
645. Type: Blume, s.n., Java (holotype L; isotype K). Synonyms: Agathidanthes javanica Hassk. l.c.
254; Nyssa sessiliflora Hook. f. & Th., Gen. Pl. 1 (1867) 952; Ilex daphniphylloides Kurz, J. As. Soc.
Beng. 39, 2 (1870) 72; Daphniphyllopsis capitata Kurz l.c. (1875) 201; Nyssa arborea Koord., Exk.
Fl. Jav. 2 (1912) 731; Nyssa bifida Craib, Kew Bull. (1913) 69.
Tree to 50 m tall, 90 cm in diameter, sometimes with very low buttresses. Bark grey,
smooth to slightly flaky, with prominent corky lenticels; inner bark dull yellow or pale
brown, fibrous or laminated, staining dark blue after slash. Sapwood heavy, yellowish
white. Twigs green, with large scattered lenticels and leaf-scars, often covered with brown
tomentum when young, gradually turning glabrous. Leaves typically crowded toward the
end of the twigs; stalk 1.54 cm long, hairy; blades slightly glaucous below, thinly leathery,
oblong-lanceolate to obovate, 519 x 27 cm; base gradually narrowed toward stalk, margin
entire to slightly wavy, apex abruptly pointed; midrib flushed reddish, slightly hairy; lateral
veins 811 pairs, hairy. Flowers in stalked, roundish, axillary heads 1218 mm across,
stalk slightly angular, 1050 mm long, glabrous or slightly hairy, bracteate; bracts
persistent in hermaphrodite flowers. Male flowers in 2040-flowered heads; calyx bellshaped, 45-toothed; petals 45, free, overlapping, recurved, shortly hairy; stamens 810 in
2 whorls. Hermaphrodite flowers in 39(18)-flowered heads; calyx campanulate, densely
appressed-sericeous, lobes 45, irregular, rounded or almost absent; petals 45; stamens
810, inner whorl sterile; ovary 1-chambered, style with 2 curving branches. Fruits
ellipsoid drupes, slightly flattened, to 22 x 15 mm, crowned by small persistent calyx. Seed
one, stone-like, grooved on one side and knobby on the other.
Vernacular name. Sarawakterang bulu (Kelabit).
Distribution. Sumatra, Peninsular Malaysia, Java and Borneo. In Sabah, the species is
known only from Mt. Kinabalu (SAN 22427, SAN 23507, SAN 42755, SAN 42785, SAN
60599, SAN 62216 and SAN 62220); in Sarawak, one collection so far (S. 35531), from
Kelabit Highlands, Bario.
255
Ecology. On gentle slopes and ridge tops in primary submontane forests at about 1400-1500 m.
Uses. The juicy mesocarp is edible. Wood heavy, occasionally used for local construction
work.
256
OCHNACEAE
K.M. Kochummen
Kepong, Malaysia
Ridley, FMP 1 (1922) 364; Merrill, EB (1921) 387; Masamune, EPB (1942) 468; Backer & Bakhuizen
f., FJ 1 (1963) 326; Kanis, Blumea 16 (1968) 1, FM 1, 7 (1971) 97; Ng, TFM 3 (1978) 253; Cockburn,
TS 2 (1980) 62; Anderson, CLTS (1980) 282; Whitmore, Tantra & Sutisna, CLK 2, 1, (1990) 278.
Shrubs or small to medium-sized trees. Leaves alternate, distichous or spiral, simple, often
toothed along the margins; stipules persistent. Inflorescences axillary or terminal, 1
many-flowered, cymose, racemose or thyrsoid. Flowers bisexual, regular; sepals 5, usually
free, persistent; petals 510, free; stamens 510many, anthers opening by longitudinal
slits or apical pores; staminodes 0many; carpels (2)510(15), superior, free with one
ovule or united with 2many ovules per cell; styles united. Fruit a drupe, berry or capsule.
Distribution. About 30 genera and 250 species, pantropical, rarely subtropical. In Sabah
and Sarawak, represented by 7 genera and 9 species.
Ecology. Lowland to montane forests to 3000 m. The species are adapted to poor sandy or
peaty soils in undisturbed forests except Schuurmansia which is a genus of pioneer species.
Pollination is probably by insects because of the brightly coloured flowers. The bluish or
black 1-seeded fruits are likely to be dispersed by birds. However in the many-seeded
capsules of those species in the subfamily Sauvagesioideae the mode of dispersal is not
understood.
Uses. Some species are used locally in traditional medicine.
Taxonomy. The Ochnaceae is divided into two subfamilies, the Ochnoideae and
Sauvagesioideae which separately appear much more natural than the family itself.
Subfamily Ochnoideae, consisting in Sabah and Sarawak of Brackenridgea and Gomphia, is
characterised by 10many stamens, free carpels that share a common style, one-seeded
drupes on a swollen receptacle, and a dimorphic shoot system consisting of orthotropic
leaders and plagiotropic branches. Subfamily Suavagesioideae, consisting of the remaining
5 genera, is characterised by 5 stamens, united carpels, a berry-like or capsular fruit with
more than one seed, and orthotropic shoots and branches. The genus Tetramerista which
Ridley included under Ochnaceae has been assigned to the family Tetrameristaceae.
257
Key to genera
1.
Shrubs to medium-sized trees. Stamens 10; carpels 510, free. Fruit a 1-seeded
drupe.........................................................................................................................2
Shrubs. Stamens 5; carpels 25, united. Fruit a capsule or berry with more than one
seed.............................................................................................................................3
2.
Leaves with distinct marginal veins, margin toothed; stipules united. Anthers opening
by apical pores............................................................................................3. Gomphia
Leaves without marginal vein, margin entire; stipules free. Anthers opening by
longitudinal slits...............................................................................1. Brackenridgea
3.
Leaf margin toothed. Ovary 5-celled. Anthers opening by apical pores. Fruit a berry
...................................................................................................................2. Euthemis
Leaf margin entire or toothed. Ovary 1-celled. Anthers opening by longitudinal slits.
Fruit a capsule.............................................................................................................4
4.
Branches hollow. Leaves shiny, lateral veins invisible. Seeds winged........4.

Schuurmansia
Not as above................................................................................................................5
5.
Stem coarsely bristly with persistent stipules and bracts. Inflorescences axillary, oneflowered........................................................................................................................
Neckia Korth.
Ned. Kruidk. Arch. 1 (1848) 358; Merrill l.c. 388; Ridley l.c. 134; Masamune l.c. 469; Kanis
l.c. (1968) 69, l.c. (1971) 110.
Monotypic genus (N. serrata Korth.) confined to Sumatra, Peninsular Malaysia,

Borneo (Sabah, Brunei, Sarawak, Kalimantan) and the Philippines. Lowland to
submontane forests to 1200 m.
Understorey shrubs. Stipules comb-shaped. Leaves spiral, blade with toothed
margin. Inflorescences axillary; rachis thread-like, 15 cm long; bracts many; only
single flower developing; pedicels jointed. Flowers 5-merous; stamens 5, anthers
opening by longitudinal slits, staminodes many; carpels 3, united, ovary 1-celled,
with distinct style and stigma. Fruit a capsule, splitting into 3. Seed not winged,
pitted.
Stem not so. Inflorescences terminal, with many flowers..............................................6
6.
Leaves oblanceolate, subsessile. Staminodes 10.............................................................

Indovethia Boerl.
Feestbundel P.J. Veth (1894) 89; Merrill l.c. 388; Masamune l.c. 469; Kanis l.c. (1968) 72,
l.c. (1971) 112; Anderson l.c. 283.
Monotypic genus (I. calophylla Boerl.). Central Sumatra and NW Borneo. Uncommon, in moist shady places in lowland forest.
Shrublets. Stipules comb-shaped. Leaves spiral, margin entire or toothed. Flowers:
sepals 5, persistent in fruit; petals 5, free; stamens 5, anthers opening by longitudinal
slits, staminodes 10; carpels 3, united, ovary 1-celled, with distinct style and
stigma. Fruit a capsule, splitting into 3 including style. Seed not winged, surface
pitted.
Leaves linear with distinct petiole. Staminodes many......................5. Schuurmansiella
258
OCHNACEAE (KOCHUMMEN)
1. BRACKENRIDGEA A. Gray
(W.D. Brackenridge, 18101893, the Scots-American botanist and horticulturist)
New Gen. Pl. (1853) 5; Merrill l.c. 387; Masamune l.c. 468; Furtado, Gard. Bull. Sing. 19 (1962) 181;
Kanis l.c. (1968) 41, l.c. (1971) 101; Ng l.c. 256; Cockburn l.c. 64; Anderson l.c. 282; Whitmore,
Tantra & Sutisna l.c. 278.
Small to medium-sized trees. Stipules small, often divided, falling off early. Leaves
alternate-distichous, glossy above; lateral veins strongly curved to the apex, often some of
the lower ones parallel to the margin. Inflorescences thyrsoid of condensed cymes; rachis
often growing vegetatively after flowering; bracts many, at base of inflorescences, falling
off early leaving distinct ring-like scars; pedicels jointed at base, filiform. Flowers arranged
in 14 tiers; sepals 5, accrescent, fleshy and red in fruit; petals 5(10), white or yellow;
stamens 10 (or many), anthers dehiscing by longitudinal slits; carpels 5(10), free, but
sharing a common style; ovules camptotropous or epitropous; stigma small. Fruit a drupe,
12(5), greenish becoming almost black on ripening, surrounded by persistent sepals.
Distribution. About 8 species, tropical E Africa, Madagascar, Malesia; 2 species in Sabah
and Sarawak.
Ecology. Everwet tropical areas to 1000 m. Dispersal is mainly by birds because of the
conspicuous black fruits on the red torus and calyx. Flowering and vegetative growth tend
to occur at alternative intervals during the growth of the stem. After a period of leaf
production, the terminal bud is covered with closely spaced scale-leaves, some of which
bear inflorescences in their axils. When flowering and fruiting is completed the terminal bud
resumes vegetative activity to bear foliage leaves again. This pattern of growth can be traced
through the series of alternating leaf-scars and peduncle-bases along the twig.
Taxonomy. Two sections are recognised in the genus. In section Brackenridgea, all flowers
in a cyme open simultaneously; the corolla is 5-merous, white; there are 10 stamens in one
whorl; and 5 carpels. In section Notochnella, the flowers of a cyme open successively; the
corolla is yellow and irregular; there are many stamens in more than 1 whorl; and 510
carpels. Section Notochnella is confined to the Philippines.
Key to Brackenridgea species

Inflorescence a many-flowered cyme; pedicels in two or more tiers..................1. B. hookeri
Inflorescence a 3(5)-flowered cyme; pedicels in one tier only.......................2. B. palustris
1. Brackenridgea hookeri (Planch.) A. Gray
Fig. 1.
(W.J. Hooker, 17851865, sometime Director of the Kew Botanic Gardens)

l.c. 6; Furtado l.c. 182; Kanis l.c. (1968) 45, l.c. (1971) 102; Ng l.c. 256; Anderson l.c. 283; Whitmore,
Tantra & Sutisna l.c. 278. Basionym: Gomphia hookeri Planch. in Hooker, Lond. J. Bot. 6 (1847) 3.
Type: Philips, s.n., Penang (K). Synonyms: Gomphia corymbosa (King) Ridl., J. Str. Br. R. As. Soc.
54 (1910) 36 p.p. excl. typus; Brackenridgea denticulata Furtado l.c. 183.
259
Tree to 30 m tall, 60 cm diameter. Bark reddish brown, smooth to slightly fissured and
scaly; inner bark reddish brown. Leaves elliptic to oblong, rarely obovate, 617 x 2.57.5
cm; base cuneate, apex pointed; midrib raised above; lateral veins of lower half of leaf
curving upwards to almost to leaf apex, running parallel to margins; petiole 0.61 cm long.
Flowers clustered in two or more tiers; pedicel 12 cm long. Fruits 69 x 5.56 mm.
Distribution. India, Andamans, Thailand, Peninsular Malaysia and Borneo. Uncommon and
of scattered occurrence in Sabah and Sarawak.
Ecology. Lowland mixed dipterocarp forest on sandy humult ultisols to submontane forests
to 1000 m, including kerangas and dry hillocks in swampy forests.
2. Brackenridgea palustris Bartelli

(Latin, palustris = inhabiting boggy or marshy ground)
Malpigia 15 (1901) 165; Merrill l.c. 387; Masamune l.c. 468; Furtado l.c. 182; Kanis, l.c. (1968) 46,
l.c. (1971) 102; Ng l.c. 256; Cockburn l.c. 64; Anderson l.c. 283; Whitmore, Tantra & Sutisna l.c. 278.
Type: Beccari PB 3472, Borneo (holotype FI; isotypes A, K, P). Synonyms: Gomphia hookeri
(Planch.) A. Gray var. corymbosa King, J. As. Soc. Beng. 62 (1893) 233; B. serrulata Bartelli l.c. 163,
Merrill l.c. 387, Masamune l.c. 469; Gomphia corymbosa (King) Ridl. l.c. (1910) 33, p.p., FMP 1
(1922) 367.
Tree to 30 m tall, 40 cm diameter. Bark brown to reddish brown, smooth to scaly, inner
bark reddish brown. Sapwood white to pale yellow. Leaves elliptic to oblong, 5.510 x 24
cm; base cuneate, apex pointed; midrib raised above; lateral veins of lower half curving
upwards, running parallel to the margin towards apex; intercostal veins invisible to very
faint; petiole 0.51 cm long. Flowers fascicled in one tier only, pedicel jointed. Fruits to 8
x 6 mm.
Distribution. Sumatra, Peninsular Malaysia, Borneo, Philippines, Celebes. Uncommon in
Sabah and Sarawak.
Ecology. Usually on lowlands, especially in peat swamp and kerangas forests, rarely in
submontane forests to 1000 m.
2. EUTHEMIS Jack
(Greek, eu = good, themis = law; the even thickness and symmetry of the leaves)
Mal. Misc. 1, 5 (1821) 5; Merrill l.c. 388; Ridley l.c. (1922) 367; Masamune l.c. 469; Kanis l.c. (1968)
62, l.c. (1971) 108; Ng l.c. 257; Anderson l.c. 283; Ashton l.c. 62.
Shrubs. Stipules falling off early. Leaves spiral, with numerous close, almost parallel
lateral veins; margin thick-rimmed, distinctly or faintly toothed. Inflorescences terminal
and axillary racemes. Flowers: sepals persistent in fruit; petals 5, white or pink; staminodes
05; stamens 5, anthers opening by apical pores; carpels 5, united, ovary 5-
260
261
262
celled, ovules 2 per cell, pendulous, axile; style distinct, stigma minute. Fruit a berry with 5
pyrenes. Seeds 12 per cell.
Distribution. 2 species; SW Cambodia, Sumatra, Peninsular Malaysia, Borneo.
Ecology. Kerangas forests, on low ridges, in peat swamp forests and also on ridges with
poor sandy soils, below 1250 m.
Uses. E. leucocarpa roots are used in traditional medicine in Peninsular Malaysia and in
Brunei; the fruits are used to treat eye diseases.
Key to Euthemis species

Leaves 840 cm long; apex pointed; margin distinctly toothed; petiole 25 cm long. Mature
fruit white......................................................................................1. E. leucocarpa
Leaves 415 cm long; apex rounded to slightly notched, with short mucro; margin faintly
toothed; petiole to 1.5 cm. Mature fruit red.........................................................2. E. minor
1. Euthemis leucocarpa Jack
Fig. 2AB.
(Greek, leuco = white, karpos = fruit)

l.c. 16; Merrill l.c. (1921) 388; Ridley, FMP 1 (1922) 368; Masamune l.c. 469; Kanis l.c. (1968) 62,
l.c. (1971) 108, Ng l.c. 257; Anderson l.c. 283. Type: Wallich 2516, Singapore (K, neotype).
Synonym: E. robusta Hook. f. in Ridley l.c. 368.
Shrub to 2 m tall. Stipules ovate. Leaves oblong to linear-oblong, 840 x 210 cm; base
tapered, margin distinctly toothed, apex acute; midrib raised above; lateral veins numerous,
12 mm apart; petiole winged, 25 cm long. Inflorescences in branched racemes. Fruits
globose, 610 mm in diameter, white when mature.
Distribution. Cambodia, Sumatra, Peninsular Malaysia, and Borneo. Widely distributed in
Sabah and Sarawak.
Ecology. Lowland to submontane forests to 1000 m, on poor soils. In Sarawak frequent in
mixed swamp and kerangas forests.
2. Euthemis minor Jack
Fig. 2CD
(Latin, minor = small; the smaller size compared to E. leucocarpa)

l.c. 18; Merrill l.c. (1921) 388; Ridley l.c. (1922) 368; Masamune l.c. 469; Kanis l.c. (1968) 65, l.c.
(1971) 108; Ng l.c. 257; Cockburn l.c. 62; Anderson l.c. 283. Type: Wallich 2517, Singapore (K,
neotype). Synonyms: E. obtusifolia Hook. f., Trans. Linn. Soc. 23 (1862) 163, Merrill l.c. (1921) 389,
Masamune l.c. 469; E. ciliata Pearson, Kew Bull. (1906) 3; E. hackenbergii Diels, Bot. Jahrb. 60
(1926) 310.
263
264
Shrub or treelet to 5 m tall. Twigs black. Leaves oblong to oblonceolate, 415 x 1.54 cm;
base tapered, margin faintly toothed, apex rounded with a short point; midrib raised above;
lateral veins c. 1 mm apart, very faint to invisible; petiole 0.51.5 cm long. Inflorescence
usually an unbranched raceme. Flowers sepals and petals pinkish when fresh. Fruits
globose, 46 mm in diameter, with 5 prominent ridges.
Distribution. Sumatra, Peninsular Malaysia, Borneo. Common in Sabah and Sarawak.
Ecology. Lowland to submontane forests to 1250 m; locally abundant in Sarawak in kerangas
and padang alan forest.
3. GOMPHIA Schreb.
(Greek, gomphos = a thorn or spike;
the form of the flower base during fruit development)
Gen. Pl. ed. 8 (1784) 291; Ridley l.c. (1922) 365; Kanis, Taxon 16 (1967) 420, l.c. (1968) 51, l.c.
(1971) 105; Ng l.c. 258; Cockburn l.c. 65; Anderson l.c. 283; Whitmore, Tantra & Sutisna l.c. 278.
Synonyms: Campylocerum Tiegh., Bull. Mus. Hist. Nat. Paris (1902) 546; Meesia Gaertn., Fruct. &
Sem. Pl. 1 (1788) 344.
Shrubs or small to medium-sized trees. Stipules united, falling off early. Leaves spiral,
margin firmly toothed; lateral veins numerous, close and almost parallel, with 23 distinct
intramarginal veins; petiole to 3 mm long. Inflorescences in terminal and axillary panicles,
peduncle persistent. Flowers: sepals 5, enlarged and persistent in fruit; petals 5; stamens
10, anthers opening by apical pores; gynophore columnar, 5-ribbed; carpels 5, free. Fruit a
drupe. Seed one.
Distribution. Mainly African; 1 species in SW India, Sri Lanka, E Thailand, Indo-China,
Hainan and W Malesia.
Ecology. Confined to everwet tropical and moderately dry monsoon areas from lowlands to
1500 m. Dispersal is presumed to be by birds.
Gomphia serrata (Gaertn.) Kanis
Fig. 3.
(Latin, serratus = with teeth; the leaf margin)

l.c. (1967) 418, l.c. (1968) 53, l.c. (1971) 105; Ng l.c. 258; Cockburn l.c. 65; Anderson l.c. 283;
Whitmore, Tantra & Sutisna l.c. 278. Basionym: Meesia serrata Gaertn. l.c. (1788) 344. Type:
Koenig 25, Ceylon (L). Synonyms: Gomphia sumatrana Jack l.c. 29; G. microphylla Ridl. l.c. (1922)
369; G. oblongifolia Ridl., Kew Bull. (1925) 281; Ouratia angustifolia (Vahl) Baill. ex Laness, Rev.
Gen. Pl. (1891) 106, Merrill l.c. 387, Masamune l.c. 470; O. borneensis Bartelli l.c. 158, Merrill l.c.
387, Masamune l.c. 470; O. neriifolia Bartelli l.c. 158, Merrill l.c. 387, Masamune l.c. 470; O.
beccariana Bartelli l.c. 159, Merrill l.c. 387, Masamune l.c. 470; O. megacarpa Ridl., Kew Bull.
(1930) 76, Masamune l.c. 470.
Shrub or medium-sized tree, to 20 m tall, 40 cm diameter; bole slightly fluted. Bark dark
grey-brown, scaly; inner bark pink, fibrous. Sapwood pale. Leaves shiny above, very
265
266
variable in shape and size, elliptic, oblong, or obovate, 5.533 x 27.5 cm; base cuneate,
margin faint to distinctly toothed, apex pointed; midrib raised above; lateral veins
numerous, close, almost parallel, very faint to almost invisible on both surfaces, forming
13 distinct wavy intramarginal veins; intercostal veins finely reticulate, very faint; petiole
to 3 mm long. Inflorescences axillary and terminal panicles. Flowers: pedicel c. 1 cm long;
sepals 5, tinged pink, enlarged in fruit; petals 5, yellow or cream, obliquely obovate to
broadly spathulate; stamens 10, subsessile or with very short filaments, anthers opening
with 2 apical pores, obovoid; carpels 5, free, with a single style. Fruits kidney-shaped,
12(5), yellowish green, ripening purplish, 0.51 x 0.40.6 cm, with persistent sepals.
Seed one.
Vernacular names. Sabahantimagas gimbaan (Murut), kolambang (Dusun), majangmajang (Kadazan), quintalai (Kadazan). Sarawakaam (Kenyah), chinaga-lampong,
keladang, kelutak (Iban), ladin (Malay).
Distribution. SW Peninsular India, Sri Lanka, E Thailand, Indo-China, Hainan, Malesia.
Common and widely distributed in Sabah and Sarawak.
Ecology. Lowland forests on well-drained infertile organic soils, including mixed dipterocarp
and kerangas forests, to submontane forest to 1500 m, sometimes occurring by stream
banks, rarely on limestone forests. Flowering and fruiting samples have been collected
throughout the year. As this species is found in a number of ecological habitats there is
considerable variation in its morphological characters. This variation is more pronounced in
Sarawak. In the more exposed habitats, on cliffs and on poor kerangas soils, this species has
very small leaves.
Uses. Roots and leaves are bitter and are decocted in S India for a stomachic and antiemetic
tonic. Young branch tissue is used to treat toothache in Cambodia.
4. SCHUURMANSIA Blume
(J. Schuurmans Stekhoven, 17921855,
horticulturist at the Leiden University Botanical Garden)
Mus. Bot. Lugd. Bat. 1 (1850) 177; Masamune l.c. 470; Kanis l.c. (1968) 74, l.c. (1971) 115;
Anderson l.c. 283; Whitmore, Tantra & Sutisna l.c. 278.
Shrubs or small trees. Twigs hollow. Stipules entire, narrow, lanceolate. Leaves spiral,
shiny, margin entire or toothed, glandular; lateral veins invisible. Inflorescences in terminal
panicles. Flowers: sepals 5; petals 5; stamens 5, anthers opening by longitudinal slits,
staminodes many in one or two whorls; carpels 3, united, ovary 1-celled. Fruit a capsule,
splitting into 3. Seeds winged.
Distribution. 3 species; Borneo, Philippines, Celebes, Moluccas, New Guinea, Solomon
Islands, Bismarks; 1 species in Borneo.
Ecology. Pioneer plants from sea-level to 3000 m.
267
Schuurmansia elegans Blume
Fig. 4.
(Latin, elegans = elegant; the growth habit)

l.c. 178; Kanis l.c. (1968) 75, l.c. (1971) 115; Anderson l.c. 238; Whitmore, Tantra & Sutisna l.c. 278.
Type: Zippelius, s.n., Amboina (L). Synonyms: S. parviflora Ridl., Trans. Linn. Soc. 2, Bot. 9 (1916)
18; S. borneensis Ridl., Kew Bull. (1930) 77, Masamune l.c. 470.
Small tree to 5 m tall. Leaves obovate to oblong, 1030 x 2.510 cm, shiny, surface of dried
leaves finely reticulate; base tapered, apex rounded to blunt; lateral veins many, close,
almost parallel, very faint on both surfaces; petiole 56 cm long. Flowers yellow. Fruits
fusiform, c. 2.5 x 0.5 cm, with pointed tip and persistent sepals. Seeds c. 3 mm long with
slender wings.
Distribution. Borneo, Philippines, Celebes, Moluccas, New Guinea. In Sabah and Sarawak
uncommon, known by only 2 collections (Sabah, SAN A 3636; Sarawak, Nooteboom 1517).
Ecology. Lowlands and hills including limestones, to 900 m.
5. SCHUURMANSIELLA Hall. f.
Rec. Trav. Bot. Neerl. 10 (1913) 344; Merrill l.c. 387; Masamune l.c. 470; Kanis l.c. (1968) 73, l.c.
(1971) 113; Anderson l.c. 283.
Shrubs. Stipules needle-like. Leaves spiral, margin finely toothed. Inflorescences terminal
racemes. Flowers: sepals 5, petals 5, staminodes many, stamens 5, anthers opening by
longitudinal slits; carpels 3, united, ovary 1-celled, style short, stigma capitate. Fruit a capsule,
splitting into 3. Seeds hairy.
Distribution. Monotypic, endemic to Borneo (Sarawak).
Ecology. Mainly lowlands, especially in kerangas forests on poor soils and on sandstone
cliffs near sea, rarely to 600 m.
Schuurmansiella angustifolia (Hook. f.) Hall. f.
Fig. 5.
(Latin, angustus = narrow, folium = leaf)

l.c. 345; Merrill l.c. (1921) 387; Masamune l.c. 470; Kanis l.c. (1968) 73, l.c. (1971) 114; Anderson
l.c. 283. Basionym: Schuurmansia angustifolia Hook. f. l.c. 157. Type: Lobb, s.n., Sarawak (K).
Shrub to 10 m tall. Stipules to 12 x 0.5 mm. Leaves linear-oblong, 817 x 0.71.45 cm;
base tapered, apex pointed; midrib raised above; lateral veins invisible to very faint, numerous,
close; petiole 0.51 cm long. Flowers: petals white with pink base; stamens purplish; ovary
ovoid, style short, purplish, stigma capitate. Fruits ellipsoid, c. 8.5 x 3 mm.
268
269
Distribution. Endemic to western Sarawak.

Ecology. Lowlands from sea-level to 600 m, mainly in kerangas forest.
270
OLACACEAE
Lesmy Tipot
Kepong, Malaysia
Hooker f., Fl. Brit. Ind. 1 (1875) 572 (under Olacineae); King, J. As. Soc. Beng. 64, 2 (1895) 108;
Merrill, EB (1921) 242; Ridley, FMP 1 (1922) 419; Sleumer in Engler & Prantl, Pfl. Fam. 2, 16b
(1935) 5, Blumea 26 (1980) 145, FM 1, 10 (1984) 1; Masamune, EPB (1942) 258; Backer &
Bakhuizen f., FJ 2 (1965) 63; Smythies, CST (1965) 113; Whitmore, TFM 2 (1973) 299; Keng,
OFMSP (1978) 212; Anderson, CLTS (1980) 284; Cockburn, TS 2 (1980) 65; Corner, WSTM 2
(1988) 600; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 279.
Trees, shrubs or climbers. Leaves spirally arranged, rarely distichous, simple, entire,
stalked, without stipules, when dry often with parchment-like and/or finely tuberculate
surface, pinnately veined with the lateral veins rather distant. Inflorescences axillary
cymes, racemes or branched spikes. Flowers bisexual, rarely unisexual, mostly radially
symmetrical, rarely heterostylus; calyx often cup-like, shortly 37-lobed or dentate, sometimes
accrescent; petals 37, free or joined below; stamens as many or twice the number of
petals; disc present; ovary mostly superior, rarely inferior, 35-loculed, ovule solitary in
each locule; style usually 1, stigma 35-lobed. Fruit a drupe; exocarp thin or fleshy; endocarp crustaceous to woody. Seeds one per fruit, with thin testa and abundant endosperm.
Distribution. Pantropical, 27 genera with c. 170 species. In Sabah and Sarawak, represented
by 8 genera with 9 species of trees, shrubs and climbers.
Ecology. Found in a wide range of habitats, from open and sandy seashores to secondary
and primary mixed dipterocarp forests. Occasionally in submontane forests.
Uses. Most species are small to medium-sized trees and are, therefore, commercially not
important, except for Scorodocarpus borneensis which produces a dark red timber. Species
of Anacolosa, Strombosia and Ochanostachys are occasionally used in local construction
works. The wood of Ximenia americana can be used as a substitute for sandal wood,
whereas the fruit kernel is said to have medicinal properties. Fruits of Scorodocarpus
borneensis are edible and the young leaves are taken as vegetable.
Taxonomy. Engler (Syllabus, 1924) recognised Olacaceae as the most primitive family in
the order Santalales. Ridley l.c., following Hooker f. l.c., on the other hand, treated
Olacaceae together with Icacinaceae under the order Olacineae. Hutchinson (Fam. Fl. Pl. 1
(1964) 329) placed Olacaceae together with Opiliaceae and a few other families in the
Olacales, an order comparatively more primitive than the Santales. Keng l.c., Sleumer l.c.
and Whitmore l.c. followed Engler's classification, a stand accepted in the present
account.The Santalales consists of a small group of families showing a tendency toward
271
272
OLACACEAE (LESMY)
hemiparasitism, reduction of the flowers and adaptation to the climbing and epiphytic habit.
In this account, the nomenclature follows that of Sleumer's revision (1984), in which he has
considerably reduced the number of species of Strombosia and Olax recognised by earlier
workers.
Key to genera
1.
Slender woody climbers, often with axillary tendrils......................................................

Erythropalum Blume
Bijdr. (1826) 921; Merrill, PEB (1929) 58; Masamune l.c. 258; Sleumer l.c. (1980) 151, l.c.
(1984) 17.
One species, E. scandens Blume, distributed from the Himalayas to Assam, Bengal,
Burma, Indo-China, Thailand and Malesia. In Sabah, recorded from Kinabatangan,
Lahad Datu, Ranau, Tambunan, Tawau, Tenom and Mt. Kinabalu at c. 1200 m. In
Sarawak, in Melinau and Bukit Jebong in Bau district.
Fruit drupaceous, ellipsoid, ripening reddish, entirely enclosed by the persistent
calyx which finally splits from top downwards into 36 reflexed segments. In open
areas and hillsides.
Trees, shrubs or scandent shrubs, without axillary tendrils...........................................2
2.
Leaves usually mucronate at apex. Twigs armed with thorns.........................................

Ximenia L.
Sp. Pl. (1753) 1193; Masamune l.c. 259; Sleumer l.c. (1980) 166, l.c. (1984) 10.
Eight species in the tropics and subtropics. 1 species, X. americana L., in Borneo
and has been recorded from Teluk Tambak, Santubong near Kuching, Lubok Jayau
and Tatau near Bintulu in Sarawak; and from Lahad Datu, Sandakan and Semporna
in Sabah.
Low-branching shrub, deciduous in dry season. Usually on sandy seashores and in
dry forest fringes, sometimes on stony ground.
Leaves not mucronate. Twigs without thorns............................................................3
3.
Leaves distichous or subdistichous. Calyx well-developed, enclosing fruit or flat and

fleshy at maturity........................................................................................................4
Leaves not distichous. Calyx not developed as above....................................................5
4.
Leaves elliptic, both surfaces parchment-like. Fruit seated on an enlarged flattish

calyx.......................................................................................................2. Harmandia
Leaves ovate, elliptic to oblong, surface not parchment-like. Fruit enclosed in the
enlarged calyx...............................................................................................................
Olax L.
Sp. Pl. (1753) 34; Merrill l.c. (1921) 242; Masamune l.c. 259; Sleumer l.c. (1984) 8.
About 40 species, Old World Tropics. 1 species, O. imbricata Roxb., distributed in

India, Sumatra, Peninsular Malaysia, Borneo, Java, New Guinea and Solomon
Islands. In Sabah, recorded from Lamag, Mt. Lotung, Bongawan, Papar, Sipitang,
and Sandakan. In Sarawak, in Lawas, Limbang, Sibu, Kapit and Kuching.
273
Scandent shrub or climber. On open sandy ground including coasted areas, and
occasionally on rocky ground along streamsides.
5.
Leaves usually with fine pellucid-dots visible against strong light................................6

Leaves not so.............................................................................................................7
6.
Young twigs zig-zag. Petals entirely free.............................................5. Strombosia

Young twigs not zig-zag. Petals fused in the lower half............................1. Anacolosa
7.
All parts reeking of garlic smell when fresh. Leaves broadly elliptic, c. 20 cm long.
Bark fissured or thinly flaky, inner bark without any sap..................4. Scorodocarpus
Garlic smell absent. Leaves elliptic, rather small. Bark dippled, distantly scaly, inner
bark with a slight white sap...............................................................3. Ochanostachys
1. ANACOLOSA Blume
(Greek, anakolos = knotted; the calyx-cup rim)
Mus. Bot. Lugd. Bat. 1 (1850) 250, t. 46; Merrill l.c. (1921) 242; Ridley l.c. (1922) 424; Sleumer in
Engler & Prantl l.c. (1935) 20, l.c. (1980) 146, l.c. (1984) 23; Masamune l.c. 258; Whitmore l.c. 300;
Anderson l.c. 283; Whitmore, Tantra & Sutisna l.c. 279.
Trees or shrubs. Leaves spiral, pinnately veined. Flowers bisexual, in axillary cymes or
fascicles; calyx cup-shaped, (5)6(7)-dentate or minutely toothed; petals (5)6(7), inserted on the margin of the cupular disc, fused in the lower part, fleshy and concave below;
stamens (5)6(7), filaments short, anthers ovoid; disc hypogenous, fused with the ovarywall, 6-denticulate or furrowed; ovary superior, incompletely 2(3)-celled below, 1-celled
above, style short and thickened at base; ovules 2(3), unitegmic, pendant, placentation
central. Fruit a drupe, surrounded by the enlarged disc, tipped by the remains of the style,
seated on the small persistent calyx; pericarp thin, fleshy; endocarp thin, crustaceous. Seeds:
embryo minute; endosperm starchy and oily.
Distribution. 15 species, in the Old World tropics, of which 3 species occur in Malesia. In
Sabah and Sarawak, one species.
Anacolosa frutescens (Blume) Blume
Fig. 1.
(Latin, frutex = a shrub; the habit)

l.c. (1850) 251, t. 46; Masamune l.c. 258; Sleumer l.c. (1980) 146, l.c. (1984) 25; Anderson l.c. 283;
Whitmore, Tantra & Sutisna l.c. 279. Basionym: Stemonurus frutescens Blume l.c. (1826) 649. Type:
Blume "2168", Java (holotype L; isotypes P, U). Synonyms: Anacolosa heptandra Maingay ex Mast.
in Hooker f. l.c. 581; A. arborea Koord. & Valeton, Bull. Inst. Bot. Buitz. 2 (1899) 9; A. luzoniensis
Merr., Philip. J. Sc. 4, Bot. (1909) 253; Anacolosa sp. Merr. l.c. (1921) 242; Salacia bartletti Ridl.,
Kew Bull. (1938) 239.
Erect shrub or small slender tree, to 30 m tall and 30 cm diameter. Bark smooth to papery
or mottled, greenish grey; inner bark reddish. Sapwood yellow to pale red. Twigs greyish
brown or whitish. Leaves very variable in shape, from elliptic to elliptic-oblong or
274
OLACACEAE (LESMY)
lanceolate, 918 x 69.5 cm, papery to leathery, glossy above when fresh, brownish and
dull in dry state, usually with tiny warts or tubercles on both surfaces, or with fine pelluciddots visible against strong light; base cuneate, slightly asymmetric, margin entire, apex
acuminate, blunt or rounded; lateral veins 57 pairs, ascending, rather distant, lowest pair
close to the base, faint on upper surface, conspicuous and slightly raised below; petiole
stout, 512 mm long. Flowers in axillary clusters of 515, pedicels 25 mm; calyx shortly
57-lobed, glabrous or pale rusty-puberulous, c. 3 mm in diameter; petals 6, ovatelanceolate, glabrous or rarely puberulous outside, 23 x 11.5 mm; stamens 6, anthers hairy;
ovary with 23 pendulous ovules. Fruits obovoid to oblong, with persistent calyx, green,
ripening yellow to orange, c. 1.2 x 0.8 cm. Seeds: endosperm copious.
Vernacular names. Sabahsalangugapit (Dusun, Tawau). Sarawakbelian landak (Malay,
Iban), jerit (Iban), ladit (Melanau).
Distribution. Burma, Andaman and Nicobar Is., Thailand, Sumatra, Peninsular Malaysia,
Java, Borneo, Philippines, Celebes and the Moluccas. In Sabah, recorded from Kalabakan,
Keningau, Kota Belud, Lahad Datu, Ranau, Sandakan, Sepulut and Tawau. In Sarawak,
known from Kuching, Sibu, Sarikei, Kapit, Bintulu, Miri to Baram.
Ecology. Common on hillslopes of primary mixed dipterocarp forests on sandy humult
ultisols, sometimes in heath and peat swamp forests. Once recorded from limestone hill, and
once from submontane forest at about 1100 m.
Uses. The wood is pale reddish brown, considerably hard and heavy, and sometimes used
locally for house-posts though not very durable.
2. HARMANDIA Pierre ex Baill.

(Jules Harmans, 18451921, French colonial officer)
Bull. Linn. Soc. Paris 2 (1889) 770; Sleumer l.c. (1935) 30, l.c. (1980), l.c. (1984) 9; Whitmore l.c.
Trees monoecious. Leaves distichous, pinnately veined. Inflorescences short, axillary

racemes. Flowers unisexual; calyx at anthesis disc-form, shortly 4-dented, much accrescent
at maturity forming a frill below the fruit; petals 4 in male, 68 in female flowers, connate
to an urceolate tube at base, free at the upper part; disc extra-staminal, annular. Male
flowers: stamens 4, epipetalous, filaments fused into a tube with the free anthers on top;
ovary rudimentary. Female flowers: staminodal tube without anthers; ovary pyramidal, 1celled, with 2 pendant ovules, placentation basal; style short-conical, stigma 3, sessile.
Fruit a drupe, concrescent with the much enlarged calyx below; pericarp fleshy; endocarp
thin and woody. Seeds: endosperm oily; embryo excentric.
Distribution. Monotypic; Indo-China and W Malesia (Sumatra, Peninsular Malaysia and
Borneo).
Ecology. Usually found in lowland forests.
275
276
OLACACEAE (LESMY)
Harmandia mekongensis Pierre ex Baill.
Fig. 2.
(of Mekong River in Laos)

l.c. 770; Sleumer l.c. (1980) 153, l.c. (1984) 9. Type: Harmand 1322, SE Laos (holotype P; isotypes
BM, K, L, SING). Synonym: Harmandia kunstleri King l.c. 100.
Medium-sized tree to 30 m tall, 30 cm diameter. Bark thin, pale brown to whitish, flaky;
inner bark pale yellow, granular. Sapwood pale yellow. Twigs slender, dark brown to black,
striate and slightly zig-zag. Leaves elliptic, 78.5 x 34 cm, subcoriaceous, parchment-like
especially on lower surface, deep green when fresh; base cuneate to slightly rounded,
margin recurved, apex pointed; lateral veins 46 pairs, rather inconspicuous on both
surfaces; reticulations very faint; petiole 810 mm long. Flowers pale green to whitish;
calyx cupular at bud stage, accrescent in fruit; petals c. 2 mm, connate below, forming an
urceolate corolla. Male flowers: stamens 4, filaments connate to a fleshy tube, c. 1.5 mm;
anthers cordate, c. 0.5 mm; pistillodes present. Female flowers: staminodal tube without
anthers; ovary conical, tapering to a short style; stigmas 3, sessile. Fruits ovoid-ellipsoid,
green, maturing black, c. 3 x 1.5 cm, with an enlarged pale red and fleshy calyx below.
Seeds: pericarp fleshy, c. 0.5 mm thick; endocarp woody, c. 0.5 mm thick.
Distribution. Indo-China (Laos, Annam) and W Malesia (Sumatra, Peninsular Malaysia
and Borneo). In Sabah very uncommon, represented by only two collections from Keningau
(SAN 107911) and Nabawan (SAN 118776). Not yet recorded from Sarawak.
Ecology. In lowland and hill primary and disturbed forests to about 300 m.
3. OCHANOSTACHYS Mast.
(Greek, okanon = shield strap, stachus = spike; the inflorescence)
in Hooker f. l.c. 576; Merrill l.c. (1921) 242, l.c. (1929) 58; Ridley l.c. (1922) 422; Masamune l.c. 259;
Whitmore l.c. 302; Anderson l.c. 284; Sleumer l.c. (1980) 153, l.c. (1984) 12; Whitmore, Tantra &
Sutisna l.c. 279. Synonym: Petalinia Becc., Malesia 1 (1883) 257.
Trees. Leaves spiral. Flowers bisexual, on simple or branched axillary spikes; calyx small,
cup-shaped, 45-lobed, not accrescent; petals 45, almost free; stamens 815, epipetalous,
(1)2(3) on each petal, filaments joined at base, anthers globular to subglobular; disc
hypogynous, fleshy, rather inconspicuous; ovary superior, incompletely 34-chambered,
style short, stigma 3-lobed; ovules bitegmic, pendulous, placentation free basal. Fruit a
drupe; pericarp thin; endocarp woody. Seeds: endosperm starchy; embryo small, apical.
Distribution. Monotypic; confined to W Malesian region (Sumatra, Bangka, Lingga Island,
Peninsular Malaysia, and Borneo).
Ecology. Lowland rain forest.
277
Ochanostachys amentacea Mast.
Fig. 3.
(Latin, amentum = a catkin; the inflorescence)

in Hooker f. l.c. 577; Merrill, l.c. (1921) 242, l.c. (1929) 58; Ridley, Kew Bull. (1931) 35, incl. var.
rufa Stapf ex Ridl.; Masamune l.c. 259; Browne, FTSB (1955) 251; Smythies l.c. 113; Burgess, TBS
(1966) 420; Whitmore l.c. 302; Anderson l.c. 284; Cockburn l.c. 67; Sleumer l.c. (1980) 153, l.c.
(1984) 14; Whitmore, Tantra & Sutisna l.c. 278. Type: Maingay 384, Malacca (lectotype K;
isolectotype P). Synonyms: Petalinia bancana Becc. l.c. 258; Ochanostachys bancana (Becc.)
Valeton, Crit. Overz. Olacin. (1886) 104.
Small to medium-sized tree, to 30 m tall, 70 cm diameter; bole straight to (often) rather

poorly formed, at base shortly fluted or buttressed. Bark grey-brown to reddish brown,
shedding off into thin, irregular scales, exposing lighter coloured patches, giving a dippled
appearance; inner bark finely fibrous, yellowish brown, with discrete droplets of white
latex. Sapwood hard, brownish yellow. Leaves usually glabrous, thinly leathery, ovateelliptic to elliptic-oblong, 613 x 37 cm, green and shiny above, yellowish green beneath
when fresh, rather dull and brownish when dry, usually with blackish dots on both surfaces;
base cuneate or rounded, margin entire, apex acute to acuminate; lateral veins 46 pairs,
rather distant, sunken above prominent below; intercostal veins scalariform, very fine and
rather faint; stalk slender, 1.52.5 cm, with a faint, darker, 5 mm, apical knee.
Inflorescences simple or branched spikes to 12 cm long. Flowers green, in opposite, wellspaced clusters; calyx 45-toothed, c. 1 mm; petals (3)4(5), ovate to ovate-oblong, c. 2.5
x 1.5 mm; ovary depressed ovoid, striate, glabrous, style short, cylindrical. Fruits
subglobose, to 2 cm across, on a slender peduncle of c. 2 mm; pericarp thin, exuding a milky
gum; endocarp woody. Seeds subglobose.
Vernacular names. Sabahtanggal (Dusun). Sarawakimah (Bidayuh Bau), petaling
(Malay), sagad berauh (Murud), santikal (Iban).
Distribution. Sumatra, Bangka, Peninsular Malaysia and Borneo. Common and widespread
throughout Sabah and Sarawak.
Ecology. Main canopy tree in primary and secondary mixed dipterocarp forest, on hillsides
and ridges on loamy usually clay-rich soils, to 800 m, occasionally in heath forest.
Uses. Produces a yellowish brown, hard, heavy, and reasonably durable wood, useful for
house construction and logging railways.
4. SCORODOCARPUS Becc.
(Greek, skorodon = garlic, karpos = fruit; the strong garlic smell in the fruit)
Nuov. Giorn. Bot. Ital. 9 (1877) 274, t.11, f.1217; Merrill l.c. (1921) 242; Ridley l.c. (1922) 424;
Masamune l.c. 259; Smythies l.c. 113; Whitmore l.c. 303; Anderson l.c. 284; Cockburn l.c. 67;
Sleumer l.c. (1980) 160, l.c. (1984) 15; Whitmore, Tantra & Sutisna l.c. 279.
Garlic-smelling tree, with straight bole, without buttresses. Leaves spiral. Flowers bisexual, in
short racemes; calyx small, cup-shaped, narrowly 45-toothed, not enlarged in fruit; petals
278
OLACACEAE (LESMY)
279
280
OLACACEAE (LESMY)
45, hairy within; stamens 810, in pairs, joined halfway down, and attached to the base
of petal; ovary superior, imperfectly 34-celled, style elongate conical, stigma 34-lobed,
ovule 1 in each cell, pendant, placentation free. Fruit a drupe; pericarp thin, fleshy;
endocarp woody. Seeds: endosperm fleshy, containing starch and tannin.
Distribution. Monotypic; S Thailand, Sumatra, Lingga Island, Peninsular Malaysia and
Borneo.
Ecology. Common in lowland forest.
Scorodocarpus borneensis (Baill.) Becc.
Fig. 4.
(of Borneo)
l.c. 274, t. 11, f. 1217; Merrill l.c. (1921) 242; Ridley l.c. (1922) 424; Masamune l.c. 259; Brown l.c.
280; Smythies l.c. 422; Whitmore l.c. 303; Anderson l.c. 284; Cockburn l.c. 67; Sleumer l.c. (1980)
160, l.c. (1984) 15; Whitmore, Tantra & Sutisna l.c. 279. Basionym: Ximenia borneensis Baill.,
Andansonia 11 (1874) 271. Type: Beccari PB 1581, Sarawak, Mt. Matang (holotype FI; isotypes P,
W).
Medium-sized to large tree, to 40 m tall, 80 cm diameter; all parts with garlic smell
especially when crushed or cut; crown small, dense; bole straight, to 25 m, occasionally
with small, low buttresses. Bark grey to dark red or brown, fissured and thinly rectangularly
flaky; inner bark purplish red, inwards with coarse orange flecks. Sapwood hard, yellow to
reddish brown. Leaves coriaceous, shiny green above when fresh, dull olive-green when
dry, glabrous, elliptic, 1022 x 49 cm; base cuneate to rounded, margin entire, apex
acuminate; lateral veins 56 pairs, distant, curving upwards towards margin, flat above, very
prominent beneath; reticulation rather faint; petiole 1.52 cm, thickened distally or almost at
the leaf-base. Inflorescences racemose, to 4 cm, rusty to greyish puberulous. Flowers
white, c. 1.5 cm long, solitary or grouped in clusters of 23 along the rachis; pedicels c. 1.5
2 mm; calyx small with wavy edges; petals white or creamy white, 810 x 2 mm, woolly
inside, reflexed; anthers 34 mm; ovary yellowish green, tapering toward the thickish white
style. Fruits globose, green, c. 5 cm across; peduncle 1 cm; pericarp thin and fleshy;
endocarp woody with numerous vertical fibre-like hard strands. Seeds subglobose.
Vernacular names. Sabahbawang hutan (Malay). Sarawaksagan-berauh (Murut),
sindok, sindu (Iban), troduh (Bidayuh Bau).
Distribution. Peninsular Thailand, Sumatra, Lingga Is., Peninsular Malaysia, Borneo. In
Sabah and Sarawak widespread. Also occurs in Brunei and Kalimantan.
Ecology. Commonly found on slopes in primary and secondary mixed dipterocarp forest on
clay loam soils, and occasionally in seasonally flooded alluvial forest.
Silviculture. Bawang hutan requires well-drained soils for good growth under normal
conditions. It is evergreen but slow-growing. Fruiting usually occurs during June
September.
281
282
OLACACEAE (LESMY)
Uses. Produces a dark brown, moderately durable, medium hardwood timber useful for
piling, making bridges and house-posts. The fruits are boiled and eaten by many
communities in Borneo (S. 36079 and SAN 10463). The young leaves are also cooked as a
vegetable in Sarawak.
5. STROMBOSIA Blume
(Greek, strombos = conical or pear-shaped; the fruit)
l.c. (1826) 1154; Merrill l.c. (1921) 242; Ridley l.c. 425; Sleumer in Engler & Prantl l.c. (1935) 21, l.c.
(1980) 163, l.c. (1984) 19; Masamune l.c. 259; Whitmore l.c. 305; Anderson l.c. 284; Whitmore,
Tantra & Sutisna l.c. 280. Synonym: Lavallea Baill., Adansonia 2 (1862) 361.
Shrubs or trees. Young twigs distinctly zig-zag, smooth, light coloured, old ones often black.
Leaves spirally arranged, pinnately veined, glabrous; lateral veins rather distant, ascending.
Flowers bisexual, in sessile or shortly stalked cymes or clusters; calyx cup-shaped, 5-lobed,
accrescent and adnate to the pericarp almost to the top of the mature fruit; petals 5, free,
inner parts hairy; stamens 5, adnate to the petals, filaments flat, with simple hairs, anthers
dorsifixed; disc prominent, 5-lobed; ovary partly sunken into the receptacle, almost entirely
covered by the 5-lobed fleshy disc, 35-chambered; ovules 35 per locule, anatropous,
placentation free central; style short, stigmas subglobular, obscurely 35-lobed. Fruits
drupaceous, crowned by the persistent calyx and style base; pericarp thin-fleshy; mesocarp
crustaceous or woody. Seeds: embryo small, apical; endosperm fleshy and oily.
Distribution. About 12 species, c. 9 of which are confined to tropical Africa; the rest occur
in India, Sri Lanka, Burma, Thailand, and Malesia (Sumatra, Peninsular Malaysia, Java,
Borneo, the Philippines and N Moluccas). In Sabah and Sarawak, represented by 2 species.
Ecology. Generally lowland forest.
Key to Strombosia species

Leaves smooth on upper surface, not pellucid-punctulate; intercostal veins usually visible.
Flowers in cymose inflorescences. Fruits with truncate apex, apiculate by the persistent,
hard style-base................................................................................................1. S. javanica
Leaves parchment-like on both surfaces, distinctly pellucid-punctulate; intercostal veins
seldom visible. Flowers in clusters. Fruits with obtuse-rounded apex, remains of style-base
inconspicuous................................................................................................2. S. ceylanica
1. Strombosia javanica Blume
Fig. 5.
(of Java)
l.c. (1826) 1155; Merrill l.c. (1921) 242; Ridley l.c. 425; Masamune l.c. 259; Browne l.c. 282;
Whitmore l.c. 306; Sleumer l.c. (1980) 164, l.c. (1984) 21; Whitmore, Tantra & Sutisna l.c. 280.
Type: Blume, s.n., West Java, Mt. Salak (holotype L; isotypes K, P).
283
Tree to 40 m tall, 100 cm diameter; bole straight, often with knobs; crown dense and
narrow. Bark yellowish grey, shallowly irregularly fissured; inner bark pinkish, fibrous.
Sapwood pale yellow. Leaves elliptic-oblong, (10)1218(24) x 48 cm, thick
membranous to subcoriaceous, drying yellowish olivaceous or brownish, smooth and shiny
above, without pellucid-dots; base obtuse to rounded, margin entire, apex subacuminate;
lateral veins 57 pairs, conspicuous and raised below, flat above; petiole slightly swollen
distally, 1.52 cm. Flowers solitary or in 37-flowered fascicles; calyx disc-shaped, 45
angular, teeth obscure, c. 3 mm; petals greenish white, reflexed at apex, densely hairy
inside, glabrous outside, ovate-lanceolate, 810 x 23 mm; filaments ciliate at the apex,
anthers ovate-oblong, c. 0.5 mm; ovary deeply 5-furrowed lengthwise; style columnar;
stigma obscurely 5-lobed. Fruits ovoid, 23.5 x 1.52 cm, green, with thin and fleshy
pericarp and woody endocarp, apex truncate, crowned by the remains of the calyx, disc and
style.
Vernacular name. Sarawakbelian landak (Iban).

Distribution. Burma, S Thailand, Sumatra, Peninsular Malaysia, Java and Borneo
(Sarawak, Brunei and Kalimantan). In Sarawak uncommon (e.g., S. 34250, S. 35255 from
Mt. Gading, Lundu, and S. 1476 from Marudi). Not yet recorded in Sabah.
Ecology. Scattered but locally common in mixed dipterocarp forest and secondary forest to
600 m, on clay-rich fertile soils.
Uses. Produces a moderately hard, durable, light yellowish timber, used locally for house
construction.
2. Strombosia ceylanica Gardner

(of Ceylon)
Calc. J. Nat. Hist. 6 (1845) 350; Sleumer l.c. (1980) 165, l.c. (1984) 22; Whitmore, Tantra & Sutisna
l.c. 280. Type: Gardner, s.n., Ceylon, Hantane (holotype K; isotype BM). Synonyms: S. lucida
Teijsm. & Binn. ex Valeton l.c. 93; Anacolosa maingayi Mast. in Hooker f. l.c. 580; S. maingayi
(Mast.) Whitmore, Gard. Bull. Sing. 26 (1973) 285; S. rotundifolia King l.c. 102; S. multiflora King
l.c. 102; S. latifolia Stapf, Kew Bull. (1906) 71, Merrill l.c. (1921) 242, Masamune l.c. 259.
Small to medium-sized tree, 40 m tall, 60 cm diameter; crown dense; bole straight, closely
branched, sometimes buttressed. Bark grey to brown, peeling off in scroll-shaped patches
or scales; inner bark reddish brown. Sapwood hard, orange-brown. Leaves subcoriaceous,
elliptic to ovate-oblong, 815(25) x 37(11) cm, shiny above when fresh, drying dull or
greenish brown or glaucous green, conspicuously parchment-like, distinctly pellucidpunctulate; base cuneate to rounded, margin entire, apex shortly acuminate; lateral veins 5
8 pairs, ascending, curved, faint above, slightly raised beneath; intercostal veins invisible.
Flowers in clusters or fascicles of 36, almost sessile or with 12 mm pedicels, arising from
small woody-warts; calyx 5-lobed, lobes ovate, obtuse, ciliate; petals 5, greenish white,
oblong, hairy inside, glabrous outside; anthers ovate; ovary semi-inferior, disc conical,
faintly 5-lobed; style filiform, 24 mm. Fruits roundish to subglobose, c. 22.5 cm across,
attenuate at base at maturity, apex with the persistent tiny style; pericarp thinly fleshy, pink
to purple, rugose or tuberculate; endocarp thinly woody. Seeds c. 1.2 cm across.
284
OLACACEAE (LESMY)
Vernacular name. Sarawakbelian landak (Iban, Malay, Melanau).

Distribution. SW India, Sri Lanka, Sumatra, Peninsular Malaysia, Java, and Borneo.
Widespread in Sarawak, from Limbang, Baram, Miri, Marudi, Bintulu, Kapit, Sri Aman to
Kuching. In Sabah, recorded from Beaufort, Beluran, Lahad Datu, Sandakan and Tenom.
Ecology. In lowland mixed dipterocarp and secondary forest on leached sandy humult
soils; also collected in hill forest up to 1000 m.
Uses. The timber is hard, heavy but only moderately durable. The wood is yellowish brown,
having rather fine and even texture, useful for house construction.
285
OXALIDACEAE
R.C.K. Chung
Kepong, Malaysia
Merrill, EB (1921) 311, PEB (1929) 111; Ridley, FMP 1 (1922) 329; Masamune, EPB (1942) 355;
Backer & Bakhuizen f., FJ 1 (1963) 244; Veldkamp, FM I,7 (1971) 151; Cockburn, TFM 1 (1972)
347, TS 2 (1980) 69; Anderson, CLTS (1980) 286; Corner, WSTM 2 (1988) 605; Whitmore, Tantra &
Sutisna, CLK 2, 1 (1990) 282.
Trees, shrubs, climbers or herbs. Leaves alternate or spiral, compound with a terminal
leaflet or simple, pinnately veined, stipules sometimes present, stalk usually with a basal
joint. Inflorescences axillary, pseudoterminal or on the branches, racemes, panicles or
cymes, 1- to many-flowered. Flowers bisexual, regular (actinomorphic), 5-merous, heterotristylous (with short, medium and long-styled forms), heterodistylous (with short and longstyled forms) or homostylous (with all styles of similar length), stalk jointed; sepals 5, free or
fused at the base, persistent; petals 5, clawed, often falling off early; stamens 10, arranged
in two whorls, 5 opposite and 5 alternate with petals, connate at the base into an annulus,
persistent, of which 5 are sometimes sterile (staminodes); anther dorsifixed, versatile,
dehiscing extrorsely by longitudinal slits; disc absent; ovary superior, 5-locular; ovules
anatropous, bitegmic, tenuinucellate (with a thin nucellus, Averrhoa) or crassi-nucellate
(with a thick nucellus, Oxalis and Biophytum), 1 or more per locule; placentation axile;
styles 5, of the same or different length, terminal, free, erect or recurved. Fruit a dry 5valved capsule or a berry. Seeds 1 to many per fruit, sometimes arillate (except for species
of Sarcotheca and Averrhoa bilimbi); endosperm fleshy; embryo straight.
Distribution. 6 genera with about 890 species, mostly in tropical or subtropical areas. In
Malesia, 5 genera with 29 species, of which 14 are endemic. In Sabah and Sarawak,
represented by 5 genera with 12 species, of which only 5 are wild and 2 are cultivated trees.
Ecology. Found in a wide variety of habitats, from lowland primary and secondary forests
to montane forests, to about 2200 m.
Uses. Species of Averrhoa are cultivated for its edible fruit, and that of Oxalis are cultivated
as ornamentals. The wood of the ligneous Oxalidaceae is not utilised as timber. In
Peninsular Malaysia, the fruits of Sarcotheca are edible.
Taxonomy. Hutchinson (Fam. Fl. Pl. 1 (1959) 356, 497; Gen. Fl. Pl. 2 (1967) 610)
considered the herbaceous taxa as the Oxalidaceae sensu stricto (in the Geraniales), and
accommodated the arboreous taxa in the Lepidobotryaceae (Malpighiales) and Averrhoaceae
(Rutales) respectively. Recent work indicates that Sarcotheca and Dapania (in the
Lepidobotryaceae) should be associated with Averrhoa rather than with Lepidobotrys.
Therefore, the Averrhoaceae would be better retained as a group in the Oxalidaceae. Species
287
of Sarcotheca have sometimes been confused with that of Rourea (Connaraceae). However,
Rourea differs from Sarcotheca in having free carpels, 2-collateral ovules, seeds with an
aril, and a dry, indehiscent, 1-celled and 1-seeded fruit.
Key to genera
1.
Herbs...........................................................................................................................2
Climbers or trees........................................................................................................3
2.
Leaves trifoliolate..........................................................................................................
Oxalis L.
Sp. Pl. (1753) 433, Gen. Pl. ed. 5 (1754) 198; Ridley l.c. 330; Backer & Bakhuizen f. l.c.
245; Veldkamp l.c. 153.
7 species in Malesia; 2 species (O. corniculata and O. corymbosa) in Sabah and

Sarawak.
Herbs. Leaves trifoliolate; leaflets usually obcordate. Inflorescences cymose to
pseudoumbellate, one- to many-flowered. Sepals shortly connate at base; petals
connivent above the claw. Capsules dehiscing loculicidally, the valves remaining
attached to the central axis. Seeds usually few, ejaculatory apparatus and testa
smooth.
Leaves pinnate, with many leaflets.................................................................................
Biophytum DC.
Prod. 1 (1824) 689; Edgewick & Hooker f., Fl. Br. Ind. 1 (1874) 436; Ridley l.c. 330; van
Steenis, Bull. Jard. Bot. Btzg 3, 18 (1950) 449; Backer & Bakhuizen f. l.c. 246; Veldkamp
l.c. 159.
7 species in Malesia; 1 species (B. sensitivum) in Sabah & Sarawak.

Herbs or usually sympodially branched dwarf shrubs. Stipules bristle-like
(setaceous). Leaves paripinnate, in tufts at the end of the stems or branches; leaflets
opposite. Flowers terminal, usually in pedunculate, bracteate pseudo-umbels,
heterodistylous, heterotristylous or homostylous; sepals free; petals twist-ed,
connivent above the claw. Capsules split loculicidally to the base, forming a 5rayed star-shaped fruit without leaving a columella. Seeds 16 per cell, ejaculatory
apparatus and aril white, thin.
3.
Climbers. Leaves unifoliolate.........................................................................................

Dapania Korth.
Ned. Kruidk. Arch. 3 (1854) 381; 2; Ridley l.c. 334; Veldkamp, Blumea 15 (1967) 523, l.c.
(1971) 166.
2 species in Malesia (Sumatra, Peninsular Malaysia, Borneo) with one species, D.

grandifolia endemic to NE Sabah and SE Kalimantan.
Common woody climbers in primary lowland forests. Leaves alternate, estipulate;
stalks jointed in the middle. Inflorescences racemose, on the branches or axillary,
solitary to fascicled. Flowers androdioecious; sepals 5, connate at the lower half,
margins ciliate; petals 5, free, imbricate. Capsules fleshy, loculicidally dehiscent.
Seeds hard; aril enveloping the seed.
Trees. Leaves pinnate, trifoliolate, or unifoliolate.........................................................4
4.
Leaves imparipinnate with 319 pairs of leaflets. Stalks of leaf and leaflet not jointed...
Averrhoa L.
288
OXALIDACEAE (CHUNG)
l.c. (1753) 428, l.c (1754) 196; Ridley l.c. 331; Backer & Bakhuizen f. l.c. 247; Veldkamp
l.c. (1971) 174; Corner l.c. 605; Whitmore, Tantra & Sutisna l.c. 282.
Two species (A. bilimbi and A. carambola) of unknown origin, currently cultivated
throughout the humid tropics and subtropics.
Shrubs or small trees, cultivated as fruit trees. Stipules absent. Leaves spirally
arranged or clustered at the end of branches, imparipinnate; leaflets entire, subopposite, subsessile, 3 or more pairs. Inflorescences panicles, axillary or borne on
the trunk. Flowers heterodistylous or heterotristylous; sepals shortly connate at
base; petals twisted, free or connivent above the claw, pink to dark red; stamens all
fertile or 5 fertile and 5 sterile; ovary syncarpous with 37 superimposed
pendulous ovules in each of the 5 locules. Fruit a large juicy berry. Seeds
numerous, elliptic, flat; aril, if present, fleshy, attached to the entire adaxial placenta,
2-lobed, and enveloping the seed; cotyledons thin, flat; germination epigeal.
Both species prefer a climate with a dry season, but also do well in wetter climates.
A. carambola has been cultivated successfully in frost-free subtropics up to 30S in
Australia and 32N in Israel. For healthy growth, both species require adequate
moisture and well-drained soils. In Malaysia, A. carambola has been planted and
thrives well in well-drained peat soils. Vernacular name: belimbing (Malay).
Leaves unifoliolate or trifoliolate. Stalks of leaf and leaflet jointed...............Sarcotheca
SARCOTHECA Blume
(Greek, sarcos = fleshy; theca = container; the fleshy fruit)
tabarus (Dusun, Sabah), piang (Iban, Sarawak)
Mus. Bot. Lugd. Bat. 1 (1850) 241; Ridley l.c. 332; Merrill l.c. (1929) 111; Veldkamp l.c. (1967) 527,
l.c. (1971) 168; Cockburn l.c. (1972) 348, l.c. (1980) 70; Anderson l.c. 286; Whitmore, Tantra &
Sutisna l.c. 282; Ng, Mal. For. Rec. 34 (1992) 470. Synonyms: Roucheria Miq. l.c. (1859) 136;
Connaropsis Planch. ex Hook. f., Trans. Linn. Soc. 23 (1860) 166.
Trees or shrubs; bole deeply fluted. Bark reddish or reddish brown, scaly or dippled; inner
bark reddish. Sapwood pale. Stipules absent. Leaves alternate, unifoliolate or trifoliolate;
margin entire; leaf-stalks jointed, proximal parts and leaflet stalks swollen and wrinkled.
Inflorescences axillary or terminal panicles, one to several together. Flowers heterodistylous, arranged in more or less stalked cymes, scattered along a simple or sparsely
branched rachis; cymes subtended by small caducous bracts; sepals unequal, persistent
(except in S. diversifolia), imbricate, shortly connate at base, abaxial surface strigose with
appressed hairs; petals twisted, imbricate, free at base, usually fused above and falling off
as a single unit, adaxial surface with minute papillae in the upper half; stamens all fertile;
ovary glabrous to hairy, ovules 2 in each locule. Fruits fleshy, red at least when dry,
usually 5-ridged. Seeds flat, without aril, testa reddish, smooth to wrinkled, hard; cotyledons
thin, flat; germination epigeal.
289
Distribution. 11 species in W Malesia (Sumatra, Peninsular Malaysia, Borneo, Celebes). In

Sabah and Sarawak, 5 species are recorded.
Ecology. Primary and secondary forest on poor soils at low altitudes.
Uses. Species of Sarcotheca produce a light, easily worked timber, which is sometimes used
for roofing and interior work, but it lacks both strength and durability. The fruit, although
sour, is eaten in curry dishes, and cooked with vegetables and sweets. It is said to be a
remedy for cough.
Key to Sarcotheca species

1.
Leaves trifoliolate; terminal leaflet larger than lateral leaflets. Mature fruits greenish
yellow when fresh, red when dry.........................................................1. S. diversifolia
Leaves unifoliolate. Mature fruits red when fresh.........................................................2
2.
Young twigs and abaxial surface of leaf-blades densely brownish velvety hairy; leaflet
margin curled inwards............................................................................4. S. ochracea
Young twigs and abaxial surface of leaf-blades glabrous to sparsely hairy; leaflet
margin not curled inwards...........................................................................................3
3.
Young twigs 4-angled. Leaves oblong to oblanceolate, 1427 x 48 cm; lateral and
intercostal veins distinctly prominent on the lower surface................3. S. macrophylla
Young twigs terete (rounded). Leaves elliptic, oblong or lanceolate, 4.515 x 26 cm;
lateral and intercostal veins prominent to inconspicuous on the lower surface..............4
4.
Leaf blades glaucous below, elliptic or oblong; veins not drying reddish; stalks long
(stalk:blade ratio 1:4.5). Panicles short and compact. Sepals dark red to crimson...........
...................................................................................................................2. S. glauca
Leaf blades not glaucous below, oblong to lanceolate; veins drying reddish; stalks short
(stalk:blade ratio 1:10). Panicles long and lax. Sepals rusty.................5. S. rubrinervis
1. Sarcotheca diversifolia (Miq.) Hallier f.

(Latin, diversifolius = with leaves of different shapes)
Med. Rijksherb. Leiden 1 (1911) 2; Veldkamp l.c. (1967) 529, l.c. (1971) 170; Cockburn l.c. (1980)
71; Whitmore, Tantra & Sutisna l.c. 282. Basionym: Rourea diversifolia Miq., Fl. Ind. Bat. 1 Suppl.
(1860) 528. Type: Teijsmann HB 707 (= ? 706), West Sumatra, Sibolga, Morsala Island (holotype L;
isotypes BO, CAL, K). Synonyms: Connaropsis diversifolia (Miq.) Kurz, J. As. Soc. Beng. 39, 2
(1870) 69, excl. syn. C. griffithii; Santalodes diversifolium (Miq.) O. Kuntze, Rev. Gen. Pl. 1 (1891)
155; Connaropsis acuminata Pears., Kew Bull. (1906) 2; S. acuminata (Pears.) Hallier f., Beih. Bot.
Centralbl. 34, 2 (1917) 27; S. subtriplinervis Hallier f. l.c. (1917) 27; Connaropsis grandiflora Ridl.,
Kew Bull. (1930) 75.
Small to medium-sized tree to 30 m tall, 50 cm diameter, sometimes with equal buttresses to

2 m tall. Bark smooth, pock-marked or occasionally scaly, reddish brown; inner bark
generally light red or pink. Sapwood white, pale yellow or sometimes pinkish. Twigs
rounded. Leaves trifoliolate, stalks 517 mm long, 12.5 mm thick, rachis 831 mm long,
0.752 mm thick; leaflets ovate to elliptic or lanceolate, lateral leaflets 2.510.5 x 14 cm,
sometime falling off early but leaving a scar, terminal leaflet often twice the size of the other
290
OXALIDACEAE (CHUNG)
leaflets, papery to thinly leathery, glabrous; base acute to broadly rounded, apex acute to
caudate; lateral veins 25 pairs, prominent on both surfaces; intercostal veins net-like;
leaflet stalk 3.58 mm long, 0.752 mm thick. Inflorescences 14 panicles together, shorter
than the subtending leaf, loosely branched; 19 cm long, rusty hairy, becoming glabrous at
mature state. Flower stalk with proximal part 2.55 mm long, distal part 13 mm long;
sepals purplish, ovate to oblong, 2.55 x 1.53 mm, apex acute to emarginate, abaxially
puberulous to glabrous, mostly falling off, seldom persistent in fruit; petals pink to red,
obovate-oblong to obovate-lanceolate, 810 x 23.5 mm, apex rounded to emarginate, 12
mm long clawed; filaments in short-styled form 2.53.5 mm and 3.54.5 mm, in long-styled
form 1.52.5 mm and 2.53 mm; ovary ellipsoid, puberulous to glabrous, styles in shortstyled form 0.51 mm long, in long-styled form 2.54 mm long. Fruits green when young,
greenish yellow when mature, ellipsoid, glabrous, 1220 x 915 mm. Seeds none to 7 per
fruit, 67.5 x 4.55 mm, testa smooth.
Vernacular names. Sabahiba jantan (Suluk), tabarus (general). Sarawakbuah piang
(Iban and Malay), jiwang (Punan). Bruneikerapa-kerapa, perapan macas, tabaus, tebarus
(Tutong Dusun). Generalbelimbing bulat (Malay).
Distribution. Sumatra and Borneo. Common in the coastal area of Sabah and throughout
Sarawak.
Ecology. Primary mixed dipterocarp forest and secondary forest from lowlands to 900 m, on
leached yellow, especially humult sandy ultisols, occasionally in mixed swamp forest but
very rare in the Rejang delta. Flowering in FebruaryNovember, fruiting in MarchJune
and SeptemberDecember.
2. Sarcotheca glauca (Hook. f.) Hallier f.
Fig. 1.
(Greek, glaukos = bluish grey; the lower leaflet surface)

l.c. (1911) 2; Merrill l.c. (1921) 312; Masamune l.c. 355; Veldkamp l.c. (1967) 535, l.c. (1971) 172;
Cockburn l.c. (1980) 71; Anderson l.c. 286; Whitmore, Tantra & Sutisna l.c. 282. Basionym:
Connaropsis glauca Hook. f. l.c. (1860) 166. Type: Lobb 1857, Borneo (holotype K).
Small to medium-sized tree to 21 m tall, 30 cm diameter; no buttresses. Bark smooth to

slightly fissured, occasionally scaly, often warty; inner bark yellow-brown, sometimes with
a pink tinge. Sapwood usually white or pale yellow, rarely pink. Twigs rounded. Leaves
unifoliolate, stalk 730 mm long, 0.51 mm thick, leaflet stalk 36 mm long, 11.5 mm
thick; blades elliptic to oblong, 4.511 x 1.55 cm, slightly larger on sterile twigs, papery to
thinly leathery, glabrous, glaucous, dull, grey-green on the abaxial surface, sometimes shiny
on the adaxial surface; base broadly cuneate to rounded or subcordate, apex acute,
acuminate or caudate; lateral veins 57 pairs, more or less looping and joining near the
margin; intercostal veins net-like. Inflorescences 12 panicles together, erect, compact,
usually many-flowered, 213 cm long, covered with minute rusty scale-like indumentum.
Flower stalk with proximal part 25.5 mm long, distal part 1.52 mm long; sepals dark red
to crimson, ovate to oblong, 1.53 x 0.51.5 mm, apex rounded, minutely hairy on the
abaxial surface, glabrescent in fruit; petals red, darker at apex, oblong to lanceolate, 3.56 x
12 mm, apex rounded to slightly emarginate, 0.51.5-mm-long-clawed; filaments in short291
styled form 22.5 mm and 2.53.5 mm, in long-styled form 11.5 mm and 1.52 mm long;
ovary ellipsoid, rusty hairy, styles in short-styled form 0.51 mm long, in long-styled form
2.53 mm long. Fruits bright pink to dark red, turning black when old, ellipsoid to nearly
round, 7.512.5 x 511 mm. Seeds 19 per fruit, 67 x 3.54 mm, testa smooth.
Vernacular names. Sabahasam daham (around Sandakan), gitan gizu (Dusun Rungus),
kandis daham (around Mempakul), rangkas-rangkas (Dusun Kinabatangan), t-izos-izos
(around Papar). Sarawakaremajuh (Dayak), medang, piang (Iban), tulang payung piang
(Malay). Bruneiasam piai (Dusun), kandis daham (Kedayan).
Distribution. NW Borneo (Sabah, Sarawak and Brunei). Common in Sabah and Sarawak.
No record from Kalimantan.
Ecology. Primary and secondary lowland forests, often on hill slopes. In Sarawak,
uncommon in mixed swamp forest, occasional or widespread in kerangas forest and mixed
dipterocarp forest on sandy humult ultisols. Flowering in FebruaryApril and
SeptemberDecember, fruiting in OctoberMay.
3. Sarcotheca macrophylla Blume

(Greek, makros = large, phullon = leaf)
Whitmore, Tantra & Sutisna l.c. 282. Type: Mller, s.n., Borneo (lectotype L; isolectotype BO).
Synonym: Roucheria macrophylla Miq. l.c. (1859) 136.
Small tree to 15 m tall, 10 cm diameter; no buttresses. Bark smooth, pale brown; inner bark
reddish. Sapwood white. Twigs 4-angled when young and rounded when old. Leaves
unifoliolate, stalk 512 mm long, 1.53 mm thick, leaflet stalk 39 mm long, 13 mm thick;
blades oblong to oblanceolate, 1526 x 39 cm, thinly leathery, adaxial surface glabrous,
abaxial surface puberulous on the veins; base rounded to truncate or subcordate, apex
acuminate to cuspidate; midrib puberulous, sunken above; lateral veins 613 pairs,
distinctly prominent on the abaxial surface; intercostal veins net-like, distinctly prominent.
Inflorescences 14 panicles together, slender, lax, pendulous; 1285 cm long, brownpuberulous. Flower stalk with proximal part to 2 mm long, distal part to 1 mm long; sepals
obovate to ovate, 1.52.5 x 11.5 mm, apex rounded, abaxially rusty hairy, puberulous in
fruit; petals dark red, obovate-oblong to oblanceolate, 35 x 12 mm, apex obtuse to
emarginate, to 0.5-mm-long-clawed; filaments in short-styled form 12 mm and 23 mm
long, in long-styled form 11.5 mm and 1.52.0 mm long; ovary subglobose to ellipsoid,
densely hairy, styles in short-styled form 0.250.75 mm long, in long-styled form 12 mm
long. Fruits dark red, subglobose, 611 x 58 mm, glabrescent, glossy. Seeds to 9 x 4 mm,
testa wrinkled.
Vernacular names. Sarawakbelimbing manik (Bakumpai-Dayak), piang (Iban).
292
OXALIDACEAE (CHUNG)
293
Distribution. Borneo (Sarawak and Kalimantan). Only one collection, Beccari PB 3166
reported from Marop, Sarawak.
Ecology. Primary and secondary forest on sandy soils at low altitude. Flowering and
fruiting in January-May.
4. Sarcotheca ochracea Hallier f.

(Greek, ochre = yellow to yellowish brown; the dry leaves)
Anderson l.c. 286; Whitmore, Tantra & Sutisna l.c. 282. Type: Haviland 2343, Borneo (holotype L;
isotypes K, SING).
Small tree to 10 m tall, 10 cm diameter; no buttresses. Bark brownish; inner bark pale
brown. Twigs rounded. Leaves unifoliolate, stalk 824 mm long, 1.52 mm thick, leaflet
stalk 36 mm long, 1.53 mm thick; blades elliptic to oblong, 7.519.5 x 410 cm, adaxial
surface glabrous, abaxial surface rusty-pubescent to velvety; base obtuse to rounded, margin
slightly wavy and curved towards the abaxial surface, apex acuminate; lateral veins 69
pairs, abaxial surface prominent; intercostal veins net-like, prominent. Inflorescences 12
panicles together, erect to pendulous, compact to elongate, branched in comb-like way,
many-flowered, 764 cm long, rusty-velvety. Flower stalk with proximal part 48 mm
long, distal part 0.51 mm long; sepals reddish brown, obovate, 2.53 x 23 mm, apex
rounded to retuse, rusty-puberulous on the abaxial surface, puberulous in fruit; petals
scarlet, darker at apex, obovate to lanceolate, 68 x 1.52 mm, apex obtuse to rounded,
1.52.0-mm-long-clawed; filaments in short-styled form c. 2.5 mm and 3.5 mm long, in
long-styled form 1.52 mm and 22.5 mm long; ovary subglobose, rusty hairy, styles in
short-styled form 0.51 mm long, in long-styled form 22.5 mm long. Fruits bright red,
subglobose to oblong with rounded apex, 815 x 615 mm. Seeds 12 per fruit, 57 x
34.5 mm, testa wrinkled.
Vernacular names. Sarawakikor mata (Iban), pechi mata (general).
Distribution. Borneo. In Sarawak uncommon (Miri, Bintulu and Tatau).
Ecology. In mixed dipterocarp forest on pale yellow infertile clay soils. Flowering and
fruiting in AprilOctober.
5. Sarcotheca rubrinervis Hallier f.

(Latin, ruber = red, nervus = nerve)
Cockburn l.c. (1980) 73; Whitmore, Tantra & Sutisna l.c. 283. Type: Amdjah 1082, E Borneo
(holotype L; isotype BO). Synonym: S. oblongifolia Merr. l.c. (1929) 111.
Small to medium-sized tree to 20 m tall, 30 cm diameter; no buttresses. Bark smooth, thin,

reddish to reddish brown; inner bark yellow or yellowish brown, sometimes with a pink
tinge, fibrous. Sapwood white or pale yellow. Twigs rounded. Leaves unifoliolate, stalk
620 mm long, 1.52 mm thick, leaflet stalk 24 mm long, 1.52 mm thick; blades oblong
to lanceolate, 6.515 x 26 cm, glabrous, abaxial surface not or sometimes slightly
glaucous; base obtuse to rounded, apex acute to acuminate; lateral veins 411 pairs, often
with reddish tinge, more or less looping and joining near the margin; intercostal veins
irregular, the more prominent ones running from vein to vein or from vein to midrib, fainter
294
OXALIDACEAE (CHUNG)
ones net-like. Inflorescences 12 panicles together, elongate, lax, pendulous, 636 cm long,
rusty puberulous. Flower stalk with proximal part 35 mm long, distal part up to 1 mm
long; sepals ovate to lanceolate, 1.52.5 x 11.5 mm, apex acute to emarginate, minutely
rusty hairy at base, puberulous in fruit; petals red or rarely white, lanceolate, twice as long
as sepals, apex rounded to emarginate, 0.51-mm-long-clawed; filaments in short-styled
form 1.52 mm and 2.53 mm long, in long-styled form 0.51 mm and 11.5 mm long;
ovary subglobose, densely hairy, styles in short-styled form c. 0.5 mm long, in long-styled
form 1.52 mm long. Fruits pink to bright red, subglobose, 610 x 510 mm, glaucous to
glossy. Seeds to 6.5 x 5 mm, testa smooth to more or less wrinkled.
Vernacular names. Sabahasem-asem (Dusun Kinabatangan), iba talon (Bajau), ira
prumpuan (Suluk), lampyos (Dusun Penampang), pinggoh, pinguh (Kalabakan), tabarus
(general).
Distribution. Borneo (Sabah and Kalimantan). In Sabah only known from Tawau.
Ecology. Primary and secondary forests near rivers on loamy soils of flat to undulating
lands. Flowering and fruiting in MarchDecember.
295
PITTOSPORACEAE
John B. Sugau
Sandakan, Malaysia
Merrill, EB (1921) 287; Masamune, EPB (1942) 324; Bakker & van Steenis, FM 1, 5 (1957) 345;
Whitmore, TFM 2 (1973) 309; Anderson, CLTS (1980) 286; Ashton, MNDTS 2 (1988) 325; Corner,
WSTM 2 (1988) 607; Sugau, Sandakania 4 (1994) 41.
Small to medium-sized trees, shrubs, or climbers; indumentum if present consisting of

simple hairs; bark resinous. Leaves spirally arranged, often crowded towards the end of the
twigs, simple, mostly entire, pinnately veined; stipules absent. Inflorescence a few- to
many-flowered, axillary or terminal, bracteate corymbose panicle, raceme or thyrse, sometimes
on bare branches, often also as fascicles of flowers. Flowers regular, mostly 5-merous,
bisexual or functionally unisexual; bracteoles often 2; sepals 5, free or connate at the base,
overlapping; petals 5, often narrowed at the base and loosely joined as a tube, overlapping,
caducous; stamens 5, inserted on the sepals, erect, free, or slightly connivent at the base,
anthers 2-celled, introrse, basifixed, dehiscing lengthwise (or by pores); ovary superior,
sometimes shortly stalked, mostly 1-chambered, sometimes completely or incompletely 2chambered, placentas 26, basal or on the wall, style simple, stigma simple and thickened,
or lobed, ovules numerous, anatropous with one integument. Fruit a berry or a loculicidal,
2- or 6-valved capsule. Seeds few to many per fruit, embedded in a resinous or oily pulp or
fluid, seldom winged, testa thin, smooth; endosperm copious, hard; embryo small, close to
the hilum; cotyledons small, 25.
Distribution. 9 genera restricted to the Old World, chiefly Australian; 6 genera are completely
restricted to Australia; and 2 in E Malesia and Australia; 1 (Pittosporum) is widely
distributed and the sole representative of the family in Sabah and Sarawak.
Ecology. Members of the family can be found in everwet or dry regions, from tropical to
warm-temperate areas.
Uses. Several genera have species that are grown as ornamentals; it has been suggested
(Bakker & van Steenis l.c.) that a few Pittosporum species might be used for re-afforestation of
bare lands, on account of their adaptability to such sites.
PITTOSPORUM Banks ex Sol.

(Greek, pitta = resin, spora = seed; the resinous seed)
in Gaertner, Fruct. 1 (1788) 286; Merrill l.c. 287; Masamune l.c. 324; Bakker & van Steenis l.c. 345;
Whitmore l.c. 309; Anderson l.c. 286; Ashton l.c. 325; Corner l.c. 607; Sugau l.c. 41.
297
Trees or shrubs, with rhythmic growth; terminal bud protected by scale leaves. Leaves
entire, often hairy when young. Flowers few to many, in fascicles, thyrses or corymbs,
seldom solitary, white or pale yellow, fragrant; sepals free or more or less connate; petals
generally ligulate, free or united in the lower part, the free segments spreading or recurved;
stamen filaments slender in male flowers, somewhat shorter and slightly broadened at the
base in female flowers, anthers generally oblong in male flowers, sagittate and smaller in
female flowers; ovary sessile or stalked, glabrous or hairy, 1-chambered, slender in male
flowers but stout in female flowers; style in female flowers short and with a 25-lobed
capitate stigma, in male flowers longer and with obscure stigmas; placentas 25, basal or on
the ovary wall, ovules few to many. Fruit a loculicidally dehiscent, 2(6)-valved capsule,
usually ripening orange. Seeds 1 to many per fruit, covered by a resinous viscous fluid;
germination epigeal.
Distribution. About 100 species, from warm-temperate to tropical areas, from Africa, South
and East Asia and Australasia, to the Pacific Islands. 6 species are documented for Borneo,
all found in Sabah, while only 2 occur in Sarawak.
Ecology. Species of Pittosporum are generally understorey shrubs or trees of the rain forest,
occurring from near sea-level to the subalpine zone on Mt. Kinabalu.
Key to Pittosporum species

(based mainly on leaf characters)
1.
Leaves very narrowly elliptic to linear, to 1.1 cm wide .....................2. P. linearifolium

Leaves elliptic, oblanceolate or obovate, typically more than 1.5 cm wide....................2
2.
Leaves persistently thickly rusty hairy on the lower side, and conspicuously bullate on
the upper side; margins markedly recurved when dry.............................6. P. silamense
Leaves glabrous or, if hairy, only when very young, and not bullate or only finely so on
the upper side; margins not or only weakly recurved when dry.....................................3
3.
Lateral veins finely impressed above. Sepals 67 mm long...............3. P. longisepalum

Lateral veins flat above. Sepals less than 5 mm long....................................................4
4.
Leaves markedly obovate, the apex abruptly acuminate to cuspidate. Inflorescence

borne on bare branches................................................................................................5
Leaves elliptic, or if oblanceolate or obovate, then the apex gradually acuminate.
Inflorescence borne in the leafy portions of branches.........................1. P. ferrugineum
5.
Leaf-stalks 818 mm long. Petals 912 mm long. Infructescence-stalks not conpicuous

or to 0.5 cm long. Mature fruits to 1.72 cm long...............................5. P. resiniferum
Leaf-stalks, 1525 mm long. Petals only 47 mm long. Infructescence-stalks 13 cm
long. Mature fruits 11.5 cm long.......................................................4. P. ramiflorum
298
PITTOSPORACEAE (SUGAU)
Key to Pittosporum species

(based on flower and fruit characters)
1.
Inflorescences mainly on bare branches....................................................................2

Inflorescences among leaves on the branches...............................................................3
2.
Petals 47 mm long. Infructescence-stalks 13 cm long. Fruit 11.5 cm long................

...........................................................................................................4. P. ramiflorum
Petals 912 mm long. Infructescence-stalks inconspicuous or to 0.5 cm long. Fruits
1.72 cm long...................................................................................5. P. resiniferum
3.
Seeds attached to the fruit-wall to halfway to the fruit-apex...............1. P. ferrugineum

Seeds attached to the basal portion of the fruit-wall......................................................4
4.
Fruits, at least when young (and ovary), glabrous, or at most slightly hairy at the base
........................................................................................................2. P. linearifolium
Fruits, at least when young (and ovary), hairy all over.................................................5
5.
Infructescence-stalks 0.81.2 cm long. Seeds to 11 per fruit...................6. P. silamense

Infructescence-stalks 23 cm long. Seeds 14 per fruit.....................3. P. longisepalum
1. Pittosporum ferrugineum Aiton

(Latin, ferrugineus = rusty; the brown leaf tomentum)
Hort. Kew ed. 2, 2 (1811) 27; Merrill l.c. 287; Masamune l.c. 324; Bakker & van Steenis l.c. 345;
Whitmore l.c. 309; Anderson l.c. 286; Ashton l.c. 322; Corner l.c. 607. Type: Hort. Kew (from a
cultivated plant in the Royal Botanic Gardens Kew; native of New Guinea and introduced before 1787
by the Right Hon. the Earl of Tankerville), fide Aiton (BM). Synonyms: Itea javanica Blume, Bijdr.
(1926) 863; Pseuditea javanica (Blume) Hassk., Flora 25, 2 (1842) Beibl. 30; Pittosporum javanicum
(Blume) Blume, Mus. Bot. Lugd. Bat. 1 (1850) 159; Pittosporum rufescens Turcz., Bull. Soc. (Imp.)
Nat. Mosc. 27, 2 (1854) 367; Pittosporum nativitatis Bakker in Andrews, Monogr. Christm. Isl. (1960)
171; Pittosporum versteeghii Merr. & Perry, J. Arn. Arb. 21 (1940) 177.
Small to medium-sized tree, to 15 m tall, occasionally reaching 20 m tall, 25 cm diameter.

Bark brownish; inner bark yellowish. Sapwood pale yellow, soft. Young shoot more or less
densely covered with caducous rusty brown hairs. Leaves typically elliptic, rarely
oblanceolate or obovate, 612 x 24 cm, papery, pale brown on both surfaces; base cuneate,
margin plane and slightly undulate, apex narrowly acute to gradually acuminate; midrib
sharp on the lower side but depressed on the upper side; lateral veins 59 pairs, flat above;
intercostal veins reticulate, fine; leaf-stalks 815 mm long, slender and grooved on the
upper side; Inflorescence a many-flowered thyrse, 1.55 cm long, borne among leaves on
the branches, or terminal; peduncle 15 cm long, rusty-hairy. Flowers pale yellow, sepals
lanceolate to linear, 24 mm long, generally rusty hairy; petals ligulate, 68 x 1 mm;
stamens 34 mm long, filaments 2.53 mm long, anthers 0.51 mm long; ovary ellipsoid or
cylindric, sessile, densely rusty hairy, style 11.5 mm long, glabrous, stigma often scarcely
capitate in male flowers, 2-lobed in female flowers. Fruits capsular, 116 in an
infructescence, often broader than long or subglobose, 0.51 cm long, 2-valved, generally
notched and mucronate apically, narrowed towards the often hairy base, rugose, ripening
orange; infructescence-stalks 15 cm long, hairy. Seeds 824 per fruit, attached on a
placenta halfway to the apex of the fruit, shiny black, fixed firmly in a sticky pulp.
299
Vernacular names. Sabahara bukit (Brunei Malay), nonok (Dusun Papar, in confusion
with Ficus), soipang/saipang (Dusun Ranau)
Distribution. SE Asia to Australia and Melanesia, usually in open vegetation, especially
along sea-shores and on mountains to 2000 m. In Sarawak, so far known only from the 1st
Division (although expected elsewhere); in Sabah widespread and known in nearly every
district.
Ecology. Common in the lowlands near the coast, especially on rocky and sandy shores, and
montane zones. Locally frequent in secondary forest, infrequent in the understorey of mixed
dipterocarp forest in Sabah; in open vegetation or limestone summits in W Sarawak.
Uses. The leaves and fruits contain saponin and are used for fish poison. The wood is
sometimes used as firewood by local people in Sabah, probably on account of the combustible
resin content.
2. Pittosporum linearifolium J. B. Sugau
Fig. 1.
(Latin, linearis = linear, folium = leaves)

Sandakania 4 (1994) 41. Type: L. Madani SAN 89542, Sabah, Ranau, Bukit Ampuan (holotype SAN;
isotypes K, KEP, SAR).
Medium-sized tree, to 10 m tall. Bark smooth, whitish; inner bark creamy grey. Sapwood
white to brown. Leaves very narrowly elliptic to linear, 47 x 0.51.1 cm, chartaceous,
glabrous; base narrowly cuneate, margins slightly recurved when dry, apex narrowly acute;
midrib slightly sunken on the upper side but raised on the lower side; lateral veins 56 pairs,
inconspicuously sunken on the lower side and flattened on the upper side; reticulations very
fine; stalks 15 mm long, slender, grooved on the upper side. Inflorescence a simple thyrse.
Flowers unknown. Fruits 14 in an infructescence, capsule subglobose, 0.81.1 cm long, 2valved, notched and with an abrupt tip, narrowed towards the base, glabrous, rugose;
infructescence-stalks less than 1 cm long, glabrous. Seeds 24 per fruit, attached on a
placenta at the base of the fruit, shiny black.
Distribution. Endemic to Sabah. Very localised, restricted to the Bukit Ampuan area in
Ranau.
Ecology. Hill forest to 1200 m.
3. Pittosporum longisepalum Bakker

(Latin, longus = long, sepalum = sepal)
in Bakker & van Steenis l.c. 354. Type: Clemens 35096, British North Borneo, Mt. Kinabalu, Ranau
(holotype L; isotype K).
Small tree to 10 m tall. Leaves in distinct clusters on the twigs, elliptic, 46 x 1.52 cm;
base cuneate, margin slightly recurved, apex acuminate, acumen 0.50.8 cm; sparsely hairy
300
301
on the lower side, not bullate on the upper side; midrib impressed on the upper side, raised
on the lower side; lateral veins 57 pairs, finely impressed on the upper side, raised on the
lower side; intercostal veins reticulate, very fine; stalk c. 5 mm long, slender. Inflorescences 3flowered cymes, terminal on leafy shoots; peduncle 23 cm long, hairy. Flowers with sepals
free, linear-lanceolate, 67 x 11.5 mm, hairy, acute, longer than the petals in bud; petals
narrowly oblanceolate, c. 8 x 12 mm, acute; stamens 45 cm long, filaments c. 3 mm long,
anthers c. 2 mm long, apiculate; ovary ellipsoid, subsessile, densely hairy to 1 mm long
from the base, stipe with 5 furrows, c. 2 x 11.5 mm. Fruits subglobose, c. 0.7 cm across, 2valved, mucronate, rugose; infructescence-stalks 23 cm long, hairy. Seeds 14 per fruit, c.
4 x 3 mm, attached on a placenta at the base of the fruit.
Distribution. Sabah (Mt. Kinabalu) and Central Celebes.
Ecology. Mainly in submontane rain forest, at 12001500 m.
4. Pittosporum ramiflorum (Zoll. & Moritzi) Zoll. ex Miq.

(Latin, rami = branches, florum = flowers; the inflorescences on the bare branches)
Fl. Ind. Bat. 1, 2 (1858) 122; Bakker & van Steenis l.c. 345. Basionym: Glyaspermum ramiflorum
Zoll. & Moritzi, Nat. Geneesk. Arch. Ned. Ind. 2 (1845) 11. Type: Zollinger 2139, Java, Gede (P).
Synonyms: Pitttosporum ramiflorum var. parviflorum K. Schum. in K. Schumann & Hollrung, Fl.
Kais. Wilh. Land (1889) 70; P. comptum K. Schum. & Lauterb., Fl. Schutzgeb. (1900) 338; P.
clementis Merr., Philip. J. Sc. 3 (1908) Bot. 137.
Shrub or tree, (3)520(30)m tall. Leaves spirally arranged or in distinct clusters along the
twigs, obovate, 518 x 2.56 cm, plane, subcoriaceous, glabrous when mature; base broadly
cuneate, margin entire, apex abruptly acuminate, acumen for 0.51.5 cm long; midrib
flattened on the upper side but raised on the lower side; lateral veins 58 pairs, flattened on
the upper side but somewhat raised on the lower side; intercostal veins reticulate, fine; stalks
1525 mm long, slender and grooved on the upper side. Inflorescences densely branched and
many-flowered, mainly on the bare parts of branches; peduncles 13 cm long, hairy; bracts
ovate, c. 11.5 mm, caducous; pedicels to 6 mm. Flowers white, sepals united at their base
in a shallow cup, 1.52 mm high, otherwise free, with rounded apices; petals narrowly
elliptic, (4)56(7) x (1)2(3) mm, rounded; stamens 45 mm long, filaments 35 mm
long, anthers oblong, 0.72 mm long; ovary ellipsoid or obovoid, 23 x 11.5 mm,
subsessile, brownish hairy, style 13 mm long, stigma 2-lobed. Fruits varying in number,
on woody stalks, ellipsoid to obovoid, compressed, 11.5 cm long, 2-valved, with an abrupt
tip, valves hard, 12 mm thick, wrinkled; infructescence-stalks 13 cm long, hairy. Seeds
many per fruit, attached to the fruit-wall from the base up to apex of the fruit, very irregular
in shape to roundish, flat, 23 mm diameter.
Distribution. Java, Borneo, Southern Philippines (Negros, Cebu, Mindano and Bohol),
Celebes, the Moluccas (Ambon, Buru, Ceram), New Guinea, Solomon Islands. In Sabah
only found on Mt. Kinabalu; not yet recorded from Sarawak.
Ecology. An understorey tree in rain forest, sometimes also riverine, occasionally to 3200 m
in montane forest.
302
5. Pittosporum resiniferum Hemsl.

(Latin, resinifer = producing resin; the capsule)
Kew Bull. (1894) 344; Bakker & van Steenis l. c. 345. Type: Holmes, s.n., Philippines (lectotype K,
here designated). Synonyms: P. epiphyticum Merr. l.c. (1908) 138; P. accuminatissimum Merr.,
Philip. J. Sc. 14 (1919) 402.
Small to medium-sized tree, 210 m tall, 1020 cm diameter. Bark thin, whitish to dark
brown; inner bark creamy pink, exudate sticky. Sapwood pale white. Leaves spirally
arranged, markedly obovate, 918 x 2.55 cm, glabrous; base narrowly cuneate, margin
sligtly recurved when dry, rather thick, apex abruptly cuspidate; midrib slender, prominent
on the lower side, flattened on the upper side, pale brown; lateral veins 710 pairs, flattened
on upper side, raised on the lower side; intercostal veins reticulate, fine; stalks 818 mm
long, slender, grooved on the upper side. Inflorescence a false umbel, 11.5 cm long, borne
on bare branches below the leaves; peduncles not conspicuous or only to about 0.5 cm long,
slightly hairy. Flowers whitish; sepals united at their lower half into a shallow cup, 12 mm
high, apices rounded, 12 mm long; petals narrowly oblong, 912 x 1.52 mm; stamens 79
mm long, filaments slender, 67 mm long, anthers 1.52 mm long; ovary ellipsoid, 45 x
1.52 mm, sessile, densely hairy, style 22.5 mm long, stigma thickened. Fruits 16 in an
infructescence, ripening orange, capsule globose to ellipsoid, 1.72 cm long, very hard
when dry, 2-valved, notched and with an abrupt tip, rounded to cordate at the base,
glabrous, rugose; infructescence-stalks inconspicuous or to about 0.5 cm long, hairy to
glabrous. Seeds many per fruit, attached to the fruit-wall from the base to the apex of the
fruit.
Vernacular name. Sabahbedung (Dusun Ranau).
Distribution. Sabah (Mt. Kinabalu) and the Philippines (Luzon, Mindoro, Panay, Catanduanes,
Leyte, and Mindanao).
Ecology. In montane forest, sometimes also at lower localities. It seems to be a lightdemanding species.
6. Pittosporum silamense J. B. Sugau

(of Mt. Silam, Sabah)
Sandakania 4 (1994) 41. Type: Joseph et al. SAN 120894, Sabah, Lahad Datu, Mt. Silam (holotype
SAN).
Small tree, bole c. 5 cm diameter. Bark smooth, brownish; inner bark yellowish. Sapwood
yellowish. Leaves generally elliptic, 5.510 x 2.54 cm, conspicuously bullate on the upper
side, persistently thickly rusty-hairy on the lower side; base cuneate, margins markedly
recurved when dry, rather thick, apex narrowly cuneate to acuminate; midrib sunken on the
upper side, raised on the lower side; lateral veins 69 pairs, deeply impressed on the upper
side, raised on the lower side; stalks 818 mm long, slender, grooved on the upper side.
Inflorescence a many-flowered thyrse. Flowers unknown. Fruits 1018 in an
infructescence, greenish to blue when unripe, black when ripe, capsule subglobose, 0.81.1
cm long, 2-valved, notched and with an abrupt tip, base narrowed to about 2 mm long,
303
covered with rusty hair; infructescence-stalk 0.81.2 cm long, hairy. Seeds to 11 per fruit,
attached to a placenta at the base of fruit, black.
Distribution. Endemic to Sabah. Apparently restricted to Mt. Silam in Lahad Datu district.
Ecology. Hill forest, about 800 m, on ultramafic soil.
304
RHAMNACEAE
Carsten Schirarend
Botanischer Garten und Botanisches Museum Berlin-Dahlem,
Berlin, Germany
Ridley, FMP 1 (1922) 461; Masamune, EPB (1942) 434; Suessenguth in Engler & Prantl, Pfl. Fam. 20
d (1953) 7; Backer & Bakhuizen f., FJ 2 (1965) 80; Anderson, CLTS (1980) 289; Keng, OFMSP
(1983) 203; Corner, WSTM 2 (1988) 609; Latiff, TFM 4 (1989) 297; Whitmore, Tantra & Sutisna,
CLK 2, 1 (1990) 293.
Deciduous or evergreen, often thorny trees, shrubs, woody climbers or very rarely herbs.
Leaves simple, alternate or rarely opposite, pinnately veined or 3-veined. Stipules general-ly
present, small, deciduous or often transformed into straight to recurved spines. Inflorescences basically cymose, cymes mostly axillary, sessile or peduncled, or reduced to manyto few-flowered fascicles. Flowers radially symmetrical, 4- or 5-merous, rarely 3- or 6- or
7-merous, bisexual or unisexual (plants monoecious and/or dioecious); hypanthium
resembling a calyx-tube, patelliform or hemispherical to tubular; sepal 4 or 5, valvate in
bud, triangular, erect to more or less recurved during anthesis, often medially keeled within; petals 4 or 5, rarely absent, usually smaller than the alternate sepals, concave or hooded
(cucullate), rarely almost flat, often shortly clawed, enfolding the epipetalous stamens;
stamens 4 or 5, in a single series opposite the petals, filaments thin, adnate to the base of the
petals, anthers minute, versatile, generally 2-thecate, dehiscing by longitudinal slits; disc
intrastaminal, nectariferous, thin to more or less fleshy, entire or lobed, glabrous or rarely
pubescent, free from the ovary or adnate around the base; ovary superior or half-inferior,
23(5)-loculate, with 1 ovule in each locule; ovules anatropous, basal and erect. Fruit a
capsule or a dry or fleshy drupe with 13 dehiscent or indehiscent stones (endo-carpids),
often winged. Seeds with scanty fleshy endosperm or rarely without endosperm; embryo
large, oily, straight or rarely bent.
Distribution. A cosmopolitan family of about 45 genera and 800 to 1000 species. In Sabah
and Sarawak, represented by 8 native genera with 21 species of which only 4 species in 4
genera are trees, the others are either erect or straggling shrubs or woody climbers.
Ecology. Tropical rain forest to moderately arid areas and from near sea-level to elevation
and latitude near the tree limit.
Uses. A rather limited number of Rhamnaceae species furnish useful and valuable timber of
local importance. Some West Indian species of Colubrina, Krugiodendron and Reynosia
yield a hard and heavy wood sold as Ironwood. Ornamental plants are restricted to the
North American genus Ceanothus, of which several species are planted for their showy
inflorescences, and the European Paliurus spina-christi, which is widely used as protective
and ornamental hedge-plants. Edible fruits are furnished by the Mediterranean jujubes
(Ziziphus), which are widely cultivated in Africa and Asia. Further economic products of
305
rhamnaceous plants are dyes (Rhamnus), laxatives and diuretics (Frangula, Paliurus,
Rhamnus), soap (Colubrina, Ziziphus) and substitute for tea (Sageretia). None of these uses
is, however, reported for taxa occurring in Sabah and Sarawak.
Taxonomy. Suessenguth (l.c.) recognised 58 genera and grouped them into five tribes,
mainly distinguished by ovary and fruit characters. The 21 genera of the largest tribe
Rhamneae extend over nearly the whole area of the family, except extratropical South
America. In Sabah and Sarawak, they are represented by Alphitonia, Colubrina and
Rhamnus. The tribe Zizipheae comprises 20 genera, and is mainly centered in subtropical
and tropical regions of the world, namely in the West Indies and Central America, East Asia
and Indo-Malesia. Berchemia and Ziziphus are the two genera native to Borneo. The tribe
Colletieae are mostly restricted to the extratropical South America, with one genus
(Adolphia) in Mexico and California and another genus (Discaria) extending to New
Zealand and Australia. The predominantly climbing or even herbaceous tribe Gouanieae are
confined to a narrow pantropical belt. Out of the five genera recognised, only Gouania is
widely distributed throughout Malesia, including Sabah and Sarawak. Smythea and
Ventilago, the only two genera of the tribe Ventilagineae, are restricted to the Paleotropics,
and both occur in Sabah and Sarawak.
Key to genera
1.
Small to medium-sized trees or shrubs; fruits never winged.........................................2

Woody climbers with winged fruits..............................................................................6
2.
Leaves 3-veined from the base; stipules often transformed into straight or recurved
spines..........................................................................................................4. Ziziphus
Leaves pinnately veined; stipules never transformed into spines...................................3
3.
Fruit a single-stoned fleshy drupe..................................................................................

Berchemia Necker
Elem. bot. (1790) 122; Backer & Bakhuizen f. l.c. 83.
22 species; North America (1 species), Minor Asia, Indo-China, New Caledonia,

Malesia; 1 species in Sabah and Sarawak.
Erect or straggling shrubs. Leaves membranous, with numerous pairs of slightly
curved, parallel lateral veins. Inflorescences raceme-like thyrses. Flowers bisexual
or unisexual by abortion. Fruit an ellipsoid to obovoid, single-stoned fleshy drupe.
Fruit a 24-stoned capsule or drupe; epicarp dry or more or less fleshy........................4
4.
Fruit a 3-locular, capsule...........................................................................2. Colubrina

Fruit indehiscent..........................................................................................................5
5.
Epicarp mealy or corky; endocarpids splitting ventrally. Seeds distinctly arillate...........

.................................................................................................................1. Alphitonia
Epicarp fleshy; endocarpids indehiscent. Seeds never arillate.....................3. Rhamnus
306
RHAMNACEAE (SCHIRAREND)
6. Branches often with tendrils. Fruit with three rounded commissural wings....................
Gouania Jacq.
Select. Stirp. Amer. Hist. (1763) 263; Ridley l.c. 468; Masamune l.c. 434; Backer &
Bakhuizen f. l. c. 85.
About 50 species; tropical America and Africa, Madagascar, India, Indo-China,

Malesia, Australia, Polynesia; 3 species in Sabah and Sarawak.
Woody climbers with solitary tendrils. Leaves membranous to coriaceous, ovate or
cordate, pinnately veined, rarely 3-veined, margin entire to serrate. Inflorescence a
multiflowered, axillary or terminal, spike- or raceme-like thyrse. Flowers bisexual
or polygamous. Fruit a 3-locular schizocarp, splitting septicidally into three 2winged indehiscent endocarpids.
Branches never with tendrils. Fruit with one apical wing.............................................7
7.
Fruit a globose indehiscent structure, bearing a long (at least twice as long as the fruit)
apical wing....................................................................................................................
Ventilago Gaertn.
Fruct. Sem. Pl. 1, l (1788) 233; Ridley l.c. 465; Masamune l.c. 434; Backer & Bakhuizen f.
l.c. 81.
About 40 species; Africa, Madagascar, India, Indo-China, Malesia, Australia; 4

Leaves chartaceous to coriaceous, elliptic to oblong-ovate, pinnately veined,
margin entire to serrate. Inflorescences axillary clusters or shortly stalked cymes
which are often combined into racemes or panicles. Flowers bisexual, rarely
apetalous.
Fruit a compressed, septicidally dehiscing capsule bearing a small apical wing..............
Smythea Seem.
Bonplandia 9 (1861) 255; Ridley l.c. 468; Masamune l.c. 434; Backer & Bakhuizen f. l.c.
81.
7 species; Indo-China, Malesia, Polynesia; 1 species in Sabah and Sarawak.

Leaves membranous to chartaceous, elliptic to ovate, pinnately veined, often somewhat asymmetric. Flowers in axillary fascicles, bisexual.
1. ALPHITONIA Reiss. ex Endl.

(Greek, alphiton = baked barley meal; the mealy nature of the ripe fruit exocarp)
Gen Pl. (1840) 1098; Braid, Kew Bull. (1925) 168; Masamune l.c. 434; Anderson l.c. 289; Whitmore
Tantra & Sutisua l.c. 293.
Evergreen trees or shrubs; buds and young stems often more or less densely brownish
pubescent. Bark and wood of several species with rather strong smell of sarsaparilla.Leaves
alternate, ovate to obovate or elliptic to lanceolate, rarely almost cordate, chartaceous to
coriaceous, venation pinnate; blade usually glabrous above, whitish or rusty pubescent
underneath; stipules small, deciduous. Inflorescences many-flowered, di- or trichotomous,
axillary or rarely terminal cymes. Flowers 57-merous; hypanthium patelliform to hemisperical; sepals 5, triangular, keeled within; petals 5, clawed, cucullate; stamens 5; disc
mainly thick, nectariferous; ovary inferior, immersed in the stout disc, 23-locular, style
short, 23-lobed, basally often densely whitish to yellowish pubescent or glabrous. Fruits
307
drupaceous, globose or broadly ovoid; mesocarp thick, mealy; endocarp splitting longitudinally
into 2 to 3, hard and coriaceous endocarpids, at the base surrounded by the persistent
hypanthium. Seeds more or less completely enclosed by a loose, membranous, reddish
brown aril; testa smooth, crustaceous; embryo straight; endosperm cartilaginous.
Distribution. About 10 species; Malesia, Polynesia and Australia. One species in Sabah and
Sarawak.
Uses. The wood of the tree species in Australia, Malesia and most of the Polynesian Island
is said to be of considerable value (Braid l.c. 170), and used for axe-handles, piling, and
cabinet-making.
Alphitonia excelsa (Fenzl) Reiss. ex Endl.
Fig. 1.
(Latin, excelsus = lofty, high; the tree)

l.c. 1098; Maiden, For. Fl. N.S.W. (1903) 38; Braid l.c. 177; Masamune l.c. 434; Anderson l.c. 289;
Whitmore, Tantra & Sutisna l.c. 293. Basionym: Colubrina excelsa Fenzl in Endl., Enum. Pl. (1837)
20. Type: Cunningham, s.n., Australia, Queensland, Brisbane, Moreton Bay (K). Synonyms:
Rhamnus incanus Roxb., Fl. Ind. (1832) 603; Alphitonia incana (Roxb.) Kurz ex Hoogland, Kew Bull.
15 (1961) 33; A. philippinensis Braid l.c. 183.
Small to medium-sized tree rarely to 30 m high; bole straight to 20 m high. Bark and wood
with a rather strong smell of wintergreen. Bark smooth, grey to brownish; inner bark thin,
greenish, more or less straw, cambium yellow. Sapwood whitish; heartwood reddish brown.
Young twigs, axillary buds, stipules, petioles and lower surface of the leaves more or less
densely yellowish to whitish pubescent. Stipules minute, acute triangular, early deciduous.
Leaves alternate, chartaceous to subcoriaceous, ovate-lanceolate or elliptic, 620(30) x
36(10) cm; base acute or obtuse to rounded, rarely cordate, margin entire, apex acute to
shortly acuminate; midrib prominent below, impressed above; lateral veins 1014 pairs;
petioles 510 mm long. Inflorescences terminal or axillary cymes. Flowers about 45 mm
in diameter, light green to yellowish, 5-merous, rarely 6- or 7-merous; hypanthium shallow
hemispherical; sepals about 1.5 mm long; petals erect to slightly recurved; stamens about
1.5 mm long; disc shallow, lining the hypanthium, ring-shaped, glandulous; ovary
obconical, whitish tomentose; style conical. Fruits globose to broadly ovoid, to 15 mm in
diameter, greenish when young, black when ripe. Seeds often permanently attached to the
receptacle after dehiscence of the fruit.
Vernacular names. Sabahbalik angin (Dusun, Malay), pati yata (Dusun), pokudata
(Dusun).
Distribution. A widely distributed species, extending from Borneo, the Philippines, New
Guinea to Australia. In Sabah and Sarawak common and widespread in the lowlands. Also
Taxonomy. Braid (l.c.) split the former A. excelsa into a group of species poorly differentiated
in leaf shape and size. Examination of numerous specimens throughout Malesia and
308
309
310
Australia supports the conclusion that there is only one highly variable and widely
distributed species.
Ecology. Fairly common element of lowland secondary forests or of depauperate, disturbed
primary forests throughout the area. Maiden (l.c.) reports that the trees grow well on poor
sandy soils.
Uses. Braid (l.c.) states that the species is used as fodder plant. He also records its medicinal
properties and notes that it is valuable for tanning.
2. COLUBRINA Rich. ex Brongn.

(Latin, colubrinus = like a snake; perhaps from the French name bois couleuvre,
the snake wood or serpent tree, C. ferruginosa, from Martinique Island)
Ann. Sci. Nat. 10 (1827) 61; Ridley l.c. 465; Backer & Bakhuizen f. l.c. 84; Johnston, Brittonia 23
(1971) 2; Latiff l.c. 298; Whitmore, Tantra & Sutisna l.c. 293.
Deciduous or evergreen trees and shrubs, rarely scandent. Stipules minute, early deciduous.
Leaves alternate to rarely opposite, chartaceous to subcoriaceous; margin entire, serrate or
crenate; venation pinnate or acrodromous from the base. Inflorescence an axillary cyme or
small thyrse, sessile or shortly peduncled, mainly few-flowered. Flowers usually 5-merous,
bisexual, perigynous; hypanthium hemispherical; sepals 5, triangular, rather densely pubescent
outside, conspicuously keeled within, deciduous; petals 5, clawed; stamens 5, about as long
as petals, dehiscing introrsely; disc massive, fleshy; ovary semi-inferior, 3(4)-locular,
styles 3-lobed to trifid, stigmas 3. Fruit a subglobose, 3-lobed capsule; mesocarp thin, dry
or rather fleshy; endocarp cartilaginous to woody, splitting into 3 ventrally dehiscent endocarpids. Seeds broadly obovate, shiny, rarely with a minute basal aril, surface smooth or
pitted; testa coriaceous to bony; endosperm fleshy, thick.
Distribution. An almost pantropical genus of about 24 species, mainly centered in the
Neotropics. Two species (C. asiatica and C. beccariana) are found in Sabah and Sarawak.
Uses. The very hard and heavy wood of several neotropical species, known as West
Ironwood (C. arborescens, C. elliptica) is of some local importance.
Key to Colubrina species

Small to medium-sized tree. Leaves subcoriaceous, oblong-elliptic to oblong-obovate, 3veined from the acute base.............................................................................C. beccariana
Small coastal shrub. Leaves chartaceous, broadly ovate to cordate, pinnately veined.............
C. asiatica (L.) Brongn.
l.c. 62; Masamune l.c. 434. Basionym: Ceanothus asiaticus L., Sp. Pl. (1753) 196.
Sprawling shrub. Leaves ovate to cordate, very thin, pinnately veined, margin dentate.
Flowers in few-flowered thyrses. Fruit a globose capsule with explosive dehiscence.
India, China, throughout Malesia. In Sabah and Sarawak scattered in coastal area, and
collected from Pulau Tiga FR (FDBNB 38625) and Timbun Mata FR (FDBNB 44652)
in Sabah, and Telok Pinang, 1st Div. in Sarawak (S. 41941).
311
Colubrina beccariana Warb.
Fig. 2.

Bot. Jahrb. Syst. 13 (1891) 367; Ridley l.c. 465; Johnston l.c. 44; Latiff l.c. 298; Whitmore, Tantra &
Sutisna l.c. 293. Type: Warburg 20187, Dutch New Guinea (holotype B, destroyed; isotype A).
Synonym: C. anomala King, J. As. Soc. Beng. 45 (1896) 377.
Small to medium-sized tree to 25 m high, bole up to 20 m, diameter to 50 cm, sometimes

with weak buttresses. Bark smooth or slightly dimpled, very brittle, reddish brown to dark
brown; inner bark light red, pale yellow near cambium. Wood very hard, sapwood pale
yellow. Young branches, axillary buds, petioles, major leaf venation, peduncles, pedicels
and gynoecium densely rusty pubescent. Stipules subulate, to 4 mm long, early deciduous.
Leaves alternate, subcoriaceous, oblong-elliptic to oblong-obovate, 1022 x 3.58.5 cm;
base acute, margin slightly revolute, entire to finely serrulate, individual teeth minute,
appressed, bearing a black, glandular apex, apex acuminate, acumen 12 cm long and 23
mm wide, apically rounded; venation pinnate, slightly impressed above, more or less
prominent below; lateral veins 46 pairs, basal ones extending into the upper half of the
lamina; intercostal veins rectangular to the midrib, strongly percurrent between
superadjacent lateral veins; petioles 615 mm long. Inflorescences axillary fascicles or
shortly peduncled cymes; pedicels 13 mm long. Flowers 5(6)-merous; hypanthium
shallow patelliform; sepals triangular to acuminate, distinctly keeled within; petals obovate,
cucullate; disc fleshy, slightly grooved; ovary immersed in the fleshy disc, densely brownish
pubescent, 23-locular; style more or less deeply bi- to tri-fid, stigmas 3, slightly capitate.
Fruits globose, 1016 mm in diameter, glabrous; exocarp flaky, black when dry; mesocarp
thin; endocarp woody, splitting into 3 explosively dehiscing endocarpids when ripe;
pedicels to 2 cm long. Seeds 810 mm across, testa glossy, reddish brown.
Vernacular names. Sabahobar-obar (Dusun), odok-odok (Kadazan), udok-udok (Malay).
Distribution. Peninsular Malaysia and Borneo (Sabah and Kalimantan). In Sabah, collections
have been made from Kinabatangan, Lamag, and Sandakan (Kabili-Sepilok FR) districts.
Ecology. A typical lowland species, mainly found in undisturbed primary rainforest.
3. RHAMNUS L.
(after the ancient Greek plant name rhamnos)
Sp. Pl. (1753) 193; Masamune l.c. 434; Anderson l.c. 290; Latiff l.c. 298. Synonym: Oreorhamnus
Ridl., J. Fed. Malay. St. Mus. 10 (1920) 131.
Deciduous or evergreen trees and shrubs, rarely climbing, often thorny. Stipules small,
caducous. Leaves alternate or rarely opposite, glabrous to rather densely pubescent,
membranous to coriaceous, venation pinnate; margin entire to toothed. Inflorescences
axillary cymes, sessile or peduncled, rarely reduced to a solitary flower. Flowers perigynous,
bisexual or unisexual by abortion; hypanthium campanulate to urceolate; sepals 45,
312
deltoid, valvate in bud; petals 45 or rarely wanting, minute, usually shortly clawed,
concave to hooded; stamens 45, sterile and rudimentary or completely wanting in female
flower; disc nectariferous, lining the floral tube; ovary superior, 23(4)-locular, each
locule with one solitary, erect, anatropous ovule; stigma 3-lobed or stigmata 23. Fruit a
globular to obovoidal drupe with 23(4) endocarpids; mesocarp fleshy to coriaceous,
often with tanniferous idioblasts or mucilage cavities; endocarp cartilaginous to woody;
endocarpids abaxially convex, adaxially slightly angular to almost flat, dehiscent or
indehiscent. Seeds narrow ellipsoidal to obovoidal, rarely suborbicular; testa membranous;
endosperm fleshy, scanty; embryo large.
Distribution. The genus extends over nearly the whole area of the family, being abundant
in Eastern Asia and Southwestern North America, rather scanty in Europe and Africa, and
absent from Madagascar, Australia and Polynesia; one species in Sabah and Sarawak.
Ecology. Primarily distributed in the temperate and subtropical regions of both hemispheres, the genus extends into tropical highlands and is found even in semiarid to arid
places. Of significance is the obvious avoidance of hot tropical lowlands.
Uses. Several species are grown as ornamentals or hedge-plants, other furnish wood of local
importance or drugs (e.g., R. cathartica).
Rhamnus borneensis Steenis
Fig. 3.
(of Borneo)
J. Bot. 72 (1934) 6; Masamune l.c. 434; Anderson l.c. 290; Latiff l.c. 298. Type: Clemens 27876,
British North Borneo, Mt. Kinabalu (L). Synonyms: R. lancifolia Steenis l.c. 7, Anderson l.c. 290; R.
borneensis Steenis var. borneensis Latiff l.c. 299.
Small tree to 15 m high; crown small; bole straight, diameter to 20 cm. Bark rugulose; inner
bark yellow, fibrous, without latex. Sapwood silky white, heartwood dark brown. Young
twigs, axillary buds, stipules, petioles, pedicels, flower buds and lower surface of the leaves
densely yellowish pubescent, indument composed of stellate hairs; branches soon
glabrescent, with smooth, red to red-brown bark, without lenticels. Leaves lanceolateelliptic to obovate, chartaceous to subcoriaceous, shiny and glabrous above, densely whitish
to yellowish pubescent below, 7.514 x 35.5 cm; base acute, margin slightly revolute,
entire to finely serrulate, apex rounded or acute to acuminate, acumen blunt or broadly
triangular; lateral veins 711 pairs, straight to slightly curved, parallel, ex-medially
anastomosing with each forming well-developed marginal loopes. Inflorescences axillary
fascicles or shortly peduncled, few-flowered cymes. Flowers 45 mm in diameter; pedicels
47 mm long; hypanthium hemispheric to campanulate; sepals deltoid to acute triangular,
1.52 mm long and wide, distinctly keeled within; petals cucullate, clawed, distinctly 2lobed, 1.52 mm long; stamens 11.5 mm long; disc thin, lining the hypanthium, glabrous;
ovary very small, conical to globose, 3-locular, glabrous; style undivided, cylindric, stigma
slightly 3-partite, inconspicously capitate. Fruits red, globose to obovoid, with 23 endocarpids; mesocarp coriaceous; endocarp stony, endocarpids lenticular. Seeds elliptic to
cylindric.
313
Distribution. Sumatra, Peninsular Malaysia and Borneo. Uncommon; in Sabah known from
a few collections (e.g., Clemens 27876 and SAN 56276 from Mt. Kinabalu area), and in
Sarawak scattered and confined to montane forest (e.g., Nooteboom & Chai 1996 from Mt.
Murut; S. 35822 and S. 38139 from Mt. Mulu; and S. 47355 from Mt. Berumput). No record
from Brunei and Kalimantan.
Ecology. A species of submontane to montane ridge and mossy forests to 2700 m.
4. ZIZIPHUS Mill.
(from the Persian plant name zizuf )
Gard. Dict. Abr. ed. 4 (1754); Ridley l.c. 461; Merrill, PEB (1929) 177; Masamune l.c. 435; Backer &
Bakhuizen f. l.c. 81; Anderson l.c. 290; Corner l.c. 610; Latiff l.c. 299; Whitmore, Tantra & Sutisna
l.c. 293.
Deciduous or evergreen, small to medium-sized trees, erect or straggling shrubs or to 30 m

high woody climbers, often armed. Stipules small, deciduous or often transformed into
straight or recurved spines. Leaves alternate, usually distinctly 3-veined from the base,
rarely pinnately veined, often slightly asymmetric; margin entire or serrate to crenate.
Inflorescences axillary fascicles or sessile or peduncled, umbel-like cymes, rarely terminal
or axillary thyrses. Flowers perigynous, bisexual, yellowish to greenish; hypanthium
shallow patelliform to hemispherical; sepals 5, keeled inside, early deciduous; petals 5,
cucullate, clawed, rarely absent; disc nectariferous, fleshy, mainly flat, 5- to 10-lobed, lining
the hypanthium; ovary 23(4)-locular, styles 24, stigma 23(4), very small, slightly
capitate. Fruits drupaceous, fleshy or almost dry, exclusively one-stoned; mesocarp fleshy
to leathery; endocarp bony to woody, surface smooth to furrowed. Seeds ellipsoid, nearly
plano-convex, raphe lateral, testa thin, membranous; endosperm scanty, fleshy; embryo
straight.
Distribution. An almost pantropical genus of about 100 species centered in tropical
America and in Southeastern Asia. In Sabah and Sarawak, 8 species, which are mostly
thorny straggling shrubs or woody climbers of the lowlands.
Ecology. The genus is widely distributed in tropical and subtropical regions of the world,
extending locally into temperate or even semi-arid to arid zones.
Uses. Two species, Z. jujuba Mill. and Z. mauritiana Lam. are widely cultivated for their
edible fruits. The wood of some other species is of local importance, e.g., for housebuilding, furnitures, cabinet-making, and for agricultural tools.
314
315
316
Key to Ziziphus species

1.
Small to medium-sized unarmed tree. Leaves to 30 cm long.................Z. angustifolius

Erect or straggling shrub or woody climber with stipulary spines. Leaves smaller........2
2.
Leaves less than 10 cm long.........................................................................................3

Leaves 1017 cm long.................................................................................................7
3.
Fruits ellipsoid, with flattened base and acuminate apex, to 4 cm long...........................

Z. borneensis Merr.
l.c. (1929) 178; Masamune l.c. 435; Whitmore, Tantra & Sutisna l.c. 294.
Straggling shrub or woody climber with recurved stipulary spines, plant often
densely ferruginous-tomentose. Leaves elliptic to oblong-elliptic, chartaceous to
subcoriaceous; base often distinctly asymmetric, margin entire to minutely dentate,
apex acuminate. Flowers in axillary fascicles. Fruits a very large ellipsoid drupe.
Endemic to Borneo (Sabah, Sarawak, Brunei, Kalimantan).
Fruits globose to obovoid, to 1 cm in diameter.............................................................4
4.
Leaves very thin, venation pattern of the lower surface dark coloured and typically
distinct. Style undivided and glabrous............................................................................
Z. cumingiana Merr.
Philipp. J. Sci. 1, Suppl. 3 (1906) 206.
Erect or straggling shrub with one recurved stipulary spine per node. Leaves ovate,
often distinctly asymmetric, very thin; margin finely dentate. Flowers in fewflowered axillary cymes, very small. Drupes glabrous. Borneo (Sabah) and the
Philippines.
Leaves chartaceous to coriaceous, venation pattern never distinctly coloured. Styles 2(
3)-lobed, densely pubescent..........................................................................................5
5.
Leaves with distinct arcs of veins diverging from the lateral veins.................................
Z. horsfieldii Miq.
Fl. Ned. Ind.1, 1 (1855) 643; Masamune l.c. 435; Latiff l.c. 300; Whitmore, Tantra &
Sutisna l.c. 294.
Straggling shrub with one short stipulary spine per node. Leaves ovate to elliptic,
very thin, often somewhat asymmetric; base obtuse, margin minutely dentate, apex
shortly acuminate. Flowers in axillary, dichotomously branched cymes. Drupes
globose, densely brownish pubescent. Sumatra, Peninsular Malaysia, Java, Borneo
(Sabah, Sarawak, Brunei, Kalimantan).
Leaves without such lateral veins.................................................................................6
6.
Leaves bearing a distinct intra-marginal vein.................................................................

Z. havilandii Ridl.
Kew Bull. (1931) 495; Masamune l.c. 435; Whitmore, Tantra & Sutisna l.c. 294.
Straggling shrub with one recurved stipulary spine per node, plant more or less
densely yellowish pubescent, glabrescent. Leaves ovate, often asymmetric, very
thin; base obtuse, margin minutely dentate. Flowers in dichotomously branched,
axillary cymes. Drupes globose to obovoid, densely yellowish to brownish
pubescent. Endemic to Borneo (Sabah, Sarawak, Brunei, Kalimantan).
317
Leaves without any intramarginal vein..........................................................................

Z. suluensis Merr.
Philipp. J. Sci. 30 (1926) 408, l.c. (1929) 177; Masamune l.c. 435; Whitmore, Tantra &
Sutisna l.c. 294.
Straggling shrub with one recurved stipulary spine per node. Leaves elliptic, subcoriaceous; base obtuse to rounded, often somewhat asymmetric, margin minutely
dentate, apex acuminate. Flowers in small, axillary, few-flowered cymes. Drupes
globose to obovoid, densely brownish pubescent. Sumatra, Borneo, Philippines and
the Moluccas.
7.
Leaves chartaceous to subcoriaceous, venation pattern typically prominent on the lower

surface. Bark of twigs cream-coloured, densely lenticellate............................................
Z. crebrivenosa C.B. Rob.
Philipp. J. Sci. 3 (1908) 201; Whitmore, Tantra & Sutisna l.c. 294.
Woody climber with one recurved stipulary spine per node and densely lenticellate
branches. Leaves obovate to elliptic, often somewhat asymmetric, chartaceous;
base obtuse to rounded, margin faintly denticulate, apex acute to obtusely
acuminate and mucronate. Flowers in axillary cymes. Drupes globose, to 3 cm
diameter, densely lenticellate. Borneo (Sabah, Sarawak, Kalimantan), Philippines
and Celebes.
Leaves coriaceous, venation pattern very delicate, never prominent. Bark of twigs
brownish, not lenticellate...............................................................................................
Z. calophylla Wall.
in Roxb., Fl. Ind. 2 (1824) 366; Merrill l.c. (1926) 364, l.c. (1929) 177; Ridley, Kew Bull.
(1931) 494; Masamune l.c. 435; Latiff l.c. 300; Whitmore, Tantra & Sutisna l.c. 294.
Sturdy straggling shrub, with short, solitary or paired stipulary spines. Leaves
coriaceous, elliptic to oblong-elliptic, base usually narrowed, margin minutely
dentate, apex acuminate. Flowers in axillary or terminal cymes. Drupes globose,
densely brownish tomentose. Sumatra, Peninsular Malaysia and Borneo (Sabah,
Brunei, Kalimantan).
Ziziphus angustifolius (Miq.) Hatusima ex Steenis
Fig. 4.
(Latin, angustifolius = having narrow leaves)

Nova Guinea, Botany 3 (1960) 13; Hatusima, Cat. Hort. Bot. (1957) 242; Merrill & Perry, J. Arn. Arb.
20 (1939) 337; Anderson l.c. 290; Latiff l.c. 299; Whitmore, Tantra & Sutisna l.c. 293. Basionym:
Solenostigma angustifolium Miq., Fl. Ned. Ind., Suppl. (1861) 412. Type: Miquel HB 758 and HB
2501, Sumatra West Coast (syntypes L). Synonyms: Celtis angustifolia (Miq.) Planch. in DC., Prod.
17 (1873) 186; C. grewioides Warb., Bot. Jahrb. 13 (1891) 287; Ziziphus inermis Merr., Govt. Lab.
Publ. Philipp. 35 (1906) 37; Z. forbesii Baker f., J. Bot. 61, Suppl. (1923) 10; Z. grewioides (Warb.)
Perry ex Steenis, Blumea 7 (1954) 595.
Small to medium-sized, unarmed tree to 30 m high and 40 cm diameter; crown narrow

conical; bole straight, rarely crooked, up to 20 m high, sometimes with buttresses of about 1
m high and 11.5 m out. Bark not fissured, not peeling, with densely spaced lenticels,
greyish to brownish, about 0.2 mm thick; inner bark yellow to brownish, blaze strawcoloured, cambium orange. Sapwood hard, light to greyish brown, to 15 cm thick;
heartwood hard and heavy, dark brown. Young twigs, axillary buds, stipules, petioles,
peduncles, pedicels and flower buds rather densely yellowish to brownish pubescent; older
branches glabrescent, greyish to brownish, lenticellate. Stipules minute, narrow lanceolate,
318
not transformed into spines, 12 mm long, early deciduous. Leaves alternate, chartaceous to
subcoriaceous, symmetric, oblong-elliptic to oblong-ovate, 1220 x 37 cm; base obtuse to
acute, margin slightly revolute, finely serrate, apex acute to acuminate, acumen about 12
cm long, 0.5 cm wide, apically distinctly mucronate; venation 3-veined from the base;
intercostal veins rectangular to the midrib, strongly parallel between the midrib and the
lateral veins, outer intercostal veins anastomosing to form a slightly looped marginal vein.
Inflorescences axillary, peduncled, branched cymes, to 3 cm long. Flowers 46 mm in
diameter, 5-merous, greenish to yellowish; pedicels 24 mm long; hypanthium patelliform;
sepals triangular, keeled within; petals clawed, cucullate; stamens about 1.5 mm long,
filaments pale greenish, anthers green; disc prominent, glandulous, glabrous, yellow to
ochre; ovary immersed in the disc, 23-locular, style 23-lobed, increasingly diverging
during anthesis, stigmas distinctly papillous. Fruits globose to obovoid, often 1-seeded,
1020 mm diameter; mesocarp comparatively thin; endocarp woody.
Vernacular names. Sabahkayu labu (Kadazan), pasil-pasil (Bajau).
Distribution. India, Burma, Thailand, Sumatra, Peninsular Malaysia, Borneo (throughout),
Philippines, Celebes, Moluccas, New Guinea, and Solomon Island. In Sabah recorded from
Kinabatangan, Tenom, Tongod, Lahad Datu, and Sandakan districts; and in Sarawak from
various localities in the 1st, 4th and 7th Divisions. Also occurs in Brunei and Kalimantan.
Ecology. Lowland mixed dipterocarp forest to c. 600 m.
319
RHIZOPHORACEAE
L. Madani & K.M. Wong
Sandakan, Malaysia
Merrill, EB (1921) 420; Masamune, EPB (1942) 515; Browne, FTSB (1955) 296; Ding Hou, FM 1, 5
(1958) 429; Burgess, TBS (1966) 431; Ashton, MNDTS 2 (1988) 342; Juncosa & Tomlinson, Ann.
Missouri Bot. Gard. 75 (1988) 1278; Kochummen, TFM 4 (1989) 302; Whitmore, Tantra & Sutisna,
CLK 2, 1 (1990) 295.
Trees or shrubs, the mangrove species often with characteristic root formations (branching
stilt-roots in Rhizophora, knee pneumatophore-roots in Bruguiera). Bark invaginating finely
and regularly into the wood in the inland genera (Carallia, Gynotroches, Pellacalyx). Twigs
solid or (in Gynotroches and Pellacalyx) hollow; branch nodes swollen. Leaves opposite
and decussate, entire or toothed, pinnately veined; stipules interpetiolar, caducous, with
colleters on the inner side. Inflorescences usually axillary, cymose or a fascicle of flowers
or flowers solitary. Flowers bisexual or (in Gynotroches) unisexual (the plants dioecious),
radially symmetrical; calyx-lobes 416, valvate; petals 416, free, fleshy, 2-fid to fimbriate
or fringed with apiculate appendages or lacerate or lobed (rarely entire); stamens free or
(in Pellacalyx) inserted on the calyx-tube, 13 times the number of petals or (in Kandelia)
many; anthers 4-celled or (in Kandelia) many-celled, dorsifixed, splitting lengthwise or (in
Rhizophora) opening by a ventral valve; ovary superior or semi-inferior or inferior, 212celled (cells often incompletely or not separated at floral maturity and then apparently 1celled), style simple, stigma capitate or lobed, ovules 2many per cell, pendulous,
placentation axile; disc usually present, fleshy, annular, entire to lobed. Fruit a berry, calyx
persistent. Seeds 1many per fruit, pendulous, endospermous; embryo with laminar
cotyledons; germination epigeal or in the mangrove species viviparous (while the seed still
enclosed in the fruit and the fruit still attached to the tree).
Distribution. 15 genera with c. 145 species, pantropical. In Sabah and Sarawak 7 genera
with 23 species.
Ecology. The mangrove species occur on coastlines with warm currents, and many also
grow in brackish water conditions within the estuaries or along the lowest stretches of
rivers. The inland species are mostly of moist (primary or secondary) tropical forests.
Uses. The mangrove species have been exploited for poles used in scaffolding and the
construction of large fishing platforms and traps. The bark of almost all mangrove species
(in Bruguiera, Ceriops and Rhizophora) are used for tanning. The mangrove species are
also important in providing poles for the local charcoal industry, the calorific value of the
wood being excellent. In the inland forests, Carallia species grow to timber size and are
exploited for logs.
321
Taxonomy. Rhizophoraceae have been traditionally placed in the Myrtales and sometimes
with the Cornales. This was based on the incorrect assumption that the Rhizophoraceae
typically have an inferior ovary, whereas in fact this is a rare and derived condition.
Sometimes, also based on a single character of the presence of interpetiolar stipules, the
Rhizophoraceae has been said to be related to the Rubiaceae and Cunoniaceae. Dahlgren
(1988) (Ann. Missouri Bot. Gard. 75: 1259), after considering a number of characters,
places the Rhizophoraceae in the Celastrales, together with Celastraceae and Elaeocarpaceae.
Cronquist (1981) (An Integrated System of Classification of Flowering Plants) separates the
Rhizophoraceae sensu stricto in its own order, the Rhizophorales.
Although a number of authors have treated the genera Anisophyllea and Combretocarpus as
part of the Rhizophoraceae, it is now well established that these two genera should be
separately classified in a family, the Anisophylleaceae (this volume). Within the Rhizophoraceae
s.s. Juncosa & Tomlinson l.c. consider three tribes, Rhizophoreae (Bruguiera, Ceriops,
Kandelia, Rhizophora), Gynotrocheae (Carallia, Gynotroches, Pellacalyx, Crossostylis) and
Macarisieae (which includes seven genera not in our area).
Some Rhizophoraceae can be confused with the Rubiaceae at a glance, as both typically
have opposite leaves with interpetiolar stipules. The Rubiaceae have fused corolla members
and have a corolla-tube, and the stamens are as many as the corolla-lobes. In the
Rhizophoraceae, the corolla members (petals) are free, and frequently the stamens are at
least twice the number of petals.
Key to genera
1.
Plants of mangrove forests. Bark not invaginating into wood. Seeds germinating while
still in the fruits and the fruits still attached to the tree (viviparous)............................2
Plants of inland forests. Bark invaginating finely and regularly into wood. Seeds
germinating only after the fruits have fallen off the tree...............................................5
2.
Leaf apex pointed........................................................................................................3

Leaf apex blunt or rounded..........................................................................................4
3.
Plants developing buttresses and knee pneumatophores (breathing roots). Calyx-lobes

816, lanceolate-linear..............................................................................1. Bruguiera
Plants developing conspicuous branching stilt-roots. Calyx-lobes 4, broadly ovate
...............................................................................................................7. Rhizophora
4.
Plants developing only short buttresses, trunk not usually thickened at base. Leaves
broadly elliptic to obovate, length at most twice the width. Hypocotyl (first elongating
axis) in the germinating fruit with sharp longitudinal ridges.........................3. Ceriops
Plants typically without buttresses, trunk developing a conically thickened base. Leaves
narrowly elliptic-oblong, length at least 2.53-times the width. Hypocotyl in the
germinating fruit smooth............................................................................5. Kandelia
5.
Twigs solid. Leaf-margin distinctly toothed or crenate. Inflorescence cymose, with a

distinct main stalk........................................................................................2. Carallia
322
RHIZOPHORACEAE (MADANI & WONG)
Twigs hollow. Leaf-margin faintly toothed to entire. Inflorescence a fascicle of flowers

without any distinct main stalk....................................................................................6
6.
Stipules with overlapping margins. Ovary superior (calyx at the base of the fruit)
.............................................................................................................4. Gynotroches
Stipules not overlapping, flat to slightly spreading. Ovary inferior (calyx at the apex of
the fruit)....................................................................................................6. Pellacalyx
1. BRUGUIERA Lam.
(J.G. Bruguieres, 17501798, who worked for Lamarcks Encyclopaedia Methodica)
beus, lenggadai (Malay)
Tab. Enc. Meth. (1797) t. 397; Merrill l.c. (1921) 421; Masamune l.c. 515; Browne l.c. 299; Ding Hou
l.c. 457; Burgess l.c. 431; Ashton l.c. 349; Kochummen l.c. 309; Whitmore, Tantra & Sutisna l.c. 295.
Medium-sized trees with buttresses and knee pneumatophores, sometimes with aerial roots
when young. Leaves usually coriaceous, entire, glabrous, finely black-dotted on the lower
side; stipules lanceolate. Inflorescence a 25-flowered cyme or reduced to a solitary flower.
Flowers bisexual, articulate at the base with the stalk; calyx thick, 815-lobed, lobes
lanceolate-linear, acute; petals 2-lobed, falling off early; disc cup-shaped and attached to the
calyx-tube; stamens twice the number of petals and paired; ovary inferior, 24-celled,
ovules 2 per cell, style filiform, stigma obscurely 24-lobed. Fruits included in and adnate
to the calyx-tube, usually 1-celled, 1(2)-seeded; germination viviparous; hypocotyl rounded or obscurely ribbed, blunt.
Distribution. 6 species, tropical east Africa to Asia, throughout Malesia to Australia and
Polynesia. In Sabah and Sarawak, 4 species.
Ecology. In mangroves, usually occurring behind Rhizophora stands that are inundated.
Forming pure stands in some sites.
Uses. Not suitable for poles and for construction. The wood, however, is used for making
charcoal.
Key to Bruguiera species

1.
Flowers solitary. Bark in mature trees usually fissured-flaky...................................2

Flowers several in a cyme. Bark usually smooth or lenticellate.....................................3
2.
Flower-stalks 1012 mm long; tips of petal acute, each with 3 or 4 distinct bristles
........................................................................................................2. B. gymnorrhiza
Flower-stalks 68 mm long; tips of petals obtuse, each with 13 bristles......4. B.
sexangula
323
3.
Flowers small, the calyx 34 mm long, smooth; stalks 23 mm long. Fruit with reflexed
calyx-lobes..............................................................................1. B. cylindrica
Flowers larger, the calyx 610 mm long, ribbed; stalks 58 mm long. Fruit with erect
calyx-lobes............................................................................................3. B. parviflora
1. Bruguiera cylindrica (L.) Blume

(Latin, cylindricus = cylindric; the hypocotyl)
En. Pl. Jav. 1 (1827) 93; Browne l.c. 300; Ding Hou l.c. 467; Burgess l.c. 432; Ashton l.c. 353;
Kochummen l.c. 310; Whitmore, Tantra & Sutisna l.c. 295. Basionym: Rhizophora cylindrica L., Sp.
Pl. (1753) 443. Type: Kari-kandel Rheede, Hort. Mal. 6 (1686) 59, t. 33. Synonyms: R.
caryophylloides Burm. f., Fl. Ind. (1768) 109; R. ceratophylloides Gmel., Syst. Veg. (1796) 749; B.
caryophylloides (Burm. f.) Blume l.c. (1827) 93; B. malabarica Arn., Ann. Mag. Nat. Hist. 1 (1838)
369; Kanilia caryophylloides (Burm. f. ) Blume, Mus. Bot. Lugd. Bat. 1 (1849) 141.
Tree to 25 m tall, 45 cm diameter; buttresses to 1 m high. Bark grey, with many large
prominent lenticels; inner bark pink. Sapwood yellowish brown. Leaves elliptic, 717 x 2
8 cm, thinly coriaceous; base cuneate, apex acute; lateral veins c. 7 pairs, distinct to obscure
on both sides; stalk 14.5 cm long; stipules 2.53.5 cm long. Flowers in 3-flowered cymes;
stalks 23 mm long; calyx-tube smooth, 34 mm long, lobes 8, as long as the tube; petals
45 mm long, the lobes 1/3 the length of the petal, apex obtuse, each with 13 bristles to 1.5
mm long, outer margin fringed with white hairs. Fruits with calyx-tube 11.2 cm long, the
calyx-lobes reflexed; hypocotyl cylindric, to 15 cm long, 0.5 cm thick.
Vernacular names. Sabahbeus (Malay). Sarawakberus ngayong (Lundu Malay), berus
puteh (Malay), ngayong (Lundu Malay).
Distribution. SE Asia, throughout Malesia to N Queensland. In Sabah recorded from Pulau
Gaya and Pulau Tiga in the west coast; in Sarawak recorded at Lundu and the Rejang delta.
Ecology. Often gregarious on newly formed sandy clays and behind Avicennia on the
seaface; it is succeeded by other species on better drained sites.
Uses. The timber is mainly used for firewood and is not durable in contact with soil, hence it
is not favoured as a pole in house construction.
2. Bruguiera gymnorrhiza (L.) Lam.

(Greek, gymnos = naked, rhiza = root; the exposed germinating hypocotyl)
Encycl. Meth. Bot. 4 (1798) 696; Browne l.c. 301; Ding Hou l.c. 461; Burgess l.c. 431; Ashton l.c.
350; Kochummen l.c. 310; Whitmore, Tantra & Sutisna l.c. 296. Basionym: Rhizophora gymnorrhiza L.,
Sp. Pl. (1753) 433. Type: Kandel Rheede, Hort. Mal. 6 (1686) 57, t. 31 & 32. Synonyms: R. palun
DC., Prod. 3 (1828) 33; B. rheedii Blume l.c. (1827) 92; R. tinctoria Blanco, Fl. Filip. (1837) 394; B.
capensis Blume l.c. (1849) 137; B. wightii Blume l.c. (1849) 138; B. rumphii Blume l.c. (1849) 138;
324
B. cylindrica sensu Hance (non Blume), J. Bot. 18 (1879) 10; B. conjugata sensu Merr. (non R.
conjugata L.), Philip. J. Sc. 9 (1914) Bot. 118, l.c. (1921) 421, Masamune l.c. 515.
Tree to 35 m tall, 45 cm diameter; often with buttresses to 1 m high. Bark dark grey to
brown to black, deeply fissured and roughly flaky, lenticels conspicuous only when young;
inner bark yellowish pink. Sapwood pale yellowish brown. Leaves elliptic-oblong, 820 x
47 cm, thinly coriaceous; base cuneate or rarely obtuse, apex acute; lateral veins 910
pairs, indistinct on both sides; stalk 24.5 cm long; stipules 34 cm long. Flowers solitary;
stalks 1012 mm long; calyx-tube distinctly ribbed at upper part, 812 mm long, lobes
1016, 1722 mm long; petals 1315 mm long, the lobes 1/31/2 the length of the petals,
apex acute, each with 3 or 4 bristles of 23 mm long, outer margin fringed with white silky
hairs. Fruits with calyx-tube to 2.5 cm long, the calyx-lobes ascending, not reflexed;
hypocotyl slightly ribbed, to 25 cm long, 2 cm thick.
Vernacular names. Sabahputut (Malay). Sarawakberus kurong, berus merah, kurong
(Malay).
Distribution. Tropical east Africa, S and SE Asia to the Ryukyus, throughout Malesia to
Australia, Micronesia and Polynesia. In Sabah and Sarawak all districts with a coastline.
Ecology. One of the largest trees of the mangrove, found at the inland margins.
Uses. The wood is hard, difficult to work and not ornamental, although it is easily
impregnated with preservative. It can be used for rail sleepers, and is locally used in
charcoal making and for firewood, and the poles are sometimes used in house construction.
The bark has been used for the manufacture of cutch but is said to be inferior to that from
Rhizophora.
3. Bruguiera parviflora (Roxb.) Wight & Arn. ex Griff.
Fig. 1.
(Latin, parvus = little, flores = flowers)

Trans. Med. Phys. Soc. Calc. 8 (1836) 10; Merrill l.c. (1921) 421; Masamune l.c. 515; Browne l.c.
301; Ding Hou l.c. 464; Burgess l.c. 431; Ashton l.c. 353; Kochummen l.c. 310; Whitmore, Tantra &
Sutisna l.c. 296. Basionym: Rhizophora parviflora Roxb., Fl. Ind. ed. Carey 2 (1832) 461. Type:
Roxburgh no. 1246 (K). Synonyms: R. cylindrica sensu Roxb. (non L.), Hort. Beng. (1814) 36;
Kanilia parviflora (Roxb.) Blume l.c. (1849) 140; B. ritchiei Merr., Publ. Govt. Lab. Philip. 6 (1904)
11.
Tree to 30 m tall, 45 cm diameter. Bark pale grey mottled, pale brown, smooth with small
obscure lenticels; inner bark pinkish red. Sapwood pale brown. Leaves elliptic-oblong,
815 x 2.55 cm, thinly coriaceous; base cuneate, apex acute; lateral veins 910 pairs,
slight-ly distinct on both sides; stalk 0.61.2 cm long; stipules 36 cm long. Flowers 25 in
a cyme; stalks 58 mm long; calyx-tube distinctly ridged to the base, 610 mm long, lobes
1012, 23 mm long; petals 23 mm long, the lobes 1/2 the length of the petals, apex obtuse
and reflexed, each with 13 bristles to 1.3 mm long, outer margin fringed with white silky
hairs. Fruits with calyx-tube to 4 cm long, the calyx-lobes ascending, not reflexed;
hypocotyl angular, to 6 cm long, 1.5 cm thick.
325
Vernacular names. Sabahbeus (Malay), langarai (Idahan, Suluk), lanjing lanjing

(Kedayan), lenggadai (Malay). Sarawakberus lenggadai, lenggadai (Malay).
Distribution. Sri Lanka, SE Asia, throughout Malesia, New Britain. In Sabah and Sarawak
common in mangroves.
Ecology. Often mixed with Rhizophora and occasional along the margins of mangrove
channels.
Uses. The wood lasts only a year in contact with soil in the tropics. A minor source of
charcoal wood and firewood, unpopular as a pole.
4. Bruguiera sexangula (Lour.) Poir

(Latin, sex = six, angulus = angle; the ridged hypocotyl)
Encycl. Supp. 4 (1816) 262; Merrill l.c. (1921) 422; Masamune l.c. 515; Ding Hou l.c. 463; Burgess
l.c. 431; Ashton l.c. 351; Kochummen l.c. 312; Whitmore, Tantra & Sutisna l.c. 296. Basionym:
Rhizophora sexangula Lour., Fl. Coch. (1790) 297. Type: Loureiro, s.n. (BM). Synonyms: B.
angularis Spreng., Syst. Veg. 2 (1825) 602; B. cylindrica sensu Blume (non R. cylindrica L.) l.c.
(1827) 93; R. polyandra Blanco, Fl. Filip. (1837) 396; R. plicata Blanco l.c. 398; B. eriopetala Wight
& Arn. ex Arn., Ann. Mag. Nat. Hist. 1 (1838) 368, Browne l.c. 301; B. australis A. Cunn. ex Arn. l.c.
368; R. australis (A. Cunn. ex. Arn.) Steud., Nomencl. ed. 2, 2 (1841) 449; R. eriopetala Steud. l.c.
449; B. parietosa Griff., Notul. 4 (1854) 670; B. 10-angulata Griff. l.c. 669; B. oxyphylla Miq.,
Sumatra (1861) 324; B. malabarica sensu F.-Vill. (non Arn.), Nov. App. (1880) 79.
Tree to 40 m tall, 60 cm diameter; buttresses to 1 m high, stilt-roots sometimes developing.

Bark pale grey-brown, conspicuously lenticellate; inner bark pale brown. Sapwood
yellowish brown. Leaves elliptic-oblong to oblanceolate, 814 x 36 cm, thinly coriaceous;
base cuneate, apex acute; lateral veins 711 pairs, slightly distinct to obscure on both sides;
stalk 0.72.5 cm long; stipules 36 cm long. Flowers solitary; stalks 68 mm long; calyxtube distinctly ridged to the base, 1012 mm long, lobes 1012, 1820 mm long; petals c.
15 mm long, the lobes 1/2 the length of the petals, apex obtuse and reflexed, each with 13
bristles, outer margin fringed with white silky hairs. Fruits with calyx-tube 1518 mm long,
the calyx-lobes ascending, not reflexed; hypocotyl angular, to 68 cm long, 1.5 cm thick.
Vernacular names. Sabahmata buaya (Malay). Sarawakberus putut, putut (Malay).
Distribution. Sri Lanka and SE Asia to New Britain and throughout Malesia. In Sabah and
Sarawak common in mangroves. Also in Brunei and Kalimantan.
Ecology. Scattered towards the inland margins of mangroves; on wetter soils together with
B. cylindrica and B. parviflora.
Uses. Mainly for charcoal making and firewood, and also in local house construction.
326
327
2. CARALLIA Roxb., nom. cons.

(from karalli, an Indian plant name)
Pl. Corom. 3 (1811) 8; Merrill l.c. (1921) 421; Masamune l.c. 515; Browne l.c. 305; Ding Hou l.c.
481; Ashton l.c. 291; Kochummen l.c. 312; Whitmore, Tantra & Sutisna l.c. 296. Synonyms:
Symmetria Blume, Bijdr. (182627) 1130; Sagittipetalum Merr., Philip. J. Sc. 3 (1908) Bot. 247.
Small to medium-sized trees, sometimes with small buttresses or stilt-roots. Bark smooth
and hoop-marked, becoming roughened by minute longitudinal fissures and transverse
cracks, sometimes scaly, grey-brown or pinkish brown; inner bark invaginating finely and
regularly into the wood. Sapwood with conspicuous radiating lines. Stipules lanceolate.
Leaves often black-dotted on lower side, margin entire or toothed. Inflorescence a
condensed or lax cyme, or flowers in pairs or solitary. Flowers sessile or stalked, calyxlobes 58, deltoid, acute to acuminate; petals 58, clawed; stamens twice the number of
petals, usually free; disc angular, fleshy; ovary inferior, 58-celled with 2 ovules per cell, or
1-celled with 1012 ovules, styles filiform or slightly conical. Fruits small, ellipsoidobovoid, pulpy, 1-celled, usually 1-seeded. Seeds ellipsoid or reniform, endospermous;
Distribution. c. 11 species, Madagascar, India, Sri Lanka, SE Asia and Malesia to N
Australia. 8 species in Sabah and Sarawak.
Ecology. Mostly lowland forests, sometimes swamps, to hills and ridges to c. 2000 m.
Uses. The timber of some species are suitable for making tool-handles and inferior construction
such as picture-frames and panelling. It is also sometimes used for firewood and charcoal.
Taxonomy. Four species enumerated here are not named but merely indicated by numbers;
their precise naming will depend on the collection of better flowering material and, in one or
two cases, will need to be based on a study of wider scope.
Key to Carallia species

1.
Leaves without conspicuous dark gland-dots all over the blade...............................2

Leaves with conspicuous dark gland-dots all over the blade.........................................5
2.
Leaves not longer than 4 cm; lateral veins 46 pairs, intercostal veins inconspicuous on
both sides. Flowers solitary in each unbranched inflorescence.........8. Carallia sp. 5
Leaves typically longer than 4 cm; lateral veins 611 pairs, intercostal veins
conspicuous and often prominent on both sides. Flowers several to many on clearly
branched cymes...........................................................................................................3
3.
Leaves stiffly leathery, margin coarsely saw-toothed; intercostal veins very prominent
on upper side.......................................................................................6. Carallia sp. 3
Leaves chartaceous to thinly leathery, margin entire to subentire or faintly toothed
towards apex; intercostal veins only slightly prominent on upper side..........................4
4.
Leaf-margin faintly toothed to entire; lateral veins looping towards the margin forming
only one clear submarginal vein. Flowers sessile, on an apparently simple axis (actually
328
a cyme with branches reduced), peduncle nil or very short and

inconspicuous.........................................................................................3. C. coriifolia
Leaf-margin always slightly but distinctly toothed; lateral veins forming 2several
series of angular loops between midrib and margin. Flowers short-stalked, in a
distinctly branched cyme, peduncle 11.8 cm long, distinct.................1. C. borneensis
5.
Leaf lateral veins forming 2several series of angular loops between midrib and
margin; intercostal veins typically net-like; margin typically with distinct fine teeth
(rarely shortly toothed to subentire)......................................................4. Carallia sp. 1
Leaf lateral veins typically looping to form only one distinct submarginal vein;
intercostal veins more or less parallel to the lateral veins; margin either shortly toothed
or entire.......................................................................................................................6
6.
Leaf-margin shortly toothed to subentire. Inflorescence peduncle 12.5 cm long.

Flowers at least shortly stalked..............................................................2. C. brachiata
Leaf-margin entire. Inflorescence peduncle to only 1 cm long at most. Flowers sessile
....................................................................................................................................7
7.
Leaves elliptic to slightly obovate, to 78 cm long at most, chartaceous to thinly

leathery, veins inconspicuous to slightly prominent on upper side........5. Carallia sp. 2
Leaves broad-obovate, larger, stiffly leathery, veins prominent on upper side.................
............................................................................................................7. Carallia sp. 4
1. Carallia borneensis Oliver
Fig. 2.
(of Borneo)
in Hooker, Ic. Pl. (1896) t. 2459; Merrill l.c. (1921) 421; Masamune l.c. 515; Ding Hou l.c. 484;
Burgess l.c. 432; Ashton l.c. 356; Whitmore, Tantra & Sutisna l.c. 296. Type: Creagh, s.n., British
North Borneo (K). Synonyms: Sagittipetalum mindanaensis Merr., Philip. J. Sc. 3 (1908) Bot. 247; S.
palawanense Elmer, Leafl. Philip. Bot. 5 (1913) 1830; Carallia mindanaensis (Merr.) Merr., En.
Philip. 3 (1923) 146.
Small or medium-sized tree to 25 m tall. Bark finely fissured, pale to brownish, inner bark
yellowish brown. Sapwood yellowish brown. Leaves broadly elliptic to obovate, 5.510 x
2.57 cm, chartaceous, without conspicuous dark gland-dots on the blade; base cuneate,
margin distinctly densely short-toothed, apex acute to acuminate to cuspidate; lateral veins
68 pairs, slightly raised on lower side, obscure or distinct above, forming 2several series
of angular loops between midrib and margin; intercostal veins indistinct to slightly distinct
on both sides; stalk 58 mm long; stipules 11.5 cm long. Inflorescences to 3.5 cm long,
(di)trichotomously branched; peduncle 11.8 cm long. Flowers 45 mm long, shortly
stalked; calyx-lobes 6, c. 4 mm long, shortly white-hairy on inner side; petals sagittate, to 5
mm long; filaments filiform, 35 mm long; disc cup-shaped, divided to halfway into deltoid
lobes; ovary 1-celled with 12 ovules, style filiform, stigma capitate. Fruits oblong-ellipsoid,
810 x 46 mm, 1-seeded. Seeds oblong-ellipsoid, 23 mm long.
Vernacular name. Sabahkemuning hutan (Dusun).
329
Distribution. Borneo, Philippines, New Guinea. In Sabah and Sarawak, common throughout.
Ecology. Primary mixed dipterocarp forest, lowlands to about 1000 m, and secondary forest
on sandy soils.
2. Carallia brachiata (Lour.) Merr.

(Latin, brachiatus = having decussate branches; the inflorescence)
Philip. J. Sc. 15 (1919) 249; Masamune l.c. 515; Burgess l.c. 432; Ashton l.c. 356; Kochummen l.c.
313; Whitmore, Tantra & Sutisna l.c. 296. Basionym: Diatoma brachiata Lour., Fl. Coch. (1790) 296.
Type: Loureiro, s.n., Habitat in sylvis Cochinchinae (Merrill, Comm. Lour., 1935, 281, states that
this specimen cannot be found in the BM). Synonyms: Symmetria obovata Blume l.c. (182627)
1131; C. celebica Blume l.c. (1849) 131, Merrill l.c. (1921) 421; C. confinis Blume l.c. (1849) 129,
Merrill l.c. (1921) 421; C. cuspidata Blume l.c. (1849) 129, Merrill l.c.(1921) 421; C. lucida Roxb.,
Corom. Pl. 3 (1811) 8, Merrill l.c. (1921) 421; C. multiflora Blume l.c. (1849) 131; C. timorensis
Blume l.c. (1849) 128; C. cerisopsifolia Miq., Analecta Pt. 3 (1852) 8; C. floribunda Miq., Fl. Ind.
Bat. 1, 1 (1858) 1088; C. calycina Benth., J. Linn. Soc. 3 (1859) 75; Stalagmites lamponga Miq.,
Sumatra (1861) 496; Garcinia cymulosa Miq., Ann. Mus. Bot. Lugd. Bat. 1 (1864) 208; C. arguta
Koord. & Valeton, Bijdr. Booms. Java 4 (1896) 301; C. scortechinii King, J. As. Soc. Beng. 66, 2
(1897) 319; C. spinulosa Ridl., J. Str. Br. R. As. Soc. 82 (1920) 184; C. cuprea Ridl., Kew Bull.
(1938) 282; C. viridifolia Ridl. l.c. (1938) 282.
Medium-sized to large tree to 35 m tall, 70 cm diameter, sometimes buttressed and sometimes with stilt-roots. Bark smooth or scaly-lenticellate, brownish; inner bark yellowish
brown. Sapwood pale yellowish to brownish. Leaves elliptic to obovate, 512.5 x 27.5
cm, chartaceous, with conspicuous dark gland-dots on the lower side when dry; base
cuneate to obtuse, margin shortly toothed to subentire, apex acuminate to cuspidate; lateral
veins 812 pairs, faint to slightly raised on both sides; intercostal veins more or less parallel
to lateral veins, indistinct on both sides; stalk 1.52.5 cm long; stipules 0.81.2 cm long.
Inflorescence a cyme to 2.54 cm long, (di)trichotomously branched, peduncle 12.5 cm
long. Flowers 2.53 mm long, shortly stalked; calyx-lobes (4)68, 11.5 mm long,
glabrous on inner side; petals suborbicular, to 1.52 mm long; filaments filiform, 23 mm
long; disc cup-shaped, divided to halfway into deltoid lobes; ovary 58-celled with 2 ovules
per cell, style filiform, stigma lobed. Fruits globose, to 7 mm across, 1-seeded. Seeds
globose, c. 2 mm across.
Vernacular names. Sarawakrabong, radipah (Melanau Matu and Oya).
Distribution. Widely distributed from Madagascar, Sri Lanka, India, SE Asia and Malesia
to N Australia. In Sabah and Sarawak common throughout. Also in Brunei and Kalimantan.
Ecology. Primary and secondary forests, lowlands to 1800 m; on infertile organic soils in
mixed dipterocarp forest, heath forest, and on the margins of freshwater swamp forests.
Uses. The wood is good for furniture and other interior finishings such as parquet flooring
and panelling, as it is hard and has an attractive oak-like figure; it is, however, difficult to
season.
330
331
3. Carallia coriifolia Ridl.

(Latin, coriaceus = leathery, -folius = -leaved)
l.c. (1938) 283; Ding Hou l.c. 483; Ashton l.c. 357; Whitmore, Tantra & Sutisna l.c. 296. Type:
Haviland 1797, Sarawak, Kuching (holotype K; isotypes L, SAR).
Small tree to 10 m tall. Bark flaky, dark grey; inner bark yellowish. Sapwood yellowish.
Leaves ovate to oblong or elliptic, 717 x 36 cm, chartaceous, without conspicuous dark
gland-dots on the lower side when dry; base cuneate or rarely rounded, margin faintly
toothed to entire, apex acuminate; lateral veins 812 pairs, slightly raised on both sides;
intercostal veins slightly prominent on upper side, visible on lower side; stalk 0.51 cm
long; stipules 1.52.5 cm long. Inflorescence a cyme to 1.5 cm long with alternate branches
reduced, peduncle nil or inconspicuous. Flowers 45 mm long, sessile; calyx-lobes 56,
11.5 mm long, glabrous on inner side; petals sagittate, 2.53 mm long; filaments filiform,
22.5 mm long; disc cup-shaped, divided to halfway into deltoid lobes; ovary 1-celled with
1012 ovules, style filiform, stigma capitate. Fruits ellipsoid, c. 12 mm long, 7 mm wide, 1seeded. Seeds oblong-ellipsoid, 22.5 mm long.
Distribution. Endemic to Borneo. In Sabah recorded from the Penampang and Beaufort
districts, in Sarawak in Kuching and Serian districts. Also in Kalimantan.
Ecology. Mixed dipterocarp forest to 1000 m.
4. Carallia sp. 1
Small tree to 15 m tall, 30 cm diameter. Bark smooth, dark grey-brown; inner bark brown.
Sapwood pale yellowish. Leaves broadly elliptic to obovate, 67 x 48 cm, chartaceous to
thinly coriaceous, with conspicuous dark gland-dots on the lower side when dry; base
cuneate, margin finely toothed, apex acuminate to cuspidate; lateral veins 612 pairs, raised
on both sides, forming 2several series of angular loops between midrib and margin;
intercostal veins net-like, indistinct on upper side, slightly distinct on lower side; stalk 0.51
cm long; stipules 12 cm long. Inflorescence a dense cyme to 2.5 cm long. Flowers 22.5
mm long; calyx-lobes 56, c. 1.5 mm long, glabrous on inner side; petals sagittate, to 2 mm
long; filaments filiform, 22.5 mm long; disc cup-shaped, divided to halfway into rounded
lobes; ovary 5-celled with 2 ovules per cell, style filiform, stigma capitate. Fruits globose,
to 6 mm across, 1-seeded. Seeds oblong-ellipsoid, 22.5 mm long.
Vernacular names. Sabahmeransi (Brunei Malay), merawai, tikolod, yulu tambang
(Dusun).
Distribution. Known so far from Sabah, where it is apparently common (all districts) and
often confused with C. borneensis, and apparently also present in Sarawak (4th Division at
Niah National Park, and 7th Division at Ulu Belaga). Also in Brunei (Niga NN 47, Belait).
Sabah collections include SAN 124068, SAN 85908, Nooteboom 1306 (from Tambunan),
SAN 63504 (Sandakan), and SAN 89461 (Ranau). The Sarawak collections, which have
entire leaf margins, are S. 43802 (Ulu Belaga) and S. 40118 (Niah).
Ecology. Lowland secondary forest and along rivers, and mixed dipterocarp forest to c. 600
m.
332
5. Carallia sp. 2
Small tree to 25 m tall, 30 cm diameter. Bark slightly fissured, brown; inner bark brown.
Sapwood pale pinkish brown. Leaves oblong-elliptic to slightly obovate, 27 x 24 cm,
chartaceous to thinly coriaceous, with conspicuous dark gland-dots on the lower side when
dry; base cuneate to obtuse, margin entire, apex obtuse, slightly cuspidate; lateral veins
610 pairs, raised on both sides; intercostal veins more or less parallel to lateral veins,
obscure above, slightly raised below; stalk 0.51 cm long; stipules 0.50.7 cm long.
Inflorescence a cyme to 1.4 cm long, peduncle to 1 cm long. Flowers known only as young
buds, sessile. Fruits not known.
Vernacular name. Sarawakrabong (Iban).
Distribution. Recorded only from Sarawak: S. 39097 (Niah, Ulu Sungai Sah) and S. 43023
(Limbang, Ulu Medamit).
Ecology. Lowland mixed dipterocarp forest on sandy clay.
6. Carallia sp. 3
Small tree to 10 m tall, 15 cm diameter. Leaves broadly elliptic to obovate, 59 x 45 cm,
stiffly coriaceous, without conspicuous dark gland-dots on the lower side when dry; base
cuneate, margin coarsely saw-toothed, apex acute; lateral veins 810 pairs, raised on both
sides, forming 2several series of angular loops between midrib and margin; intercostal
veins prominent on both sides; stalk 0.50.8 cm long; stipules 0.51 cm long. Inflorescence a
cyme, to 2 cm long. Flowers not known. Fruits not known.
Distribution. Recorded only from Sarawak at the Nyabau Catchment area, Bintulu, Miri (S.
24594).
Ecology. Lowland mixed dipterocarp forest on yellow red sandy humult ultisols, at about
150 m.
7. Carallia sp. 4
Shrub or small tree to 6 m tall, 5 cm diameter. Leaves broadly obovate, 811 x 47 cm,
stiffly coriaceous, with conspicuous dark gland-dots on the lower side when dry; base
cuneate, margin entire, apex obtuse and very slightly cuspidate to rounded; lateral veins
810 pairs, raised on both sides; intercostal veins prominent on both sides; stalk 0.71 cm
long; stipules c. 1 cm long. Inflorescence a cyme to 1.8 cm long, peduncle to 1 cm long.
Flowers 34 mm long, sessile; calyx-lobes 5, c. 2 mm long, glabrous on inner side; petals
sagittate, to 2.5 mm long; filaments filiform, to 2 mm long; disc cup-shaped, divided to
halfway into deltoid lobes; ovary 5-celled with 2 ovules per cell, style filiform, stigma
capitate. Fruits not known.
333
Distribution. Known only from Bukit Tawai, Telupid in Sabah (SAN 39321), on ultramafic
soil.
Ecology. Secondary forest.
8. Carallia sp. 5
Shrub or small tree 12 m tall. Leaves oblong-elliptic to obovate, to 2.7 x 2.4 cm, thinly
coriaceous, without conspicuous dark gland-dots on the lower side when dry; base cuneate,
margin faintly toothed, apex acute; lateral veins 46 pairs, slightly raised on both sides;
intercostal veins inconspicuous on upper side, slightly prominent on lower side; stalk c. 0.5
cm long; stipules 0.51 cm long. Inflorescences to 1.2 cm long, with a solitary flower.
Flowers (buds) with 4 calyx-lobes. Fruits (young) ellipsoid, c. 4 mm long.
Distribution. Recorded only from Sarawak at the summit of Batu Lawi in the Kelabit
Highlands area (Nooteboom & Chai 2259).
Ecology. Forest on sandstone at about 2000 m.
3. CERIOPS Arn.
(Greek, ceras = horn, ops = obscure; alluding to the fruit
from which the hypocotyl has yet to elongate)
Ann. Mag. Nat. Hist. 1 (1838) 363; Merrill l.c. (1921) 420; Masamune l.c. 516; Browne l.c. 302; Ding
Hou l.c. 468; Burgess l.c. 432; Ashton l.c. 357; Kochummen l.c. 315; Whitmore, Tantra & Sutisna l.c.
296.
Small or medium-sized trees with stilt-roots to 1 m high; pneumatophores prominent.

Leaves broadly elliptic to obovate, coriaceous, sometimes black-dotted on lower side;
margin entire; stipules lanceolate. Inflorescence a condensed cyme, shortly stalked or
subsessile, (2)4many-flowered. Flowers with calyx deeply 56-lobed; petals 56, each
embracing 2 stamens; ovary semi-inferior, 3-celled, ovules 2 per cell, style simple, stigma
simple or obscurely 23-lobed. Fruits ovoid, almost entirely superior. Seed germination
viviparous; hypocotyl prominently ridged or grooved.
Distribution. 2 species, tropical east Africa to Malesia, N Australia and the Pacific. Both
Ecology. At the mouths of estuaries and bays, and behind the sea beach.
Uses. The bark is a source of tannin and a dye for the batik industry, and is used in the
manufacture of an alcoholic drink. The wood is used for firewood and charcoal.
334
Key to Ceriops species

Petals fringed with many long narrow processes at the apex; anthers as long as or longer
than the filament. Fruit with erect to ascending calyx-lobes; hypocotyl 915 cm long when
fully extended at fruit-fall.............................................................................1. C. decandra
Petals with 3 short appendages at the apex; anthers less than half the length of the filament.
Fruit with reflexed calyx-lobes; hypocotyl 1535 cm long when fully extended at fruitfall.......................................................................................................................2. C. tagal
1. Ceriops decandra (Griff.) Ding Hou

(Greek, deca = ten, -andrus = male; with ten stamens in the flower)
l.c. 471; Burgess l.c. 432; Ashton l.c. 357; Kochummen l.c. 315; Whitmore, Tantra & Sutisna l.c. 296.
Basionym: Bruguiera decandra Griff. l.c. (1836) 10. Type: Griffith, s.n., Burma (K). Synonyms: C.
roxburghiana Arn. l.c. 364, Merrill l.c. (1921) 420, Masamune l.c. 516, Browne l.c. 303; C. zippeliana
Blume l.c. (1849) 143; Rhizophora glomerulata Zipp. ex Blume l.c. pro syn. R. decandra Roxb. ex
Griff. l.c. (1854) 663.
Shrub to small tree to 15 m tall, 30 cm diameter. Bark smooth to papery-flaky to

lenticellate, pale brown; inner bark red. Sapwood pale yellowish brown. Leaves obovate to
elliptic-oblong, 4.510 x 2.56 cm; base cuneate, margin slightly recurved when dry, apex
obtuse, rounded or notched; lateral veins 910 pairs, obscure, joining near margin to form a
submarginal vein; stalk 12.5 cm long; stipules 1.52.5 cm long. Flower petals white,
fringed with many long narrow processes at apex; stamens 1 mm long, anthers ovoid, more
than half the length of the filaments; style c. 1 mm long, stigma capitate. Fruits ovoidconical, c. 1.8 cm long, calyx-lobes erect to ascending; hypocotyl club-shaped, 915 cm
long, sharply ridged.
Vernacular names. Sabahtengar (Brunei Malay). Sarawakbakau lali, tengar tikus
(Malay).
Distribution. India, Indo-China, Peninsular Malaysia, Banka, Java, Borneo, Philippines,
Celebes, Moluccas and New Guinea. In Sabah recorded from Kudat (Balambangan Island
also), Sandakan and Lahad Datu; in Sarawak from Selabat to the Rejang delta. Also in
Kalimantan
Ecology. Edges of mangrove swamps and along tidal creeks.
2. Ceriops tagal (Pers.) C.B. Rob.
Fig. 3.
(a Philippine plant name)

Philip. J. Sc. 3 (1908) 306; Merrill l.c. (1921) 420; Masamune l.c. 516; Browne l.c. 302; Ding Hou l.c.
469; Burgess l.c. 432; Ashton l.c. 360; Kochummen l.c. 316; Whitmore, Tantra & Sutisna l.c. 297.
Basionym: Rhizophora tagal Pers., Mem. Soc. Linn. Paris 3 (1824) 138. Type: Perrottet, Philippines,
Zamboanga (No specimen seen by Merrill, see En. Philip. Fl. Pl. 3, 1923, 144). Synonyms:
Rhizophora timorensis DC., Prod. 3 (1828) 32; C. candolleana Arn. l.c. 364; C. pauciflora Benth. in
Hooker f., Lond. J. Bot. 2 (1843) 218; C. fosteniana Blume l.c. (1849) 143; C. boviniana Tul., Ann.
Sc. Nat. 4, 6 (1856) 112; C. lucida Miq., Sumatra (1861) 325; C. timoriensis Domin, Bibl. Bot. 8
(1928) 444.
335
Small to medium-sized tree, to 25 m, 45 cm diameter, with small stilt-roots. Bark smooth to

flaky or dippled, grey-brown; inner bark pale yellowish white to pinkish. Sapwood
yellowish white to pale brown. Leaves obovate to obovate-oblong or elliptic-oblong, 512 x
27 cm; base cuneate, margin slightly recurved when dry, apex obtuse to slightly notched;
lateral veins 912 pairs, faint, joining at margin to form a submarginal vein. Flower petals
white, with 3 club-shaped appendages at the apex; stamens 35 mm long, anthers sagittate,
less than half the length of the filaments; style c. 2 mm long, simple. Fruits ovoid, 1.52.5
cm long, calyx-lobes recurved; hypocotyl club-shaped, 1535 cm long, sharply ridged.
Vernacular names. Sabahtagal (Suluk), tangal (Suluk), tanug (Bajau) tengar (Brunei
Malay), tenug (Ilanun). Sarawaktengar samak (Malay).
Distribution. East Africa, Madagascar, Seychelles, Sri Lanka, India, Indo-China, Taiwan,
throughout Malesia to Micronesia and Australia. In Sabah, recorded from the southwest part
around Sipitang to Kota Kinabalu, and the Sandakan and Lahad Datu mangroves. In
Sarawak, recorded from the Sarawak and Rejang deltas. Also in Brunei and Kalimantan.
Ecology. In brackish water environments, on relatively well-drained sites within the reach
of tides in the inner fringes of mangroves. It is much more common than the preceeding
species.
Uses. A most durable mangrove wood. The bark is used for tanning fishing lines, nets and
sails. The trunk is used in house-building. It is also used for charcoal and known as an
excellent firewood.
4. GYNOTROCHES Blume
(Greek, gune = woman, trochos = wheel; alluding to the shape of the stigma)
Bijdr. (1825) 218; Merrill l.c. (1921) 420; Masamune l.c. 516; Ding Hou l.c. 488; Ashton l.c. 362;
Kochummen l.c. 317; Whitmore, Tantra & Sutisna l.c. 297.
Small trees; young branches hollow. Leaves with entire to faintly toothed margin; stipules
lanceolate, margins overlapping. Flowers very small, solitary on short stalks or fascicled in
dense axillary clusters, stalks jointed; calyx deeply 45-lobed; petals 45, obovate to
elliptic; stamens 810, free; ovary superior, slightly ridged, 46-celled, ovules 38 per cell,
style simple, stigma a 48-lobed discoid structure or slender with spreading lobes. Fruit a
berry, globose to oblong, few- to many-seeded.
Distribution. One species, from Upper Tenasserim, throughout Malesia, to the Western
Pacific.
Taxonomy. Although the wider concept of Ding Hou l.c. is followed here, there is some
suggestion (Juncosa & Tobe, Ann. Missouri Bot. Gard. 75 (1988) 1410) that the material
from Sarawak may represent at least two ecotypes which probably constitute distinct
species.
336
337
338
Gynotroches axillaris Blume
Fig. 4.
(Latin, axillaris = axillary; the flowers)

l.c. (1825) 219; Merrill l.c. (1921) 420; Masamune l.c. 516; Ding Hou l.c. 488; Ashton l.c. 362;
Kochummen l.c. 317; Whitmore, Tantra & Sutisna l.c. 297. Type: Blume, s.n., Java (L). Synonyms:
Dryptopetalum coriaceum Arn. l.c. 373; G. dryptopetalum Blume l.c. (1849) 127; G. micrantha Blume
l.c. (1849) 127, Masamune l.c. 517; G. reticulata A. Gray in Wilkes, Bot. U.S. Expl. Exped. 1 (1854)
607; G. parvifolia Merr., Publ. Gov. Lab. Philip. 35 (1905) 46; G. puberula Merr., Philip. J. Sc., Bot.
10 (1915) 333; G. lanceolata Merr., Philip. J. Sc., Bot. 11 (1916) 21.
Small or medium-sized tree, to 35 m tall, 70 cm diameter. Bark smooth, often hoop-marked,

grey to blackish; inner bark yellowish to reddish brown, fibrous, finely and regularly
invaginating into the wood. Sapwood yellowish brown to pale reddish brown. Leaves ovate
to elliptic-oblong or lanceolate, 515 x 37.5 cm, usually coriaceous, glabrous or (rarely)
pubescent on the midrib and lateral veins below; base cuneate, apex acute to acuminate;
midrib flat on upper side, raised on lower side; lateral veins 812 pairs, flat on upper side,
raised on lower side, forming distinct loops just behind the margin; intercostal veins netlike, flat on upper side, distinctly prominent on lower side; stalks 0.51.5 cm long; stipules
to 1.5 cm long. Flowers greenish white, 116 in each leaf axil, bisexual but sometimes male
by abortion; calyx-lobes with tufts of hairs at apex; petals clawed, divided into appendages
towards the apex; stamens 12 mm long; disc cup-shaped or nearly flat, 810-lobed; ovary
ovoid; style to 2 mm long. Fruits usually globose, c. 3 mm across, green ripening red to
shiny black, persistent calyx-lobes erect or reflexed.
Vernacular names. Sabahbulu bulu (Malay), kandis batu (Malay), kupi kupi (Brunei
Malay), pahau pahau (Murut). Sarawakkelalud (Melanau), sawar bubu (Iban). Brunei
kerakas payau (Kedayan), kerakas payoh (Dusun), sawar bubu (Iban).
Distribution. Burma, Thailand, Malesia except the Lesser Sunda Islands, to Melanesia and
Micronesia. In Sabah and Sarawak, in all districts. Also recorded for Brunei and
Kalimantan.
Ecology. Lowlands to about 2200 m, in marshy places especially along rivers, and also
well-drained sandy sites. Occasionally also in secondary forests.
5. KANDELIA Wight & Arn.

(from a Malabar (Indian) plant name)
Prod. (1834) 310; Merrill l.c. (1921) 421; Masamune l.c. 517; Ding Hou l.c. 473; Ashton l.c. 363;
Kochummen l.c. 319.
Small trees. Leaves with linear stipules. Inflorescence a peduncled cyme bearing 49
flowers. Flowers with calyx deeply 5(6)-lobed; petals 5(6), bilobed, with a long bristle
339
340
between lobes, each lobe laciniate to fimbriate; stamens many, free, of unequal length,
exserted, anthers many-celled; ovary 1-celled, ovules 6, apically attached, style simple.
Fruit ovoid. Seed solitary; germination viviparous; hypocotyl cylindric to spindle-shaped.
Distribution. One species; India, Indo-China to S China and western Malesia (NE Sumatra,
Peninsular Malaysia, N and W Borneo).
Kandelia candel (L.) Druce
Fig. 5.
(a Malabar plant name)

Rep. Bot. Exch. Club. Br. Isl. 3 (1914) 420; Merrill l.c. (1921) 421; Masamune l.c 517; Ding Hou l.c.
473; Ashton l.c. 363; Kochummen l.c. 319. Basionym: Rhizophora candel L., Sp. Pl. (1753) 443.
Shrub or small tree, to 10 m tall; trunk with swollen conical base but no buttresses. Bark
flaky, lenticellate, greyish to reddish brown; inner bark thin, pink. Sapwood pale yellowish
brown. Leaves narrowly oblong-elliptic to obovate-oblong, 613 x 26 cm, coriaceous;
base cuneate, margin frequently recurved, apex acute-blunt to rounded; lateral veins 67
pairs, inconspicuous on both sides; stalks 11.5 cm long; stipules 0.51.5 cm long. Flowers
c. 2 cm long, white; calyx 56-lobed, reflexed after anthesis; petals c. 14 mm long; stamens
with small narrow anthers on slender filaments; style filiform, stigmas 3. Fruits ovoid,
12.5 cm long, 1-seeded; hypocotyl to 40 cm long, terete, smooth.
Vernacular names. Sabahberus berus (Malay), beus (Bajau, Malay), linggayong,
linggayong laut (Brunei Malay). Sarawakbakau aleh aleh (Malay).
Distribution. India, Burma, Thailand, Indo-China, China, Ryukyus, S Japan, NE Sumatra,
Peninsular Malaysia, W and N Borneo. In Sabah, recorded so far from the Papar and
Sandakan areas; in Sarawak from the Kuching, Sajingkat and Kelepu mangroves. Also in
Brunei and W Kalimantan
Ecology. Scattered along the banks of brackish water (tidal) rivers and channels in
mangrove areas.
6. PELLACALYX Korth.
(Greek, pella = hide, kalux = calyx; the hair-covered calyx)
merbuloh (Iban, Malay; Sarawak), sawar (sambar) bubu (Iban; Sarawak)
Tijd. Nat. Gesh. Phys. 3 (1836) 20; Merrill l.c. (1921) 422; Masamune l.c. 517; Ding Hou l.c. 490;
Ashton l.c. 363; Kochummen l.c. 320; Whitmore, Tantra & Sutisna l.c. 297. Synonyms: Plaesiantha
Hook. f. in Benth. & Hooker f., Gen. Pl. 1 (1865) 861; Craterianthus Valeton ex Heyne, Nutt. Pl.
Indon. 4 (1917) 196.
Small to medium-sized trees; young branches hollow. Bark invaginating finely and
regularly into the wood. Leaves oblong to ovate; sparsely to densely pubescent with stellate
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342
or simple hairs (rarely a mixture of the two), sometimes glabrescent to glabrous; margin
entire or obscurely to distinctly serrulate; stipules with margins not overlapping. Flowers
fascicled, 28 in a leaf axil, or on short dense inflorescences; calyx tubular, lobes
(3)45(6), tube hairy inside at the lower part; petals 45, inserted on the margin of the
calyx-tube, densely short-hairy outside; stamens twice as many as petals, inserted on the
margin of the calyx-tube, filaments connate at the base or free, in 1 or (rarely) 2 series,
unequal, anthers suborbicular; ovary inferior, 910(12)-celled, ovules 825 per cell, style
columnar and usually hairy, stigma discoid or capitate, obscurely 810-lobed. Fruit a berry,
sub-globose. Seeds few to many.
Distribution. 7 or 8 species; Burma and S China to Sumatra, Borneo, Philippines, and
Celebes. In Sabah and Sarawak, 4 species.
Ecology. In Sabah and Sarawak in mixed dipterocarp forest to 1300 m.
Key to Pellacalyx species

1.
Leaf lateral veins 57 pairs. Calyx 4-merous, the outside covered in very minute hairs
not easily visible to the unaided eye (appearing glabrous).............................3. P. lobbii
Leaf lateral veins 912 pairs. Calyx 5-merous, the outside conspicuously hairy............2
2.
Lower leaf surface with a mixture of simple hairs and stellate hairs, both types similarly
abundant. Inside of calyx-tube with scanty short straight hairs at the base. Stamens in
two series.....................................................................4. P. symphiodiscus
Lower leaf surface with predominantly stellate scales (mixed with a few simple hairs)
or predominantly simple hairs (mixed with a few stellate scales). Inside of calyx-tube
with either a band of woolly hairs and scanty short straight hairs below the band, or a
zone of villous hairs at the base. Stamens in one series................................................3
3.
Lower leaf surface with predominantly stellate scales mixed with a few simple hairs.
Calyx-lobes reflexed at floral maturity; inside of calyx-tube with a band of woolly hairs
and scanty short straight hairs below the band. Petal tips fringed with linear segments
attenuating into fine processes..................................................................1. P. axillaris
Lower leaf surface with predominantly simple hairs mixed with a few stellate scales.
Calyx-lobes ascending at floral maturity; inside of calyx-tube with a zone of villous
hairs near the base. Petal tips with very fine hair-like processes...............2. P. cristatus
1. Pellacalyx axillaris Korth.
Fig. 6.
(Latin, axillaris = axillary; the inflorescences)

l.c. 20, t. 2; Ding Hou l.c. 493; Ashton l.c. 364; Kochummen l.c. 320; Whitmore, Tantra & Sutisna l.c.
297. Type: Korthals, s.n., Sumatra (L).
Small to medium-sized tree, to 25 m tall, 60 cm diameter; buttresses if present to 60 cm

high. Bark pale reddish brown, finely cracked to rough-fissured; inner bark pinkish yellow.
Sapwood yellow. Leaves oblong to obovate-oblong, 817 x 3.57 cm, lower side densely
or sparsely covered with stellate scales mixed with a few simple hairs; base broadly cuneate
to rounded, margin entire or toothed (with a tuft of short hairs on each tooth), apex acute to
343
acuminate; lateral veins 912 pairs, flat on upper side, raised on lower side; stalk 58 mm
long; stipule 11.5 cm long. Flower calyx conspicuously hairy outside, the tube 56 mm
long, inside with a few short hairs at the base and a band of woolly hairs above these, lobes
5, 22.5 mm long, reflexed at anthesis; petals 1.52.5 mm long, the apices fringed with
linear segments attenuating into fine processes; stamens in one series; style 1.52 mm long.
Fruits ovoid or subglobose, c. 10 mm across.
Vernacular name. Sarawakdanguh (Bidayuh).
Distribution. Sumatra, Peninsular Malaysia, Borneo and Mindanao (Philippines). In Sabah
recorded from Kudat, Beaufort and Telupid. In Sarawak recorded from the Padawan area
(1st Div.) and Sg. Beria in the Kapit District (7th Div.). Also in Brunei and Kalimantan.
Ecology. Common where it occurs in lowland primary mixed dipterocarp forest and secondary
forest, usually in valleys and moist sites.
2. Pellacalyx cristatus Hemsl.

(Latin, cristatus = crested; the persistent calyx crowning the fruit)
in Hooker f., Ic. Pl. 16 (1886) sub t. 1546; Masamune l.c. 517; Ding Hou l.c. 493; Ashton l.c. 364;
Whitmore, Tantra & Sutisna l.c. 297. Type: Beccari PB 1258, Sarawak (G, K).
Small tree to 10 m tall, 10 cm diameter. Bark yellowish brown, smooth; inner bark pale
brown. Sapwood pale yellow. Leaves narrowly oblong to ovate-oblong, 1017 x 35 cm,
lower side densely or sparsely covered with simple hairs mixed with a few stellate scales, or
glabrescent; base obtuse to rounded, margin toothed (with a tuft of short hairs on each
tooth), apex acuminate; lateral veins 912 pairs, flat on upper side, raised on lower side;
stalk 710 mm long; stipule 11.5 cm long. Flower calyx conspicuously hairy outside, the
tube 910 mm long, inside with a few short hairs at the base and a zone of villous hairs
above these, lobes 5, 1.52 mm long, ascending at anthesis; petals 2.53.5 mm long, the
apices fringed with fine hair-like processes; stamens in one series; style 2.53 mm long.
Fruits subglobose, to 10 mm across.
Distribution. Endemic to Borneo. In Sabah, recorded from Nabawan northwards to Mt.
Kinabalu; in Sarawak from the Kuching area, Belaga and Mt. Api, Mulu National Park.
Also in Kalimantan.
Ecology. Hills and mountain ridges to c. 900 m.
3. Pellacalyx lobbii (Hook. f.) Schimp.

(Thomas Lobb, 18201894, plant collector)
in Engl. & Prantl, Pfl. Fam. 3, 7 (1893) 54; Masamune l.c. 517; Ding Hou l.c. 491; Ashton l.c. 365;
Whitmore, Tantra & Sutisna l.c. 297. Basionym: Plaesiantha lobbii Hook. f. in Benth. & Hooker f.,
Gen. Pl. 1 (1865) 145. Type: Lobb, s.n., Brunei (K).
Medium-sized tree, to 45 m tall, 65 cm diameter; stilt-roots at times present. Bark medium

brown, cracking-fissured; inner bark pale brown. Sapwood yellow. Leaves obovate to
obovate-oblong, 717 x 26 cm, lower side sparsely covered with stellate hairs; base
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cuneate, margin entire or faintly toothed (glabrous on each tooth), apex acute to abruptly
acuminate; lateral veins 57 pairs, flat on upper side, raised on lower side; stalk 58 mm
long; stipule 0.51 cm long. Flower calyx very minutely hairy outside (appearing glabrous
to the unaided eye), the tube 67 mm long, inside with a few short hairs at the base, lobes 4,
2.53 mm long, ascending at anthesis; petals 1.52 mm long, the apices finely serrulate;
stamens in one series; style 11.5 mm long. Fruits subglobose, to 8 mm across.
Vernacular names. Sarawakpaserujan gunugo, rasu (Iban).
Distribution. Sumatra and Borneo. In Sabah and Sarawak, in most districts. Also in Brunei
and Kalimantan.
Ecology. Primary and secondary forests.
4. Pellacalyx symphiodiscus Stapf

(Greek, sumphuein = fused, discus = disc; of fused parts forming the floral disc)
Kew Bull. (1898) 224; Merrill l.c. (1921) 422; Masamune l.c. 517; Ding Hou l.c. 492; Ashton l.c. 365;
Whitmore, Tantra & Sutisna l.c. 297. Type: Haviland 2206, Sarawak (K).
Small to medium-sized tree, to 25 m tall, 40 cm diameter. Bark pale brown, smooth; inner
bark pale brown. Sapwood yellow. Leaves ovate-oblong to obovate-oblong, 919 x 3.57
cm, lower side densely or sparsely covered with a mixture of stellate hairs and simple hairs
(both types similarly abundant); base acute or rounded, margin toothed (with a tuft of short
hairs on each tooth), apex acute to shortly acuminate; lateral veins 912 pairs, flat on upper
side, raised on lower side; stalk 610 mm long; stipule 0.81.2 cm long. Flower calyx
conspicuously hairy outside, the tube 45 mm long, inside with a few short hairs at the
base, lobes 5, 1.52 mm long, reflexed at anthesis; petals 1.52.5 mm long, the apices
fringed with fine hair-like processes; stamens in two series; style 1.52 mm long. Fruits
subglobose, to 9 mm across.
Distribution. Endemic to Borneo. In Sabah, known only from the Penampang and Lamag
districts; in Sarawak from the Kuching, Selang and Pengkalan Ampat areas. Also in Brunei
and Kalimantan.
Ecology. Mixed dipterocarp forest to c. 650 m.
7. RHIZOPHORA L.
(Greek, rhiza = root, pherein = bearing; the stilt-roots)
bakau (Malay)
Gen. Pl. ed. 5 (1754) 202; Merrill l.c. (1921) 420; Masamune l.c. 517; Browne l.c. 296; Ding Hou l.c.
448; Burgess l.c. 433; Tomlinson & Womersley, Contrib. Herb. Austr. 19 (1976) 1; Ashton l.c. 366;
Kochummen l.c. 321; Whitmore, Tantra & Sutisna l.c. 298. Synonyms: Mangium Rumph. ex Scop.,
Introd. Hist. Nat. (1777) 218.
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346
Small to medium-sized trees with conspicuous branching stilt-roots but no pneumatophores. Leaves succulent, entire, apex acute, lower side usually black-dotted; stipules
lanceolate, conspicuous. Inflorescence a cyme, 23-branched. Flowers bisexual; calyx
leathery, deeply 4-lobed, lobes broadly ovate, ascending but reflexed in the fruit; petals 4,
falling off early; stamens 812, filaments very short or hardly distinct; ovary semi-inferior,
2-celled, ovules 2 per cell, style obscure or to 6 cm long, stigma simple or somewhat 2lobed. Fruits ovoid, usually 1-seeded; germination viviparous.
Distribution. 7 species, mainly along tropical coasts and throughout Malesia. In Sabah, 3
species and in Sarawak, 2.
Ecology. Mangrove swamps and tidal parts of rivers; the most widespread mangrove genus.
Uses. The poles are used in fishing platforms, traps and house frames. The wood is durable
as piling timber and is also used in concrete foundations for buildings. Excellent as firewood
and for making charcoal. The bark is also used for tanning.
Key to Rhizophora species

1.
Inflorescences 2-flowered, shorter than the leaf-stalks, mostly in the axils of leaf-scars
on the branches just below the current cluster of leaves. Petals glabrous save for the
occasional presence of a few scattered short-hairs....................................1. R. apiculata
Inflorescences 216-flowered, longer than the leaf-stalks, in leaf axils within the
current cluster of leaves. Petals long-hairy on the margins and inner side....................2
2.
Leaves typically 1118 x 510.5 cm. Style obscure or very short, not exceeding 1.5
mm......................................................................................................2. R. mucronata
Leaves typically 612 x 36 cm. Style filiform, 46 mm long.....................3. R. stylosa
1. Rhizophora apiculata Blume
Fig. 7.
(Latin, apiculatus = with a short abrupt point; the leaf apex)

l.c. (1827) 91; Ding Hou l.c. 452; Burgess l.c. 433; Ashton l.c. 366; Kochummen l.c. 322; Whitmore,
Tantra & Sutisna l.c. 298. Type: Pee-Kandel Rheede, Hort. Mal. 6 (1686) 61, t. 34. Synonyms: R.
candelaria DC., Prod. 3 (1828) 32, Merrill l.c. (1921) 420, Masamune l.c. 517; R. conjugata Arn. (non
L.) l.c. 363, Browne l.c. 298.
Medium-sized tree to 30 m tall, 75 cm diameter. Bark pinkish grey to grey, smooth to

shallowly fissured. Sapwood pale brown. Leaves narrowly elliptic to sublanceolate, 716 x
37 cm, coriaceous; base cuneate, apex acute to pointed, with a short mucro; stalk 1.53 cm
long; stipules 48 cm long. Inflorescences 2-flowered, shorter than the petioles, peduncle
0.51.5 cm long, mostly in the axils of leaf-scars on the branches just behind the current
cluster of leaves. Flowers sessile, calyx-lobes 1014 x 68 mm; petals 811 x 1.5 2 mm,
glabrous except for the occasional presence of a few scattered short-hairs; stamens 1012,
sessile; style 0.51 mm long. Fruits obpyriform (inverted pear-shaped), 22.5 cm long,
hypocotyl to 33 cm long.
347
Vernacular names. Sabahbangkita (Malay). Sarawakbakau minyak (Malay).

Distribution. Tropical SE Asia, Sri Lanka, throughout Malesia to Micronesia and New
Britain, the Solomons and New Hebrides. In Sabah, the largest stands occur between the
Bay of Brunei and Klias in the SW, but generally common in mangroves. In Sarawak, with
good stands at the Sarawak River Delta, the Lower Rejang and generally elsewhere. Also in
Ecology. Dominant species of the mangroves, forming pure stands on soft recent muddy
substrates, immediately behind the river banks or up to them, in the lower parts of major
river deltas.
2. Rhizophora mucronata Lam.

(Latin, mucronatus = with an abrupt short point; the leaf apex)
Encycl. 6 (1804) 189; Merrill l.c. (1921) 421; Masamune l.c. 517; Browne l.c. 298; Ding Hou l.c. 453;
Burgess l.c. 433; Ashton l.c. 367; Kochummen l.c. 322; Whitmore, Tantra & Sutisna l.c. 298. Type:
Commerson, s.n., isle de France (Mauritius) (P). Synonyms: R. mangle Roxb. (non L.), Hort. Beng.
(1814) 36; R. macrorrhiza Griff. l.c. (1836) 8; R. lingissima Blanco, Fl. Filip. (1837) 398.
Medium-sized tree to 30 m tall, 210 cm diameter. Bark grey to black, roughly grid-cracked.
Sapwood pale brown. Leaves broadly elliptic to oblong, 1118 x 510.5 cm, coriaceous;
base cuneate, apex acute to slightly obtuse, with a short mucro; stalk 2.55.5 cm long;
stipules 5.58.5 cm long. Inflorescences 25(12)-flowered, longer than the petioles,
peduncle 2.55.5 cm long, in leaf axils within the current cluster of leaves. Flowers with
stalks 48 mm long, calyx-lobes 1315 x 57 mm; petals 810 x 23 mm, long-hairy on
the margins and inner side; stamens 8, subsessile; style 0.51.5 mm long. Fruits narrowly
ovoid, 57 cm long, hypocotyl to 62 cm long.
Vernacular name. Sabah and Sarawakbakau kurap (Malay).
Distribution. E Africa through S and SE Asia to Queensland, Melanesia and Micronesia,
also in Tonga. In Sabah and Sarawak, common in mangrove areas. Also in Brunei and
Kalimantan.
Ecology. Usually in small stands or scattered groups along the lower parts of tidal rivers,
both on clay-rich and sandy deposits.
3. Rhizophora stylosa Griff.

(Latin, stylosus = with a conspicuous style; the flower)
l.c. (1854) 665; Ding Hou l.c. 456; Kochummen l.c. 323. Type: Griffith, s.n., Malacca, Pulo Bissar
(K). Synonym: R. mucronata var. stylosa Schimp., Bot. Mitt. Trop. 3 (1891) 92.
Small tree to 15 m tall, 25 cm diameter. Bark dark brown to black, fissured. Sapwood pale
yellowish brown. Leaves elliptic to sublanceolate, 6.512 x 36 cm, coriaceous; base
cuneate, apex acute to pointed, with a short mucro; stalk 1.52.5 cm long; stipules 46 cm
348
long. Inflorescences 48(16)-flowered, longer than the petioles, peduncle 2.55 cm long,
in leaf-axils within the current cluster of leaves. Flowers with stalks 0.51 mm long, calyxlobes 912 x 35 mm; petals 78 x 2.53.5 mm, long-hairy on the margins and inner side;
stamens 8, filaments short but distinct; style 46 mm long. Fruits ovoid to obpyriform,
2.54 cm long, hypocotyl to 54 cm long.
Distribution. Taiwan, Peninsular Malaysia, Java, Borneo, Philippines, Celebes, Moluccas,
New Guinea to Melanesia, New Britain and Queensland. In Sabah, recorded only from the
Lahad Datu area (SAN 26145) and Pulau Balambangan in the north (SAN 85589). Not
recorded for Sarawak, Brunei and Kalimantan.
Ecology. Exclusively along sandy shores and coral terraces facing open sea.
349
RUTACEAE
David T. Jones
South Florida Natural Resources Center,
Everglades National Park, Homestead, Florida, U.S.A.
Hooker f., Fl. Brit. Ind. 1 (1875) 484; Kurz, For. Fl. Brit. Burma 1 (1877) 178; Merrill, EB (1921) 313,
PEB (1929) 113; Ridley, FMP 1 (1922) 340; Craib, Fl. Siam Enum. 1 (1931) 215; Masamune, EPB
(1942) 357; Browne, FTSB (1955) 315; Backer & Bakhuizen f., FJ 2 (1965) 94; Swingle, Citrus
Industry 1 (1967) 190; Stone, TFM 1 (1972) 367; Anderson, CLTS (1980) 307; Perry, MPESA (1980)
361; Corner, WSTM 2 (1988) 656; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 304; Ng, MFFSS 2
(1992) 493.
Aromatic trees, shrubs, or woody climbers, or seldom herbs; twigs and branches sometimes
armed with spines or thorns; tissues usually dotted with lysigenous oil-glands containing
volatile essential oils. Leaves pinnate or trifoliolate or unifoliolate, less commonly simple;
opposite, alternate or spirally arranged, rarely whorled, often conspicuously pellucid-dotted;
petioles sometimes winged, usually articulated with the blade, without stipules.
Inflorescences terminal or axillary panicles, cymes or racemes, or flowers seldom solitary,
rarely epiphyllous. Flowers bisexual or (in dioecious plants) unisexual, regular or rarely
somewhat irregular; sepals 45, rarely 23, usually imbricate, free or fused basally to form
a cup-like calyx; petals 45, rarely 23 or 6, alternate with the sepals, imbricate or valvate,
free or fused basally, rarely united to form a tube; stamens 45 or 810 or more (to 60),
equal or unequal, sometimes intermixed with staminodes, filaments free or fused basally,
rarely united to form a tube; anthers 2-celled, often gland-tipped; disc surrounding base of
ovary, intrastaminal, usually annular, nectariferous, sometimes modified into a gynophore,
rarely lacking; ovary superior, carpels 45, rarely 13 or 6 or more, completely to partially
fused or free except for the united styles, ovules 12 per locule, rarely more, placentation
axile, stigma small or broad, sometimes lobed. Fruit a capsule, follicle, samara, drupe,
berry or hesperidium (Citrus), dry or fleshy; hesperidia composed of juice-filled pulpvesicles, with or without oil-droplets; pericarp leathery to woody or fleshy, usually
glandular. Seeds usually 1 per carpel, ovoid to ellipsoid or oblong, with or without endosperm, sometimes winged, sometimes hairy; testa firm or membranous; monoembryonic or
polyembryonic; embryo straight or curved; cotyledons white or green, flat or folded.
Distribution. A cosmopolitan family comprising approximately 160 genera and 1650
species, distributed largely in tropical and subtropical parts of the world, with good
representation in arid areas of the warm temperate zone (Australia and S Africa). In Sabah
and Sarawak, 23 genera with about 75 species, of which 17 genera and 43 species are native
trees and shrubs. The remaining taxa are scandent shrubs (15 species), or are cultivated (c.
17 species). The genera Fortunella (23 species), Limonia (1 species), and Triphasia (1
species) are represented only by introduced species found under cultivation and are
excluded from this treatment.
351
Ecology. Native species are found in a wide range of natural habitats occurring in the
lowlands, hills, and mountains, along coastal areas, and on offshore islands. These habitats
include primary and secondary forests (flat or undulating lands, ridges, hillsides), forest
margins, open places and disturbed sites, mangrove forest (Merope), heath forest, swamp
(freshwater, peat) forest, streamsides and riverbanks, and beach forest (Severinia). Some
species show affinities for specific soil types such as ultramafic (Lunasia) and limestone
(some Clausena, Glycosmis, Murraya and Micromelum). The majority of species are found
below 1300 m elevation, and a few ascend to higher altitudes (Citrus and Tetractomia to
1800 m, and Melicope to 2400 m). Thirteen tree and shrub taxa are endemic to Sabah and/or
Sarawak, four of which are found in each of the genera Glycosmis and Melicope, followed
by Maclurodendron (2 species), Clausena, Monanthocitrus, and Pleiospermium. Abundance
varies among species, ranging from widespread and common to infrequent and rare.
Merrillia is very rare in Sabah, and Burkillanthus is endangered, if not already extinct, in
Sarawak.
The typically small size and drab colour (white or cream to greenish or yellowish) of the
flowers of the Rutoideae (Melicope, Tetractomia, Zanthoxylum) and Toddalioideae
(Acronychia, Maclurodendron), and the larger, often fragrant flowers of the Aurantioideae
(Citrus, Murraya, Pleiospermium, etc.), suggests insect pollination. The watery nectar
secreted through stomata in the outer portions of the floral disc serves as both an attractant
and food source. Fruit and seed dispersal is by birds (Melicope, Zanthoxylum), wind
(Tetractomia), water (Merope), and fruit- and seed-eating animals (Citrus, Clausena,
Glycosmis, Micromelum, Severinia, and most other Aurantioideae).
Uses. The family includes the wild and cultivated citrus fruits of the subfamily
Aurantioideae (chiefly from Citrus and Fortunella) which serve as a source of food and
flavourings, essential oils, and medicines. A number of other genera provide lesser known
edible fruits (Clausena, Limonia, and Triphasia), flavours (Murraya), and traditional
medicines and poisons (Glycosmis, Merope, Micromelum, Murraya, Paramignya, Severinia).
Although few of the species reach timber size, the wood of some is valued for its special
qualities such as grain, colour, or hardness (Citrus, Merrillia, Micromelum, and Murraya).
A number of species are prized as ornamentals or hedge-plants due to their attractive foliage
and fragrant flowers (Citrus, Fortunella, Glycosmis, Luvunga, Murraya, and Triphasia).
Certain species of Pleiospermium and Severinia have been suggested as a source of
rootstock material for Citrus breeding and improvement programmes.
More than half the species of the non-aurantioid genera (subfamilies Rutoideae and
Toddalioideae) attain timber size, yet only Acronychia (1 species), Maclurodendron (1
species) and Melicope (1 species) have reportedly been used for construction purposes. All
of the genera (except Maclurodendron), including Toddalia, are said to have minor
medicinal importance. Lesser-known uses are as a source of resin (Melicope), food and
flavouring (Acronychia and Melicope), and fish poison (Acronychia).
Taxonomy. The Rutaceae is generally accepted as closely allied to the Meliaceae,
Simaroubaceae, and in some characters to other families included in the order Sapindales,
e.g., the Anacardiaceae, Burseraceae, and Sapindaceae. From the Meliaceae and Simaroubaceae
(its closest allies), the family differs by its glandular-punctate leaves and the presence of
secretory cavities containing aromatic esthereal oils in the parenchymatous and pericarp
tissues. In the most detailed treatment of the family, Engler (in Engler & Prantl, Pfl. Fam. 19
352
RUTACEAE (JONES)
a (1931) 187359) recognised 7 subfamilies. Airy-Shaw (in Willis, Dict. Fl. Pl. & Ferns 8th
ed. (1973) 1014), however, raised the subfamily Flindersioideae and Rhabdodendroideae to
family status, thus reducing the number of subfamilies to 5. Of these 5 or 7 subfamilies,
only the Aurantioideae, Rutoideae and Toddalioideae are represented by native species in
Sabah and Sarawak. More recently, Da Silva et al. (Pl. Syst. Evol. 161 (1988) 97134), on
the basis of phytochemical, morphological and geographical evidence, proposed the
inclusion of the subfamily Toddalioideae in the Rutoideae, and the reorganisation of tribes
and genera in the subfamily Aurantioideae.
Key to subfamilies
1.
Carpels usually 35, usually joined only by the styles, separated in fruit. Fruits follicular,
woody, dehiscing along the upper suture...............................................subfam. Rutoideae
(Genera: Lunasia, Melicope, Tetractomia, Zanthoxylum)
Carpels 220, joined completely or nearly so. Fruit a drupe or berry, dry or fleshy,
indehiscent..............................................................................................................................2
2.
Carpels 25, rarely 1, rarely completely joined. Fruit a single, several-loculed drupe, or
24 druplets joined at the base. Seeds with endosperm, cotyledons obscure..................
.................................................................................................subfam. Toddalioideae
(Genera: Acronychia, Maclurodendron, Toddalia)
Carpels usually 420, completely joined. Fruit a dry or fleshy berry or hesperidium,
with or without pulp-vesicles. Seeds lacking endosperm, cotyledons well-developed
......................................................................................................subfam. Aurantioideae
(Genera: Burkillanthus, Citrus, Clausena, Fortunella,
Glycosmis, Limonia, Luvunga, Merope, Merrillia,
Micromelum, Monanthocitrus, Murraya, Paramignya,
Pleiospermium, Severinia, Triphasia).
Key to native tree and shrub genera

1.
Twigs, branchlets, and older branches unarmed. Leaves opposite.................................2

Twigs and branchlets and/or older branches armed with spines or prickles, or unarmed.
Leaves alternate or spirally arranged............................................................5
2.
Leaves trifoliolate and/or unifoliolate, rarely bifoliolate; petioles 130 cm long, not
swollen apically..........................................................................................8. Melicope
Leaves strictly unifoliolate; petioles not longer than 5 cm, generally swollen
apically........................................................................................................................3
3.
Leaves generally leathery. Fruit a group of 14 dehiscent follicles.......16. Tetractomia

Leaves thinly leathery. Fruit a 4-loculate, indehiscent drupe........................................4
4.
Younger branchlets glabrous to finely hairy. Leaflets usually elliptic; lateral veins and
reticulations conspicuous below. Flowers bisexual...................................1. Acronychia
353
Younger branchlets brown- to rusty-hairy. Leaflets usually obovate to oblanceolate;

lateral veins and reticulations inconspicuous below. Flowers unisexual..........................
......................................................................................................7. Maclurodendron
5.
Twigs and branchlets unarmed (or with spine-like paraphylls in some Severinia)........6
Twigs and branchlets and/or older branches armed with spines or prickles (or unarmed
or nearly so in some Pleiospermium)..........................................................................12
6.
Leaves simple; blades often rigid when older, not articulated at the base......................7
Leaves pinnate and/or unifoliolate; leaflet blades not as above, articulated at the
base..........8
7.
Leaf-margins sinuate or coarsely dentate. Petioles swollen apically. Fruit a group of

ribbed follicles..............................................................................................6. Lunasia
Leaf-margins subentire to obscurely crenulate. Petioles not as above. Fruit a fleshy
berry.........................................................................................................15. Severinia
8.
Young branch-tips densely rusty-hairy. Leaves pinnate and/or unifoliolate, often leathery
..................................................................................................................5. Glycosmis
Young branch-tips glabrous or hairy (if hairy, not as above). Leaves pinnate, thinly
leathery........................................................................................................................9
9.
Leaf-rachis narrowly winged. Leaflets alternate to subopposite. Fruit a large berry, 7 cm

in diameter or more; pericarp thick, leathery.......................................10. Merrillia
Leaf-rachis wingless. Leaflets alternate. Fruit a small berry, 12 cm in diameter or less;
pericarp thin......................................................................................................10
10. Leaflets generally 37 (or more in some of the cultivated species); bases not obliquely
asymmetric (or asymmetric in some of the cultivated species). Inflorescence an axillary
panicle (or terminal and/or corymbose in some of the cultivated species)...13. Murraya
Leaflets generally 515 (or more in some Clausena); base obliquely asymmetric.
Inflorescence a terminal panicle or corymb................................................................11
11. Leaflets 915. Inflorescence a flat-topped corymb. Petals valvate in bud....11.
Micromelum
Leaflets 531. Inflorescence a panicle. Petals imbricate in bud....................4. Clausena
12. Trees or scandent to erect shrubs. Twigs and branchlets and/or older branches armed
with numerous thick spines or small (sometimes hollow or recurved) prickles. Leaves
pinnate. Fruit of 14 follicles.............................................................17. Zanthoxylum
Trees or erect shrubs. Twigs and branchlets and/or older branches armed with straight,
terete, axillary spines, or unarmed or nearly so. Leaves 23-foliolate and/or unifoliolate,
or simple. Fruit a fleshy berry or hesperidium.........................................13
13. Leaves 23-foliolate (or unifoliolate in juvenile growth), rather stiff when older
...........................................................................................................2. Burkillanthus
Leaves strictly unifoliolate or simple..........................................................................14
14. Shrubs or small trees. Petioles wingless. Fruits lacking pulp-vesicles.........................15
Shrubs or small to large trees. Petioles narrowly to broadly winged, or rarely wingless.
Fruits containing pulp-vesicles...................................................................................16
354
RUTACEAE (JONES)
15. Leaves oblong-ovate, thick, fleshy; lateral veins inconspicuous. Petiole articulated with
the blade. Mangrove swamp plants................................................................9. Merope
Leaves oblong-oblanceolate, thin, not fleshy; lateral veins conspicuous. Petiole not
articulated with the blade. Inland lowland forest plants..................12. Monanthocitrus
16. Spines solitary or paired, generally only on older branches, or branches unarmed or
nearly so. Fruits 1.52.5 cm in diameter..........................................14. Pleiospermium
Spines solitary, predominantly on younger branches and twigs, older branches often
unarmed. Fruits mostly 4 cm in diameter or larger..........................................3. Citrus
Key to native genera

1.
Carpels 34, seldom 2, often joined by the styles but separated in fruit. Fruits follicular,
leathery or woody, dehiscing along the upper suture; endocarp sometimes detaching...2
Carpels 218, incompletely or completely united. Fruit an indehiscent, dry or fleshy
drupe or berry..............................................................................................................5
2.
Flowers 45-merous, unisexual or bisexual. Branches armed with spines or prickles, or

unarmed......................................................................................................................3
Flowers 34-merous, unisexual (or bisexual in some Melicope). Branches unarmed....4
3.
Ovary 4-carpellate. Follicles boat-shaped; exocarp leathery, not glandular. Seeds winged,
testa parchment-like. Flowers bisexual. Branches unarmed..................16. Tetractomia
Ovary 24-carpellate. Follicles subglobose; exocarp conspicuously glandular. Seeds unwingless, testa black, glossy. Flowers unisexual. Branches armed with spines or
prickles..............................................................................................17. Zanthoxylum
4.
Flowers 3-merous, unisexual. Follicles flattened laterally, ribbed, with a short beak.
Leaves simple, alternate................................................................................6. Lunasia
Flowers 4-merous, unisexual or bisexual. Follicles subglobose, not ribbed, without
beak. Leaves trifoliolate and/or unifoliolate, opposite..................................8. Melicope
5.
Carpels rarely completely united; ovary usually shallowly lobed. Fruit a drupe............6
Carpels always completely united; ovary not lobed. Fruit a berry or hesperidium.........8
6.
Scandent shrubs, armed with prickles. Leaves trifoliolate, alternate or spiral. Flowers
unisexual.......................................................................................................................
Toddalia Juss.
Gen. Pl. 3 (1789) 371; Hooker f. l.c. (1875) 497; Kurz l.c. (1877) 183; Craib l.c. 220; Backer
& Bakhuizen f. l.c 101; Perry l.c. 368.
1 species; Africa, Madagascar, India, Sri Lanka, S China, Indonesia, Borneo

(Sabah and Sarawak), and the Philippines.
Climbers with prickly branchlets. Leaves trifoliolate; leaflets with prickly midrib
and toothed margin. Inflorescence an axillary or terminal panicle. Flowers 45merous, unisexual; sepals small; petals to 5 mm long, oblong; stamens 45, female
flowers with staminodes; ovary 45-carpellate, on short stalk. Fruit a small, round
drupe. Seeds smooth, angular; embryo curved, embedded in fleshy endosperm.
355
Erect shrubs or trees, unarmed. Leaves unifoliolate, opposite. Flowers unisexual or

bisexual ......................................................................................................................7
7.
Flowers unisexual; buds globose; petals imbricate; filaments glabrous; ovaries and
fruits glabrous................................................................................7. Maclurodendron
Flowers bisexual; buds cylindrical; petals valvate; filaments ciliate at base; ovaries and
fruits hairy or sparsely so........................................................................1. Acronychia
8.
Twigs and branchlets unarmed. Ovary 25-carpellate (or rarely 6-carpellate in

Merrillia); ovules 12 per locule (or 810 in Merrillia). Fruits rather small (or large in
Merrillia), without pulp-vesicles, dry or slightly juicy (or mucilaginous in Merrillia)
....................................................................................................................................9
Twigs and branchlets (especially in young plants) and/or older branches armed (or
unarmed or with spine-like paraphylls in some Severinia; unarmed or nearly so in some
Pleiospermium). Ovary 25-carpellate (or 618-carpellate in Citrus), ovules 2 per
locule or more (or rarely 1). Fruits small to large, with or without pulp-vesicles, usually
juicy or mucilaginous.....................................................................................13
9.
Inflorescence a terminal, flat-topped corymb. Petals valvate in bud. Cotyledons thin,

folded.................................................................................................11. Micromelum
Inflorescence a terminal or axillary panicle, raceme or cyme, or 12-flowered. Petals
imbricate in bud. Cotyledons thick, flat......................................................................10
10. Inflorescence 12-flowered. Petals c. 2.5 cm long. Ovary 56-carpellate. Fruits 9 cm or

more in diameter; pericarp thick, leathery.................................................10. Merrillia
Inflorescence a many-flowered panicle, raceme or cyme. Petals rarely exceeding 2 cm
long. Ovary 25-carpellate. Fruits less than 2.5 cm in diameter; pericarp thin, glandular
..................................................................................................................11
11. Style persistent, thick, shorter than the ovary. Locules 12-ovulate. Young growth
densely reddish hairy................................................................................5. Glycosmis
Style deciduous or persistent, shorter than the ovary or as long or longer. Locules 2ovulate (rarely 1-ovulate). Young growth not as above...............................................12
12. Petals 1021 mm long or more. Style as long as or longer than the ovary, stigma
capitate. Floral buds cylindrical or oblong.................................................13. Murraya
Petals 36 mm long. Style shorter than the ovary, stigma flattened. Floral buds globose
or ovoid......................................................................................................4. Clausena
13. Stamens twice as many as the petals. Fruits 13 cm in diameter; locules containing
mucilage or scant flesh, pulp-vesicles lacking. Erect or scandent shrubs or trees........14
Stamens 24 times as many as the petals or more. Fruits variable in size, to 20 cm in
diameter; locules containing juice-filled pulp-vesicles. Erect shrubs or trees..............17
14. Scandent shrubs armed with solitary, recurved or straight spines. Leaves trifoliolate
and/or unifoliolate.....................................................................................................15
Erect shrubs or small trees armed with solitary or paired, straight spines. Leaves
unifoliolate or simple.................................................................................................16
356
RUTACEAE (JONES)
15. Leaves trifoliolate (or unifoliolate on juvenile stems). Petioles 730 cm long, rarely
shorter...........................................................................................................................
Luvunga Buch.-Ham. in Wall. ex Wight & Arn.
Prodr. 1 (1834) 90; Hooker f. l.c. (1875) 508; Kurz l.c. (1877) 193; Merrill l.c. (1921) 315,
l.c (1929) 115; Ridley l.c. (1922) 354; Craib l.c. 235; Masamune l.c. 360; Backer &
Bakhuizen f. l.c. 105; Burkill, EPMP (1966) 1396; Swingle l.c. (1967) 265; Anderson l.c.
308; Perry l.c. 371; Stone, Proc. Ac. Nat. Sc. Phil. 137 (1985) 221.
c. 12 species; India, Sri Lanka, Burma, Indo-China, Malesia; 78 species in Sabah

and Sarawak.
Woody climbers with hook-like or (in juvenile plants) straight axillary spines.
Leaves trifoliolate or (in juveniles) unifoliolate. Leaflets coriaceous; margin entire;
petiolules pulvinate. Inflorescence an axillary panicle or raceme. Flowers 45merous; calyx cup-like, slightly lobed; petals oblong-linear; stamens usually 8 or
10; ovary 24-carpellate, on short stalk, style thick. Fruit a glandular berry, thickwalled. Seeds 13 per fruit.
Leaves unifoliolate. Petioles to 2.5 cm long, usually shorter...........................................
Paramignya Wight
Ill. Ind. Bot. 1 (1840) 108, 110; Hooker f. l.c. (1875) 509; Kurz l.c. (1877) 193; Ridley l.c.
(1922) 355; Burkill, Gard. Bull. S.S. 5 (1931) 213, l.c. (1966) 1691; Craib l.c. 235; Swingle
l.c. (1967) 270; Stone l.c. (1972) 384; Perry l.c. 371; Corner l.c. 669.
c. 12 species; India, Sri Lanka, Burma, S China, Indo-China, Thailand, Malesia,

Australia; 34 species in Sabah and Sarawak.
Woody climbers or (rarely) shrubs, with hook-like or short straight axillary spines.
Twigs zigzag. Leaves unifoliolate, margins entire; petioles short, pulvinate.
Inflorescences axillary, often solitary. Flowers 45-merous, often large; calyx cuplike, lobed; petals lanceolate-oblong; stamens 8 or 10; ovary 35-carpellate,
stalked. Fruits round or lobed, glandular, resinous. Seeds 1 to several per fruit,
ovoid or flattened.
16. Ovary ovoid, glabrous. Fruits ovoid to ellipsoid, angular, green when ripe. Seeds
flattened. Leaves unifoliolate.......................................................................10. Merope
Ovary bottle-shaped, sparsely hairy. Fruits obovoid to pyriform, yellow-orange when
ripe. Seeds thick. Leaves simple.....................................................12. Monanthocitrus
17. Stamens 48 times as many as the petals. Pulp-vesicles tapered at both ends (fusiform),
on slender stalks............................................................................................13. Citrus
Stamens twice as many as the petals. Pulp-vesicles cylindrical or conical or irregular in
shape, stalkless or nearly so.......................................................................................18
18. Ovary 5-carpellate, locules 2226-ovulate. Fruits 811 cm in diameter, obovoid to oblong
...........................................................................................................2. Burkillanthus
Ovary 2- or 45-carpellate, locules 12-ovulate. Fruits to 2.5 cm in diameter, usually
less, round or nearly so..............................................................................................19
19. Disc small, annular to cup-like. Fruits 1525 mm in diameter, pericarp rough-glandular.
Petioles winged or narrowly so, articulated with the leaf-blades.......14. Pleiospermium
Disc cup-like, enclosing the base of the ovary. Fruits 615 mm in diameter, pericarp
smooth. Petioles wingless, not articulated with the leaf-blades.................15. Severinia
357
1. ACRONYCHIA J. R. Forst. & G. Forst.

(Greek, akron = tip, onychos = claw; the claw-like tip of the petals)
Char. Gen. Pl. ed. 1 (1775) 27; Hooker f. l.c. (1875) 498; Kurz l.c. (1877) 183; King, J. As. Soc. Beng.
62, 2 (1893) 214; Ridley l.c. (1922) 347; Merrill l.c. (1929) 113; Craib l.c. 221; Masamune l.c. 357;
Backer & Bakhuizen f. l.c. 101; Burkill l.c. (1966) 40; Stone l.c. (1972) 371; Hartley, J. Arn. Arb. 55
(1974) 469; Anderson l.c. 307; Perry l.c. 361; Whitmore, Tantra & Sutisna l.c. 304.
Shrubs or trees or rarely climbers. Branches unarmed. Leaves opposite, trifoliolate or

unifoliolate, petioles wingless; leaflets pinnately veined, margin entire, articulated at the
base. Inflorescences axillary, paniculate, subcorymbose, or 1- to few-flowered. Flowers
bisexual, 4-merous; sepals 4, free or fused at base, imbricate; petals 4, free, valvate, pelluciddotted, white or cream to greenish or yellowish, becoming reflexed; stamens 8, unequal,
alternately long and short, filaments flattened and tapering to a sharp apex, usually densely
ciliate at the base, becoming reflexed, anthers ovoid to ellipsoid, basifixed; disc shallowly
8-lobed; ovary 4-carpellate, with septicidal fissures or without, ovules 2 per locule, style
twisted, stigma shallowly 4-lobed. Fruit a 4-loculate drupe, with septicidal fissures or
without; epicarp semi-fleshy, spongy or woody when dry, with or without evident mesocarp;
endocarp cartilaginous to parchment-like. Seeds 12 per locule, narrowly ellipsoid, slightly
bent; testa dull to shiny, smoothish to finely rugose; endosperm fleshy; embryo straight.
Distribution. 42 species; from India to SW China, and throughout Malesia to E Australia
and New Caledonia. The greatest number are found in Australia, Irian Jaya (Indonesia) and
Papua New Guinea, including several rare and endemic species. One species, A. pedunculata,
occurs in Sabah and Sarawak.
Ecology. Found in primary and secondary rain forests, from the lowlands and hills to cloud
forests and subalpine habitats above 3000 m and in coastal scrub.
Acronychia pedunculata (L.) Miq.
Fig. 1.
(Latin, pedunculatus = slender-stalked; the inflorescence)

Fl. Ned. Ind., Suppl. (1861) 532; Merrill l.c. (1929) 113; Craib l.c. 221; Masamune l.c. 357; Stone l.c.
(1972) 371; Hartley l.c. 549; Perry l.c. 361; Whitmore, Tantra & Sutisna l.c. 305. Basionym:
Jambolifera pedunculata L., Sp. Pl. 1 (1753) 349. Type: Jambolifera L., Fl. Zeyl. (1747) 139.
Synonyms: Acronychia laurifolia Blume, Cat. (1823) 63; A. arborea Blume, Bijdr. (1825) 244;
Melicope conferta Blanco, Fl. Filip. ed. 2 (1845) 205; Acronychia apiculata Miq. l.c. (1861) 532.
Shrub or small to large tree to 30 m tall and 50 cm in diameter; bole to 23 m tall. Younger
branchlets glabrous to finely hairy. Leaves unifoliolate; leaflet usually elliptic or ellipticoblong to obovate or oblanceolate, 3.524.5 x 28.5 cm, thinly leathery; base cuneate or
occasionally rounded, apex acuminate or occasionally blunt or rounded; lateral veins 714
pairs; reticulations conspicuous above and below; petioles 0.55 cm long. Inflorescences
few- to many-flowered, 225 cm long, axes glabrous or nearly so, pedicels 212 mm long.
Flowers 412 mm long; sepals triangular or rounded, minute; petals linear or oblanceolate,
glabrous to sparsely hairy; disc glabrous; ovary hairy, rarely only at apex; style hairy at
base. Fruits to 15 mm in diameter, usually subglobose to ellipsoid or conical, more or less
358
RUTACEAE (JONES)
359
360
RUTACEAE (JONES)
4-lobed or longitudinally ribbed, usually sparsely hairy, base with a ring of dense hairs,
apex rounded to shortly tipped; epicarp when dry to 3 mm thick; mesocarp woody or nearly
so; endocarp cartilaginous. Seeds reddish black to black, 37 mm long.
Distribution. India, Sri Lanka, eastward to Taiwan and throughout Malesia to Papua New
Guinea. In Sabah, frequent in primary and secondary forests, often on hill-sides, from near
sea-level to 1500 m. In Sarawak, collected only twice (Baram and Lundu Districts). Also
occurs in Brunei and Kalimantan.
Uses. The wood, roots, bark and leaves are used medicinally to treat rheumatism, scabies
and colic, and as pain reliever (China, Indo-China, Indonesia). The roots are used as a fish
poison (Indo-China). The wood is rarely used in construction and for making charcoal, and
its young leaves are eaten as a condiment (Java).
2. BURKILLANTHUS Swingle
(I. H. Burkill, 18701965, one time director of the Singapore Botanic Gardens)
J. Arn. Arb. 20 (1939) 255, l.c (1967) 294; Stone l.c. (1972) 373; Whitmore, Tantra & Sutisna l.c. 305.
Trees; crown broad, dense. Branches armed, drooping in older trees. Leaves alternate, 13foliolate; petioles narrowly winged to nearly wingless, pulvinoid at base; leaflets thinly
leathery, becoming stiff when older, glabrous, articulated at the base. Inflorescences
axillary, few-flowered clusters, pedicels short. Flowers bisexual, large, 5-merous; sepals 5,
nearly free to the base, with large oil-glands; petals 5, gland-dotted distally; stamens 10,
free, slender, anthers small, oblong; disc cylindrical; ovary 5-carpellate, obovoid, sparsely
hairy, glandular on upper half, fluted with 5 grooves; ovules 2226 per locule in 2 rows,
style slender, glabrous, stigma globose. Fruit a large, obovoid to oblong berry; exocarp
thick, leathery, light green becoming pale yellow, roughened with numerous oil-glands;
endocarp thin, woody; pulp-vesicles sessile, cylindric, tips acute, pale yellow. Seeds numerous,
large, cream with brown cap, embedded in mucilage; monoembryonic.
Distribution. 1 species, restricted to N Sumatra, Peninsular Malaysia and Sarawak. Uncommon,
and possibly endangered.
Ecology. Found as solitary trees or small populations of several trees, on stream banks,
slopes, and ridge tops in primary and secondary forest, on rocky and sandy soils, from 5 to
200 m.
Burkillanthus malaccensis (Ridl.) Swingle
Fig. 2.
(of Malacca)
l.c. (1939) 257, l.c. (1967) 295; Whitmore l.c. 373; Stone l.c. (1972) 373, Fed. Mus. J. 23 (1978) 114;
Whitmore, Tantra & Sutisna l.c. 305. Basionym: Citrus malaccensis Ridl. l.c. (1922) 359. Type:
Goodenough 1273, Malacca, Nyalas (holotype SING; isotypes K, US).
361
Small tree to 15 m tall and 40 cm in diameter; bole to 4 m tall, fluted. Branches generally
armed with stout, straight, usually paired spines. Leaves unifoliolate (on juvenile growth) or
23-foliolate; terminal leaflets lanceolate or elliptic-lanceolate, 12.527 x 4.511.5 cm; base
broadly cuneate, margins subentire to slightly coarse-crenate, apex acute or acuminate;
lateral veins 1218 pairs, deeply sunken above; petioles 45 cm long; petiolules 22.5 mm
long, pulvinoid; lateral leaflets to 8 x 4 cm, petioles 14 cm long, otherwise similar to
terminal leaflet. Inflorescences 14-flowered, pedicels 57 mm long. Flower 56 cm
across; sepals lanceolate-acuminate; petals 22.5 cm long, subspathulate; ovary 6.57 mm
long. Fruits 108 x 811 cm; exocarp c. 1 cm thick; endocarp 23 mm thick, pulp-vesicles
1.53 cm long. Seeds broadly obovoid, tapering to an acute base, 2.22.7 cm long and c. 8
mm thick; testa thin, slightly wrinkled; cotyledons pale buff.
Vernacular name. Sarawaklimau hantu (Malay).
Distribution. Throughout the range of the genus. In Sarawak very uncommon, collected
once in 1961 (S. 15848) from primary riparian forest in the Labang FR, Bintulu.
3. CITRUS L.
(Latin name for the citron tree)
Sp. Pl. 1 (1753) 401; Hooker f. l.c. (1875) 514; Kurz l.c. (1877) 195; Merrill l.c. (1921) 315; Ridley
l.c. (1922) 358; Burkill l.c. (1931) 220, l.c. (1966) 568; Craib l.c. 236; Masamune l.c. 357; Santiago,
Bull. Mal. Div. Agr. 111 (1962) 1; Backer & Bakhuizen f. l.c. 107; Swingle l.c. (1967) 358; Stone l.c.
(1972) 374, l.c. (1985) 226; Perry l.c. 361; Corner l.c. 659; Whitmore, Tantra & Sutisna l.c. 305;
PROSEA 2 (1991) 119, 325.
Shrubs or small to large trees. Branches generally armed with axillary, solitary spines, older
branches often unarmed. Young twigs angular, flattened, becoming terete. Leaves alternate,
simple or unifoliolate, blades thinly leathery to leathery, glabrous or short-hairy below;
margins entire to serrate or crenulate; petioles often winged or emarginate, usually
articulated with the blade. Inflorescences axillary, short corymbose racemes or cymes, or
solitary. Flowers bisexual or male only, 45-merous, fragrant or not; calyx cup-like; sepals
45; petals 48, commonly 5, oblong-linear, thick; stamens about 48 times the number of
petals, free or fused at the base and in bundles; disc annular; ovary subglobose, (6)818carpellate, usually glabrous, ovules (1)48(12) per locule in 2 rows, style cylindric,
stigma globose-capitate. Fruit a fleshy hesperidium; pericarp leathery, outer layer (exocarp or
flavedo) densely glandular and pigmented, middle layer (mesocarp or albedo) white, dry,
inner layer (endocarp) membranous, thin, flesh of stalked fusiform, pulp-vesicles filled with
watery, sweet or bitter juice, with or without distinct oil-droplets. Seeds few to many,
angular-obovoid, often flattened, pale; mono- or polyembryonic; embryo white or green.
Distribution. 17 species; from India, Sri Lanka and Burma to Japan, China, Taiwan and
Indo-China, and throughout Malesia eastward to the Pacific Islands. Two species, C. halimii
and C. macroptera, are native to Sabah, the latter species also occurs in Sarawak. In
addition, a number of species have long been introduced and are cultivated in Sabah and
Sarawak. Numerous hybrids and variants of these species exist, many having unknown
origins which makes classification difficult.
362
RUTACEAE (JONES)
Stone (l.c. 1985) reported a collection of Citrus grandis from Ulu Segama, Sabah (SAN
75565) and suggested that it might be indigenous there, although he admitted that the
species could have been introduced into that area by early gold prospectors. On this basis
and until more specimens are available, this species is excluded from the present treatment.
Uses. Citrus fruits are eaten fresh, canned, preserved or candied. The juice is extracted and
made into drinks or concentrates. Pectin and essential oils are made from the rind, while
waste fruit-pulp is processed into cattle-feed. Citric acid is manufactured from certain
species. The leaves, flowers, and fruits of some species are sources of expensive essential
oils (oil of neroli, bergamot oil) used in perfumery. Various plant parts (leaves, flowers,
fruits, and seeds) are used to flavour foods and often enter into native medicinal remedies.
Some species are reported to show insecticidal and germicidal activity. Flavonoids from the
inner cortex are said to have anti-tumour properties. Some species are utilised as rootstock
for the cultivation of commercially important citrus because of their resistance to certain
diseases. Citrus is less commonly used as a source of wood for cabinetry. In Malaysia, the
plants have reportedly been used for occult purposes.
Taxonomy. The taxonomic delimitation of species, subspecies, and forms of Citrus is
complicated by the peculiar reproductive behaviour found in the genus including
hybridisation, polyembryony (asexual seed production), the spontaneous production of
autotetraploids, and mutations. Two widely opposing systems of classification exist. That of
Tanaka (Revisio Aurantiacearum - IX, Jap. Soc. Prom. Sci., Ueno, Tokyo, 1954) proposes 145
species (increased to 157 a few years later), while that of Swingle (l.c.1967) recognises only
16 species. This difference reflects a lack of agreement on what constitutes a species and
whether supposed hybrids should be assigned the rank of species. Swingle, whose system is
followed in this treatment, did not consider valid many of Tanaka's species which were
clearly horticultural forms, and mostly the result of interspecific hybridisation. Several other
systems of classification put forward by various authors are intermediate. Recent studies
employing numerical taxonomy, chemosystematics, and other experimental methods have
attempted to elucidate relationships among taxa and clarify the "species" problem in Citrus.
Several such studies support the notion that the genus centres around three basic species or
"biotypes", viz. C. grandis, C. medica, and C. reticulata.
Key to Citrus species

1.
Petioles narrowly winged or wingless, or if broadly winged, never as broad as the leaf
blades. Flowers large, usually 2.54.5 cm wide, fragrant. Stamens free or united in
bundles........................................................................................................................2
Petioles broadly winged, often as broad as the leaf-blades. Flowers small, usually less
than 2 cm wide, not fragrant. Stamens free..................................................................9
2.
Petioles wingless, not articulated with the leaf-blade. Flowers perfect or staminate.
Fruits usually oblong, pericarp thick. Cultivated............................................................
C. medica L.
Sp. Pl. 2 (1753) 782. Synonym: C. aurantium var. medica Wight & Arn. l.c. 98.
Probably native to NE India and upper Burma. Cultivated widely in the tropics and
subtropics for its fruits. Vernacular names: citron, limau susu.
363
Petioles winged, often narrowly so, articulated with the leaf-blade. Flowers all perfect
(except in C. limon). Fruits ovoid or globose or pear-shaped, pericarp thin (except in C.
grandis).......................................................................................................................3
3.
Petioles very narrowly winged. Flowers perfect or staminate; stamens usually more than
4 times the number of petals. Fruits ovoid. Cultivated............................................
C. limon (L.) Burm. f.
Fl. Ind. (1768) 173. Basionym: Citrus medica var. limon L. l.c. (1753) 782. Synonym: Citrus
limonum Risso, Ann. Mus. Hist. Nat. Paris 20 (1813) 201.
Probably native to S Asia and now widely cultivated in the tropics and subtropics
for its fruits. Vernacular names: lemon, limau mata kerbau.
Petioles narrowly to broadly winged. Flowers usually perfect; stamens usually 4 times
the number of petals. Fruits globose or pear-shaped, sometimes slightly flattened........4
4.
Fruits with loose pericarp, easily detached. Seeds small, plump. Embryo green............5
Fruits with adherent pericarp, not easily detached. Seeds various. Embryo not green....6
5.
Leaves somewhat rhomboid, tips acute. Fruits small to medium-sized, sometimes

flattened. Cultivated......................................................................................................
C. reticulata Blanco
Fl. Filip. (1837) 610. Synonyms: Citrus nobilis Andrews, Bot. Repos. 9 (1809) 608, non
Lour. (1790); Citrus deliciosa Tenore, Ind. Sem. Hort. Neap. (1840) 9.
Native to SE Asia. Widely grown for its edible fruits. Vernacular names: mandarin,
limau manis.
Leaves elliptic, tips rounded. Fruits small, globose. Cultivated.......................................
C. microcarpa Bunge
Mem. Ac. Imp. Sc. St. Petersb. 2 (1833) 84. Synonyms: Citrus mandurensis Lour., Fl.
Cochinch. 2 (1790) 467; Citrus mitis Blanco l.c. (1837) 610.
Originated in China as a natural hybrid between Citrus reticulata var. austera

Swingle (sour mandarin) and a species of Fortunella (kumquat); currently classified as
xCitrofortunella microcarpa (Bunge) Wijnands, Baileya 22 (1984) 134. Widely
grown in SE Asia for its acid juice. Vernacular names: calamondin, limau kesturi.
6.
Petioles broadly winged. Fruits large to very large, usually 1020 cm in diameter, round
to pear-shaped. Seeds large, rough, monoembryonic. Cultivated..........................
C. grandis (L.) Osbeck
Dagb. Ostind. Resa (1757) 98. Basionym: Citrus aurantium var. grandis L. l.c. (1753) 783.
Synonyms: Citrus decumana L., Syst. ed. 12 (1767) 508; Citrus maxima (Burm. f.) Merr.,
Interp. Rumph. Herb. Amb. (1919) 296.
Native to Malesia; widely cultivated in SE Asia for its edible fruits. Vernacular
names: pummelo, limau besar.
Petioles narrowly winged to nearly wingless. Fruits small to medium-sized, generally
49 cm in diameter. Seeds various, usually polyembryonic (except in C. halimii).........7
7.
Fruits medium-sized, 59 cm in diameter; pulp orange. Cultivated................................
364
RUTACEAE (JONES)
C. sinensis (L.) Osbeck

Reise Ostind. China (1765) 250. Basionym: Citrus aurantium var. sinensis L. l.c. (1753)
782.
Probably native to the region between China and Vietnam. Cultivated widely in the
subtropics and tropics for its edible fruits. Vernacular names: sweet orange, limau
manis.
Fruits small, 47 cm in diameter; pulp greenish to yellowish. Cultivated or wild.........8
8.
Leaves small, mostly 57.5 cm long. Stamens 2025. Fruits globose to ovoid,
sometimes with apical papilla; pericarp thin, smooth. Cultivated...................................
C. aurantifolia (Christm. & Panz.) Swingle
J. Wash. Ac. Sc. 3 (1913) 465. Basionym: Limonia aurantifolia Christm. & Panz., Pfl. Syst.
1 (1777) 618. Synonym: Citrus acida Roxb., Fl. Ind. ed. 2, 3 (1832) 390.
Native to N India and Burma, or N Malesia. Cultivated throughout the tropics and
warm subtropics for its fruits. Vernacular names: lime; limau nipis.
Leaves large, 815 cm long or more. Stamens 1820. Fruits subglobose to slightly
pyriform; pericarp thick, smooth to bumpy. Hills and mountains...............1. C. halimii
9.
Petiolar wings with subentire margins. Fruits large, 815 cm in diameter; pericarp
smooth...............................................................................................2. C. macroptera
Petiolar wings with serrate-crenulate margins. Fruits small, 57 cm in diameter;
pericarp irregularly bumpy. Cultivated...........................................................................
C. hystrix DC.
Cat. Pl. Hort. Bot. Monsp. (1813) 97. Synonym: Citrus torosa Blanco l.c. (1832) 609; Citrus
papeda Miq., Fl. Ned. Ind. 1 (1859) 530.
Of unknown origin but widely naturalised in Malesia, Sri Lanka and Burma.
Cultivated for its fruits. Vernacular names: Mauritius papeda, limau purut.
1. Citrus halimii Stone

(Sultan Abdul Halim Nu'azzam Shah Ibni Almarhum Sultan Badlishah, Sultan of Kedah and
former King of Malaysia)
Biotropica 5 (1973) 102, l.c. (1978) 114; Stone & Jones, Proc. Ac. Nat. Sc. Phil. 140 (1988) 267; Ng
l.c. 495; Whitmore, Tantra & Sutisna l.c. 305; PROSEA 2 (1991) 119. Type: Stone 9550, Peninsular
Malaysia, Pahang, G. Nuang (holotype KLU; isotypes KEP, L, SING, US).
Medium-sized or large tree to 25 m tall and 60 cm in diameter; trunk straight, cylindric;

crown meagre, of few ascending branches. Spines on youngest branches of saplings straight
to 2.5 cm long, absent on adult plants. Leaves unifoliolate, elliptic or slightly ovate, 815 x
3.57.5 cm (larger on vigorous young shoots), thinly leathery; margin subentire or
subcrenulate, apex acute to shortly acuminate and emarginate; lateral veins 711 pairs;
petioles usually 12 cm long, very narrowly winged or wingless, articulated with the blade.
Inflorescences solitary, pedicels 13.5 mm long. Flowers 5-merous; sepals deltoid; petals
ovate-elliptic, white; stamens 1820, free or in 23 bundles; disc thin; ovary (6)910365
carpellate, ovules 13(5) per locule, style columnar, stigma flat, angled. Fruits subglobose to
slightly pyriform, 57 cm in diameter, green ripening glossy deep yellow, smooth or bumpy;
pericarp c. 6 mm thick, glandular; pulp-vesicles pale greenish to yellowish, juicy, acrid oildroplets absent. Seeds numerous, to 2.1 cm long, testa veiny-reticulate; monoembryonic;
cotyledons white.
Vernacular name. Sabahlimau hutan (Malay).
Distribution. S Thailand, Peninsular Malaysia, and Borneo. In Sabah, uncommon (Kota
Marudu, Ranau and Tambunan districts), occurring as solitary trees on slopes and ridges in
primary submontane forest, at 9001800 m. Not reported from Sarawak, Brunei and
Kalimantan.
Uses. The fruits are eaten as a condiment (rinds?), and are said to be thirst-quenching
(Peninsular Malaysia). The flesh is said to be acidic and pleasant to taste.
2. Citrus macroptera Montr.
Fig. 3.
(Greek , makros = large, pteron = wing; the leaf-stalk)

Mem. Ac. R. (Imp.) Lyon Sc. 10 (1860) 187; Burkill l.c. (1931) 220, l.c. (1966) 576; Santiago l.c. 131;
Swingle l.c. (1967) 395; Stone l.c. (1972) 374; Corner l.c. 661; PROSEA 2 (1991) 326. Type:
Montrouzier, s.n., New Caledonia, Ile Art (LY). Synonyms: Citrus papuana F.M. Bail., Contr. Fl.
Brit. N. Guin. (1903) 1; C. aurantium subsp. saponacea Safford, Contr. US. Nat. Herb. 9 (1905) 226;
C. hystrix (non DC.) Ridl. l.c. (1922) 358.
var. macroptera
Small or medium-sized tree to 20 m tall and 50(100) cm diameter, generally much smaller;
crown broad, dense. Branches and twigs armed with straight spines to 1.5 cm long, longer
on juvenile growth. Leaves unifoliolate, to 30 cm long, blade ovate-lanceolate, 515 x 36.5
cm; base rounded, margin shallowly crenate, apex acuminate; petioles broadly winged, with
wing as large as the blade, obovate to nearly spathulate, margin subentire to crenate in the
upper half, articulated with the blade. Inflorescence a few-flowered cluster. Flowers 1.32
cm in diameter; buds subglobose; sepals triangular, small; petals oblong, concave; stamens
c. 20, free; ovary 1012-carpellate, slightly hairy, style thick, stigma depressed. Fruits
globose to pyriform, sometimes depressed, 815 cm in diameter or larger, smooth, green
turning pale yellow; pericarp to 2.5 cm thick, segments 1012, pulp-vesicles greenish
yellow, juice scant, bitter, acrid oil-droplets present. Seeds 12 per segment, flattened, to c.
2 cm long; monoembryonic.
Vernacular names. Sabah and Sarawaklimau hantu, limau hutan (Malay).
Distribution. The species and the variety are distributed in Indo-China, Malesia, New
Caledonia, and Polynesia. Three varieties are known to occur, only one of which (var.
macroptera) is in Sabah and Sarawak. Found occasionally in SW Sabah (Pinangah,
Kinabatangan, Lahad Datu, and Tawau districts) as solitary trees along the banks of streams
and on forested ridges and hills in primary forest, at 150600 m. In Sarawak,
366
RUTACEAE (JONES)
367
368
RUTACEAE (JONES)
known only from a single collection made from lowland riverine forest (Taufik, s.n., Kg.
Bedup, Serian; KLU).
Uses. The fruits are reported to be edible (Philippines, Indonesia, Sabah), and are said to
cause impotence (Indonesia). The leaves are eaten as a vegetable and used medicinally
(Indonesia). Sticks are made from its wood (Papua New Guinea).
4. CLAUSENA Burm. f.
(Clausen, a botanist known to Burmann)
Fl. Ind. (1768) 243; Hooker f. l.c. (1875) 503; Kurz l.c. (1877) 187; Merrill l.c. (1921) 314, l.c. (1929)
115; Ridley l.c. (1922) 352; Craib l.c. 231; Masamune l.c. 358; Backer & Bakhuizen. f. l.c. 103;
Burkill l.c. (1966) 584; Swingle l.c. (1967) 209; Stone l.c. (1972) 375; Anderson l.c. 307; Perry l.c.
363; Corner l.c. 662; Ng l.c. 495; Whitmore, Tantra & Sutisna l.c. 305; PROSEA 2 (1991) 141.
Shrubs or small trees. Branches unarmed; branchlets short-hairy. Leaves alternate, pinnate,
rachis sometimes winged; leaflets alternate, usually 59(31), often with asymmetric base,
articulated with the petiolules. Inflorescences lax panicles or racemes, terminal or axillary.
Flowers bisexual, small, 45-merous; buds small, subglobose or ovoid; sepals 45, calyx
cup-like, lobed; petals elliptic or oval, free, imbricate; stamens (7)8 or 10, free, unequal, in
2 whorls, the outer opposite the sepals and longer, filaments inflated or flattened at base,
often narrowed at apex, glabrous, anthers ovate or elliptic, dorsifixed; disc annular,
glabrous; ovary 25-carpellate, sometimes obscurely lobed, usually hairy, glandular, seated
on a glabrous, hourglass-shaped gynophore, ovules (1)2 per locule, style short, stout,
deciduous or persistent, usually narrowed where it joins the ovary, stigma flattened,
obscure. Fruit an ovoid or oblong berry. Seed usually 1 per fruit; cotyledons green.
Distribution. c. 29 species; from tropical Africa to India, S China, Taiwan, Indo-China,
throughout Malesia and N Australia. Two species, C. calciphila and C. excavata, are in
Sabah, the latter occurring also in Sarawak, Brunei and Kalimantan. A third species, C.
lansium, is sparingly cultivated.
Key to Clausena species

1.
Leaves 1531-foliolate. Flowers mostly 4-merous. Stamens 8..................2. C. excavata

Leaves 511-foliolate. Flowers 5-merous. Stamens 10..................................................2
2.
Leaves 57-foliolate. Stigma flat. Fruits oblong-ellipsoid, to 1.5 cm long.....1. C.

calciphila
Leaves 911-foliolate. Stigma 5-lobed. Fruits ovoid-globose, to 2.5 cm long. Cultivated
.......
C. lansium (Lous) Skeels
USDA Bur. Pl. Ind. Bull. 168 (1909) 31. Basionym: Quinaria lansium Lour., Fl. Cochinch.
(1790) 272. Synonyms: Cookia wampi Blanco l.c. 358; Clausena wampi (Blanco) Oliv., J.
Linn. Soc. Bot. 5, Suppl. 2 (1861) 34.
Native to S China and Indo-China. Widely cultivated in the tropics and subtropic
for its edible fruits. Vernacular names: wampee, wampi.
369
1. Clausena calciphila Stone

(Latin, calx = chalk, Greek, philos = loving; referring to its preference for limestone habitats)
l.c. (1978) 111. Type: Erwin & Paul S. 27430, Sarawak, Kuching, Bukit Pa'it (SAR).
Small tree to 10 m tall. Leaves to 60 cm long, commonly 57-foliolate; leaflets ellipticovate, the lowest ones 713 x 47 cm , the lateral and terminal ones larger, 1525 x 1216
cm; base rounded, obliquely asymmetric, margin entire to subentire, apex acuminate; lateral
veins 58(10) pairs, together with midrib raised below; petioles 1012 cm long, rachis
glabrescent, wingless; petiolules 59 mm long. Inflorescence a loose, terminal panicle, 20
cm long or more during flowering, to 40 cm long in fruit; buds subglobose, c. 2 mm long.
Flowers 5-merous; sepals 5, broadly triangular, minute, margin ciliate; petals 5, elliptic, c. 3
mm long, glabrous, margin hyaline; stamens 10, alternately long and short, filaments
narrowly elliptic, anthers oblong-ovoid, connectives 1-glandular; disc annular, yellowish;
ovary 5-carpellate, globose, glabrous, papillate-glandular, style shortly cylindrical. Fruits
oblong-ellipsoid, 1.51.6 cm long, gland-dotted.
Distribution. Endemic to Sarawak. Uncommon, known from only three localities in
Kuching (Bt. Pa'it, Bt. Bra'ang) and Baram (Bt. Mentagai) districts; on rocky limestone
slopes, at 150300 m.
2. Clausena excavata Burm. f.
Fig. 4.
(Latin, excavatus = to make hollow; the bases of the filaments)

l.c. 243; Hooker f. l.c. (1875) 504; Kurz l.c. (1877) 188; Merrill l.c. (1921) 314, l.c. (1929) 115; 314;
Ridley l.c. (1922) 352; Craib l.c. 231; Masamune l.c. 358; Backer & Bakhuizen f. l.c. 104; Burkill l.c.
(1966) 585; Swingle l.c. (1967) 212; Stone l.c. (1972) 375; Anderson l.c. 308; Perry l.c. 363; Corner
l.c. 662; Whitmore, Tantra & Sutisna l.c. 306. Type: Burmann 29, India, Dauhon Kongeere (L).
Synonyms: Clausena javanensis Raeusch. ex DC., Prodr. 1 (1824) 538; Murraya burmanni Spreng.,
Syst. Veg. 2 (1825) 315; Amyris graveolens Buch.-Ham. ex Steud., Nom. Bot. ed. 2, 1 (1840) 81.
var. excavata
Small tree to 15 m tall and 20 cm diameter. Branchlets softly hairy. Leaves 2050 cm long,
rachis slender, cylindrical, not winged; leaflets 1531, oblong-ovate to lanceolate or slightly
crescent-shaped, 29 x 1.54 cm, thinly leathery, glabrous above, sparsely hairy below;
base rounded, obliquely asymmetric, margins toothed, apex tapered or acuminate; petiolules
short. Inflorescence a much-branched panicle, terminal, 1030 cm long, pyramid-shaped,
branches hairy. Flowers 4(5)-merous, buds round; calyx minute, hairy; petals oval, 3.55
mm long, glabrous, yellowish or greenish; stamens 8, filaments inflated and concave at
base; ovary ovoid to ellipsoid, hairy or hirsute, slightly lobed, style cylindrical, persistent,
not narrowed where it joins the ovary. Fruits broadly ellipsoid, 11.8 cm long, smooth,
glabrous; peel green ripening pink, translucent. Seed 1 per fruit, oblong.
Distribution. India, Burma, S China, Taiwan, Indo-China, and throughout Malesia. Two
varieties exist with only the widespread var. excavata in Sabah and Sarawak. Widespread
and common in Sabah and parts of Sarawak (Kuching and Baram districts). Also occurs in
370
RUTACEAE (JONES)
Ecology. In forests and open places in the lowlands and hills, especially disturbed areas
around villages and forest margin, and on limestone, from sea-level to 1500 m.
Uses. The leaves are reported to have insecticidal properties, and the roots, stems, leaves
and flowers are used to treat paralysis, stomach disorders, intestinal worms, and fever
(Taiwan, Indo-China, Peninsular Malaysia, Indonesia). Its wood has been used to make axehandles (Java).
5. GLYCOSMIS Corra
(Greek, glukus = sweet, osme = scent; sweet-smelling plant)
Ann. Mus. Hist. Nat. Paris 6 (1805) 384; Hooker f. l.c. (1875) 499; Kurz l.c. (1877) 184; Merrill l.c.
(1921) 314, l.c. (1929) 113; Ridley l.c. (1922) 348; Craib l.c. 222; Narayanaswami, Rec. Bot. Surv.
Ind. (1941) 1; Masamune l.c. 359; Backer & Bakhuizen f. l.c. 102; Burkill l.c. (1966) 1103; Swingle
l.c. (1967) 206; Stone l.c. (1972) 380, l.c. (1978) 75, l.c. (1985) 1; Anderson l.c. 308; Corner l.c. 665;
Whitmore, Tantra & Sutisna l.c. 306.
Shrubs or trees. Branches unarmed. Shoot tips and young inflorescences densely hairy, the
hairs reddish. Leaves alternate or opposite, pinnate or unifoliolate, rarely simple, petioles
wingless; leaflets 15(15), alternate or sometimes opposite, chartaceous or leathery;
margin entire or obscurely toothed, articulated at the base. Inflorescences axillary or
pseudoterminal racemes or cymose panicles. Flowers bisexual, 45-merous; sepals fused at
base, imbricate, glandular; petals ovate to elliptic, imbricate, greenish to whitish; stamens 8
or 10, usually unequal, filaments abruptly narrowed at the apex, anthers small, ovate to
elliptic, generally with a terminal gland, often with 1 or several glands on the connective;
disc annular, with small lobes; ovary 25-carpellate, club- to flask-shaped to nearly conical,
often raised on a stout gynophore, ovules 12 per locule, style short, thick, persistent, stigma
slightly broadened, faintly lobed. Fruits globose to oblong or ellipsoid berries, dry or
fleshy, outer wall glandular, white to pink or purplish. Seeds 12(3) per fruit, round to
plano-convex, thin-walled, cotyledons green, fleshy.
Distribution. c. 40 species; India, Sri Lanka, SE Asia, S China, Taiwan, Malesia, and Australia.
Seven species occur in Sabah and Sarawak, two of which (Glycosmis chlorosperma and G.
macrantha) are widespread and common. Glycosmis parviflora is occasionally cultivated in
Sabah and possibly also in Sarawak.
Taxonomy. Although readily distinguished from its closest relatives (Clausena and
Murraya), the chaotic state of its constituent species (65 names have been published to date)
has persisted due to lack of any critical taxonomic and nomenclatural review. Stone (l.c.
1978 and 1985), whose interpretation of the genus is followed here, admits that Glycosmis is
still imperfectly known and several taxonomic problems remain to be solved.
371
Key to Glycosmis species

1. Leaves predominantly pinnate, never uniformly unifoliolate. Ovary smooth.................2
Leaves predominantly unifoliolate, or some of them 23-foliolate. Ovary papillateglandular..............................................................................................3. G. lanceolata
2.
Ovary glabrous, 45-carpellate....................................................................................3

Ovary densely hairy, 23-carpellate...................................................6. G. sapindoides
3.
Inflorescences pseudoterminal.....................................................................................4
Inflorescences axillary.................................................................................................7
4.
Flowers large, 711 mm long; petals glabrous.............................................................5

Flowers medium-sized to small, rarely over 6 mm long; petals hairy, at least below.....6
5.
Leaves alternate; leaflets not over 15 cm long, lateral veins 59, petiolules to 7 mm
long....................................................................................................5. G. macrantha
Leaves sometimes opposite, leaflets over 15 cm long, lateral veins 1014, petiolules
1020 mm long.......................................................................................7. G. superba
6.
Lateral veins not prominently raised below. Sepals narrowly triangular, c. 2.5 mm long.
Fruits subellipsoid, c. 7.5 mm long...........................................4. G. longisepala
Lateral veins prominently raised below. Sepals deltoid-ovoid, 11.5 mm long. Fruits
globose, over 10 mm long...............................................................1. G. chlorosperma
7.
Leaflets predominantly entire. Inflorescences corymbose. Fruits ellipsoid, usually

purplish to black................................................................................2. G. cyanocarpa
Leaflets sometimes obscurely toothed. Inflorescences narrow, somewhat elongate.
Fruits globose, white to yellowish or pinkish. Cultivated................................................
G. parviflora (Sims) Little
Phytologia 2 (1948) 463. Basionym: Limonia parviflora Sims, Bot. Mag. (1823) t. 2416.
Synonym: Glycosmis citrifolia (Willd.) Lindl., Trans. Hort. Soc. Lond. 6 (1826) 72.
Native to S China and Indo-China. Introduced to horticulture and botanical gardens in

various parts of the World.
1. Glycosmis chlorosperma (Blume) Spreng.

(Greek, khloros = green, sperma = seed)
Syst. Veg. ed. 16, 4 (1827) 162; Narayanaswami l.c. 40; Masamune l.c. 359; Backer & Bakhuizen f.
l.c. 102; Swingle l.c. (1967) 207; Stone l.c. (1972) 382, l.c. (1978) 93, l.c. (1985) 2; Anderson l.c. 308;
Corner l.c. 666; Whitmore, Tantra & Sutisna l.c. 306. Basionym: Cookia chlorosperma Blume. l.c.
(1825) 135. Type: Blume, s.n., Java, Mt. Salak (L). Synonyms: Glycosmis monticola Ridl., J. Str. Br.
R. As. Soc. 75 (1917) 14; G. malayana Ridl. l.c. (1917) 12.
Shrub or small tree. Leaves pinnate, rarely unifoliolate, opposite or alternate, glabrous, with
or without axillary lanceolate paraphylls to 2.5 cm long; leaflets (1)35(7), ovate to
372
RUTACEAE (JONES)
oblong-lanceolate, or narrowly so, 520(30) x 29(14) cm, leathery; base acute, margin
entire, apex acuminate; lateral veins 511 pairs, rather distant, strongly inarching near the
margin, depressed above, conspicuously raised below; petiolules 35 mm long. Inflorescences pseudoterminal cymose-panicles, 216 cm long, branches rusty hairy, buds small,
minutely hairy. Flowers at tips of short branchlets, 5-merous; sepals ovate, to 2 mm long,
margin hyaline, ciliate; petals whitish, obovate-oblong, to 6 mm long, rusty hairy below,
margin membranous; stamens 10, alternating long and short, anthers ovate-cordate,
sometimes minutely gland-tipped; disc ring-like, yellow; ovary (4)5-carpellate, glabrous,
flask-shaped, on a thick gynophore, style short, equalling the ovary in length. Fruits
globose, 10 12.5 mm long.
Key to varieties
1.
Leaflets ovate, 57 cm long, lateral veins 5 pairs. Inflorescences 23 cm long...............

var. bidiensis Stone
l.c. (1985) 3. Type: J. & M.S. Clemens 20663, Sarawak, Kuching, Bidi Cave (holotype K).
Small tree. Leaves 35-foliolate; leaflets ovate, 57 x 22.3 cm; lateral veins 5
pairs. Inflorescences 23 cm long. Flowers small; buds 23 mm long; anthers
glandular; disc distinctly constricted, 0.80.9 mm wide; ovary 45-carpellate.
Endemic to Sarawak, where it is known only from the type locality, on limestone.
Leaflets oblong-lanceolate to elliptic, 530 cm long, lateral veins 911 pairs. Inflorescences longer than 5 cm...............................................................................................2
2.
Leaflets oblong-lanceolate, mostly less than 15 cm long. Inflorescences 7.511 cm

long...............................................................................................................................
var. chlorosperma
Shrub or small tree to 10 m tall and 15 cm diameter. Leaves 36-foliolate; leaflets
oblong-lanceolate, 515(20) x 28.5 cm; base sometimes slightly asymmetric;
lateral veins 910 pairs. Inflorescences 7.511 cm long. Flowers small; sepals c. 1
mm long; petals c. 2.5 mm long; ovary 5-carpellate, style almost lacking. Vernacular
name: segera (Malay). Throughout the range of the species. Widespread in Sabah
and Sarawak, in mixed dipterocarp forest on fertile clay loam soils, especially on
basic volcanic substrates, occasionally on limestone, from near sea-level to 800 m.
Leaflets oblong-elliptic, 1530 cm long. Inflorescences generally larger, to 16 cm
long..........
var. elmeri (Merr.) Tanaka
Med. Rijksherb. 69 (1931) 3; Stone l.c. (1978) 94, l.c. (1985) 3. Basionym: Glycosmis
elmeri Merr., Philip. J. Sc. 30 (1926) Bot. 400, Merrill l.c. (1929) 113, Masamune l.c. 359.
Type: Ramos & Edano 44150, Philippines, Sulu Archipelago, Tawitawi (UC).
Shrub or small tree to 10 m tall and 20 cm diameter. Leaves 34-foliolate; leaflets

oblong-elliptic, 1520(30) x 69(14) cm; base broadly acute; lateral veins 911
pairs. Inflorescences 616 cm long, side branches to 5 cm long. Flowers sessile,
buds brownish. Philippines (Sulu Archipelago), Borneo. Common in Sabah and
Sarawak, occurring in lowland and hill forest, usually on ridges and slopes, and
occasionally on limestone, from near sea-level to 1280 m.
373
Distribution. S Burma, S Thailand, Peninsular Malaysia, Sumatra, Java, Borneo (Sabah,

Sarawak, Kalimantan) and S Philippines. A variable species with seven recognised
varietes, three of which occur in Sabah and Sarawak.
2. Glycosmis cyanocarpa (Blume) Spreng.

(Greek, kuanos = dark blue, karpos = fruit)
l.c. (1827) 161; Kurz l.c. (1877) 184; Masamune l.c. 359; Backer & Bakhuizen f. l.c. 102; Swingle l.c.
(1967) 207; Stone l.c. (1978) 97, l.c. (1985) 5; Whitmore, Tantra & Sutisna l.c 306. Basionym:
Cookia cyanocarpa Blume l.c. (1825) 136. Type: Blume, s.n., Java, Mt. Salak (L). Synonyms:
Glycosmis longifolia Tanaka, Bull. Soc. Bot. Fr. 75 (1928) 709; G. cymosa (Kurz) Narayanaswami l.c.
26
var. platyphylla (Merr.) Stone

l.c. (1978) 98, l.c (1985) 6. Basionym: Glycosmis platyphylla Merr., Philip. J. Sc. 12 (1917) Bot. 273.
Type: Wenzel 1611, Philippines, Leyte (US). Synonym: Glycosmis clemensii Tanaka l.c. (1931) 4.
Shrub or small to medium-sized tree to 15 m tall and 30 cm diameter. Leaves 34-foliolate,

alternate, 1120 cm long; leaflets (1)39, elliptic or lanceolate, 430 x 110 cm, glabrous
or rusty-scurfy, sometimes only on midrib, below; base acute, margin usually entire or
sometimes finely toothed, apex caudate-acuminate; lateral veins 510(20) pairs, sometimes
strongly raised below, inarching near the margin; petiolules 24 mm long. Inflorescences
predominantly axillary corymbs, 112 cm long, open-branched, branches spreading, hairy,
with linear, leaf-like bracts. Flowers 45-merous; buds glabrous, c. 5 mm long; sepals
broad, rounded, c. 1.3 mm long, hairy, especially towards apex, margin rusty-ciliate; petals
oblong to oblong-obovate, 3.54 mm long, hairy, especially towards apex; stamens 10,
nearly equal to unequal, filaments broad, narrowed at apex, anthers ovate, glandularapiculate; disc elevated, minutely lobed or glandular; ovary 45-carpellate, ellipsoid,
constricted at base and in the middle, glabrous, not glandular, seated on a well-developed
gynophore, style short, stigma broad. Fruits oblong or ellipsoid, 11.5 cm long, purplish.
Seed 1 per fruit.
Distribution. India, Nepal, SW China, Burma, Indo-China, and throughout W Malesia
(Sumatra, Peninsular Malaysia, Java, Borneo and the Philippines). A variable species
comprising 10 varieties, one of which (var. platyphylla) occurs in Borneo, Philippines, and
the Talaud Islands. In Sabah and Sarawak (Lundu district), var. platyphylla occurs on flat
and undulating land in primary and secondary lowland forest, from near sea-level to 200 m;
also in Kalimantan.
3. Glycosmis lanceolata (Blume) Spreng.

(Latin, lanceolatus = spear-shaped; the leaves)
Cat. Hort. Bog. (1866) 208; Narayanaswami l.c. 61; Stone l.c. (1985) 10; Whitmore, Tantra & Sutisna
l.c. 306. Basionym: Sclerostylis lanceolata Blume l.c. (1825) 134. Type: Blume, s.n., Batavia (L).
Synonyms: Glycosmis simplicifolia Spreng. l.c. (1827) 162; G. montana Pierre, Fl. For. Cochinch. 4
(1893) 285; G. greenei var. simplex Stone l.c. (1978) 90.
Shrub or small to medium-sized tree to 18 m tall and 25 cm diameter. Branches with shiny
and cracked bark. Leaves 15-foliolate, alternate; leaflets, oblong-ovate or oblanceolate,
8.517 x 38 cm; base rounded, margin entire, apex shortly acuminate or emarginate; lateral
374
RUTACEAE (JONES)
veins 812 pairs; petiolules 34 mm long, 79 mm long in unifoliolate leaves.

Inflorescences short, axillary cymose-panicles, 13 cm long, branches short, rusty scurfyhairy. Flowers small, 5-merous; sepals l1.5 mm long, glabrous below; petals longer,
oblong-obovate, rusty scurfy-hairy below, margins hyaline; stamens 10, alternating long and
short, anthers oblong-cordate, connectives obscurely glandular; ovary (2)3-carpellate,
ellipsoid-ovoid, glabrous, distinctly papillate glandular, seated on a well-developed
gynophore, tapering into style, stigma 3-lobed. Fruits globose.
Distribution. Indo-China, Hainan, Sumatra, Java, Borneo, S Philippines, and the Lesser
Sunda Islands. Widespread in Sabah on ridges and slopes in lowland forest, occasionally on
limestone, from near sea-level to 240 m. In Sarawak, known from only one locality in
Kuching district (Stone 13739, Padawan, Bt. Pa'it). Also known from Kalimantan.
4. Glycosmis longisepala Stone

l.c (1985) 11; Whitmore, Tantra & Sutisna l.c. 306. Type: J. & M.S. Clemens 20001, Sarawak, Mt.
Pueh (holotype K).
Small tree. Leaves 57-foliolate, to 30 cm long; leaflets ovate-lanceolate or elliptic, 716 x

2.86 cm; base acute, margin entire, apex acuminate; lateral veins 68 pairs; petiolules 38
mm long. Inflorescences pseudoterminal cymose-panicles, elongate or pyramid-shaped, to
13 cm long. Flowers 5-merous; sepals narrowly triangular, c. 2.5 mm long, surfaces hairy,
margin ciliate; petals c. 3 mm long, hairy below; stamens 10, anthers c. 1 mm long,
glandular-apiculate, connective glandular dorsally; ovary (4)5-carpellate, to 2 mm long,
glabrous, glandular, seated on a well-developed gynophore. Fruits subellipsoid, c. 7.5 mm
long. Seeds 2 per fruit.
Distribution. Endemic to Sarawak; known only from the type locality, in forest, at 1000
1400 m.
5. Glycosmis macrantha Merr.
Fig. 5.
(Greek, makros = large, anthos = flower)

l.c. (1929) 114; Masamune l.c. 360; Swingle l.c. (1967) 207; Stone l.c. (1978) 92, l.c. (1985) 11;
Whitmore, Tantra & Sutisna l.c. 306. Type: Elmer 21456, British North Borneo, Tawau (holotype
UC; isotypes MO, US). Synonym: Glycosmis oliveri Stapf ex Ridl., Kew Bull. (1930) 80, Masamune
l.c. 360.
Small to medium-sized tree to 20 m tall and 20 cm diameter; bole to 12 m tall. Leaves 59foliolate, 1730 cm long, alternate; leaflets oblong to elliptic, 714 x 36 cm, glabrous,
chartaceous; base acute, apex acuminate, acumen 12 cm long; lateral veins 59 pairs,
strongly raised below; petiolules 47 mm long. Inflorescences terminal or axillary cymes.
Flowers large, 45-merous, whitish, on short stalks or nearly sessile; sepals orbicular, 2.53
mm long, sparsely hairy or glabrous; petals 5, oblong, glabrous, c. 7 x 3 mm; stamens 10,
equal, filaments thick, c. 4 mm long, anther connectives dorsally glandular; ovary 5carpellate, glabrous, seated on a well-developed gynophore, style cylindrical. Fruits ovoid,
c. 1 cm long.
375
Distribution. Endemic to Borneo. In Sabah, widespread and abundant in primary and

secondary forest, predominantly on hillsides and ridges, and often in disturbed habitats, to
1400 m. In Sarawak, uncommon and rarely collected. Also in Kalimantan.
6. Glycosmis sapindoides Lindl. in Wall. ex Oliver

(resembling the genus Sapindus)
J. Linn. Soc. Bot. 5, Suppl. 2 (1861) 38; Hooker f. l.c. (1875) 501; King l.c. 217; Ridley l.c. (1922)
351; Craib l.c. 225; Narayanaswami l.c. 55; Backer & Bakhuizen f. l.c. 102; Swingle l.c. (1967) 207;
Stone l.c. (1972) 381, l.c. (1985) 18; Whitmore, Tantra & Sutisna l.c. 307. Type: Wallich Cat. 6376,
Penang (lectotype K). Synonyms: Glycosmis cyanocarpa var. sapindoides (Lindl.) Kurz, J. Bot. 14
(1876) 34; G. elata Ridl., J. Fed. Mal. St. Mus. 10 (1920) 130; G. macrophylla (Blume) Miq., Fl. Ind.
Bat. 1, 2 (1859) 522.
var. sapindoides
Shrub or small tree to 10 m tall and 10 cm diameter. Young branches yellow, shiny. Leaves
pinnate, alternate; leaflets (3)59, oblanceolate, 1422 x 48 cm; base acute, margin entire,
apex acuminate; lateral veins 911 pairs, prominently raised below; petiolules to 3 mm
long. Inflorescences axillary cymose-panicles, 1.52(7.5) cm long; buds rusty hairy.
Flowers small, 5-merous, in clusters at tips of short branchlets; sepals broadly ovate, to 0.5
mm long, hairy below, margin ciliolate; petals oblong-elliptic, c. 2.5 mm long, hairy below;
stamens 10, alternating long and short, filaments slender, c. 1.5 mm long, glabrous, anthers
crescent-shaped, gland-tipped; ovary (2)3-carpellate, ellipsoid, densely rusty hairy, stigma
broad. Fruits ellipsoid.
Distribution. Mainland SE Asia and W Malesia, to Papua New Guinea and Australia. Of
the three known varieties, only the commonest (var. sapindoides) occurs in Sabah. This
variety occurs in the western and central parts of the range of the species, as far east as
Papua New Guinea. In Sabah, known only from Kota Kinabalu district (Pulau Sipanggar)
where it is found on ridge tops and hillsides in primary forest, to 200 m. Also in Kalimantan.
7. Glycosmis superba Stone

(Latin, superbus = splendid; the large, attractive leaves)
l.c. (1978) 95, l.c. (1985) 20; Anderson l.c. 308. Whitmore, Tantra & Sutisna l.c. 307. Type:
Anderson, Tan & Wright S. 26059, Sarawak, Ulu Sg. Sekaloh (SAR).
376
RUTACEAE (JONES)
377
378
RUTACEAE (JONES)
Shrub or small tree to 10 m tall and 10 cm diameter. Leaves large, opposite, 36-foliolate,
to 33 cm long; leaflets elliptic-oblanceolate, 2031 x 812 cm, leathery; base cuneate,
margin obscurely wavy to subentire, apex acuminate-caudate; lateral veins 1014 pairs;
petiolules 12 cm long. Inflorescences pseudoterminal, pyramid-shaped, densely cymosepaniculate, subglabrous, 3345 mm long. Flowers 5-merous; sepals broadly orbicularovate, c. 3 mm long, glabrous below, margin hyaline; petals at least 6 mm long, glabrous,
glandular; stamens 10, anther connectives 46-glandular, apex glandular-apiculate; disc 5lobulate; ovary 5-carpellate, cylindrical, glabrous, stigma 5-lobed, style glandular.
Distribution. Endemic to Sarawak (Kuching, Bintulu and Miri districts) and Brunei.
Ecology. Found on ridges and undulating ground in primary mixed dipterocarp forest and
secondary forest on sandy humult ultisols, from near sea-level to 100 m.
6. LUNASIA Blanco
(Tagalog, lunas; a native name for L. amara)
Fl. Filip. ed. 1 (1837) 783; Merrill l.c. (1929) 113; Masamune l.c. 360; Backer & Bakhuizen f. l.c. 99;
Hartley, J. Arn. Arb. 48 (1967) 460; Anderson l.c. 308; Perry l.c. 366.
Erect shrubs or small trees; dioecious. Branches unarmed. All vegetative and reproductive
parts with grey to reddish brown scale-like and/or star-shaped hairs. Leaves alternate,
simple; leaf blades pinnately veined, leathery, becoming stiff when older; petioles wingless,
swollen at apex. Inflorescences axillary panicles, flowers in small globose-clusters, 36 mm
in diameter. Flowers unisexual, 3-merous, fragrant; sepals 3, free, valvate; petals 3, free,
valvate, white to greenish or yellowish; stamens 3, opposite the sepals, rudimentary in
female flowers, anthers dorsifixed; ovary 3-carpellate, carpels fused at base, rudimentary in
male flowers, ovules 1 per locule; styles 3. Fruits of 13 dehiscent follicles, the undeveloped follicles persistent in fruit; pericarp dry at maturity; endocarp cartilaginous, discharged
when the follicle dehisces. Seeds 1 per locule; endosperm lacking; cotyledons oily.
Distribution. One species; E Java, Borneo, Philippines, Celebes, Papua New Guinea and
Australia (Queensland).
Ecology. Occurs in well-drained, primary and secondary rain forests, thickets, and garden
regrowth, from near sea-level to 900 m.
Lunasia amara Blanco
Fig. 6.
(Latin, amarus = bitter; referring to the alkaloid content)

l.c. 783; Merrill l.c. (1929) 113; Masamune l.c. 360; Backer & Bakhuizen f. l.c. 99; Hartley l.c. (1967)
464; Anderson l.c. 308; Perry l.c. 366. Type: Escritor BS 20776, Philippines, Luzon Island (neotype
A). Synonyms: Pilocarpus amara (Blanco) Blanco l.c. (1845) 540; Lunasia reticulata Elmer, Leafl.
Philip. Bot. 4 (1912) 1511; L. gigantifolia Merr., Philip. J. Sc. 21 (1922) Bot. 519.
379
var. amara
Shrub or small tree to 15 m tall (usually smaller), 30 cm diameter, sparsely branched.
Leaves crowded toward the branchlet tips; blades oblanceolate to obovate or elliptic, 660
x (up to) 18 cm; base cuneate to narrowly rounded or cordate, margin subentire to sinuate
or coarsely dentate, apex rounded to acuminate; lateral veins 935 pairs, strongly raised
below; petioles 1.515 cm long. Male inflorescences to 28 cm long and 8 cm wide; sepals
ovate, minute; petals obovate-acuminate, c. 1 mm long; staminal filaments glabrous. Female
inflorescences to 25 cm long and 2 cm wide; sepals broady ovate, 11.5 mm long; petals
ovate-acuminate, c. 2 mm long; staminodes 3; styles fused at base, stigmas flattened and
spreading over the tops of the carpels. Fruits obovate follicles, 615 x 510 mm, flattened
laterally, ribbed on the sides, with a beak to 5 mm long, densely hairy. Seeds nearly
obovoid; testa dark brown to reddish, papery.
Distribution. One variety (var. amara) with the same range of distribution as that of the
genus and species, occurs in Sabah and Sarawak. The other known variety, babuyanica
(Merr.) Hartley, is endemic to the Babuyan Islands in the Philippines. In Sabah, var. amara
is widespread and found in a variety of habitats, including offshore islands, ridges and
slopes in primary forest, and forest on ultramafic soils, from near sea-level to 900 m. In
Sarawak, frequent on limestone cliffs and slopes at low elevations (Miri and Baram
districts).
Uses. Bark, leaves, and seeds have been used medicinally to treat skin diseases, swollen
limbs, snake bite, and stomach ailments (Philippines, Indonesia). A number of alkaloids are
reported from the plant. Lunasin and lunacrin, two alkaloids extracted from the bark, were
shown to have negative effects on the voluntary and smooth muscles, blood vessels, and
hearts of laboratory animals (Wirth, J. Am. Pharm. Assoc. 20 (1931) 1254), causing death
due to the simultaneous cessation of the respiratory and circulatory systems.
7. MACLURODENDRON T.G. Hartley

(Floyd A. McClure, 18971970, American botanist and plant explorer)
Gard. Bull. Sing. 35 (1982) 1.
Small to medium-sized trees; dioecious. Branches unarmed. Young branchlets hairy,

trichomes brownish to rust-coloured. Leaves opposite, unifoliolate; blade pinnately veined,
margin entire; petioles wingless, swollen at the apex, articulated with the blades. Inflorescences axillary, paniculate or racemose. Flowers unisexual, 4-merous, round in bud; sepals
4, triangular, fused at base, valvate; petals 4, ovate, free, imbricate, white or greenish or
yellowish; stamens 8, unequal, alternately long and short, filaments glabrous, nearly linear,
curved inward, anther ovoid to ellipsoid, dorsifixed, pollen lacking in female flowers; disc
irregularly 8-lobed; ovary 4-carpellate, shallowly 4-lobed, glabrous, rudimentary in male
flowers, 2 ovules per locule, style straight, stigma capitate, 4-lobed. Fruit a 4-loculate
drupe, glabrous; exocarp somewhat leathery, glandular; mesocarp spongy, thin when dry;
endocarp parchment-like, shiny. Seeds ovoid to kidney-shaped, dark brown to black, shiny,
12 per locule; outer testa usually spongy, inner testa bony; endosperm fleshy; embryo
straight or bent.
380
RUTACEAE (JONES)
Distribution. 6 species; Thailand, Sumatra, Peninsular Malaysia, Borneo, the Philippines,

Vietnam, and Hainan Island. One species (M. porteri) commonly occur in Sabah and
Sarawak, while M. parviflorum is endemic to Sarawak and M. pubescens to Sabah.
Ecology. Mainly in well-drained primary rain forests and occasionally in secondary forests
and heath forests, from near sea-level to 1500 m.
Taxonomy. Several species were previously described under Acronychia (e.g., A. porteri).
The two genera are similar in a number of characters including the opposite leaves, 4merous flowers, 2-ovulate locules, and drupaceous fruits. However, Maclurodendron differs
in its exclusively unifoliolate leaves (sometimes trifoliolate in Acronychia), unisexual
flowers, imbricate petals, glabrous staminal filaments, and glabrous ovaries and fruits, never
with septicidal fissures.
Key to Maclurodendron species

1.
Fully developed leaf-blades hairy below, especially on the midrib and lateral veins
............................................................................................................3. M. pubescens
Fully developed leaf-blades glabrous or nearly so.........................................................2
2.
Inflorescences 13 cm long. Petals c. 1.5 mm long. Seed surface irregularly roughened

.........................................................................................................1. M. parviflorum
Inflorescences 215 cm long. Petals 22.5 mm long. Seed surface minutely reticulate
.................................................................................................................2. M. porteri
1. Maclurodendron parviflorum T.G. Hartley

(Latin, parvus = small, flos = flower)
l.c. (1982) 14. Type: Anderson S. 25426, Sarawak, Kuching district (holotype L; isotype SAR).
Small tree to 5 m tall. Branchlets glabrous or nearly so. Leaves elliptic or obovate to
oblanceolate, 916 x 39 cm, thinly leathery, drying pale green, glabrous or nearly so; base
narrow, apex acuminate or occasionally rounded; lateral veins 711 pairs; intercostal veins
and reticulations faint; petioles 12 cm long. Inflorescences 13 cm long; peduncle axis and
branches sparsely hairy to glabrous, pedicels sparsely hairy, 13 mm long. Flowers 11.5
mm wide in bud; sepals nearly 1 mm long, sparsely hairy; petals c. 1.5 mm long, glabrous.
Fruits c. 7 mm in diameter, nearly round to ovoid, shallowly 4-lobed, acute at the apex.
Seeds c. 5 mm long, surface irregularly roughened; outer testa not spongy.
Distribution. Endemic to Sarawak, where it is known from only four collections in Kuching
district.
Ecology. In primary heath (kerangas) forest and secondary forest on podzolised soils, at low
elevations.
381
2. Maclurodendron porteri (Hook. f.) T.G. Hartley
Fig. 7.
(G. Porter, first Curator of Waterfall Garden, Penang)

l.c. (1982) 8. Basionym: Acronychia porteri Hook. f. (1875) 498; Anderson l.c. 307. Type: Maingay
Kew Distr. No. 280, Penang (lectotype K; isolectotypes BM, GH, L). Synonyms: Jambolifera porteri
(Hook. f.) Kuntze, Rev. Gen. Pl. 1 (1891) 102; Melicope unifoliolata Merr., Philip. J. Sc. 13 (1918)
Bot. 74, l.c. (1921) 314, Masamune l.c. 360.
Small to medium-sized tree to 25 m tall and 40 cm diameter; bole to 12 m tall. Branchlets

glabrous or nearly so. Leaves obovate to oblanceolate or elliptic, 524 x 210 cm, thinly
leathery, drying pale brown to dark brown, glabrous or nearly so; base narrow or acute,
apex abruptly acuminate or occasionally rounded or acute; lateral veins 611 pairs;
intercostal veins and reticulations faint; petioles 0.75 cm long. Inflorescences 215 cm
long; peduncle axis and branches sparsely hairy to glabrous, pedicels nearly glabrous, 17
mm long. Flowers 1.52 mm wide in bud; sepals 0.51 mm long, glabrous to hairy; petals
22.5 mm long, glabrous to hairy. Fruits 611 mm in diameter, nearly round to ovoid, often
shallowly 4-lobed, often tipped. Seeds 48 mm long, surface minutely reticulate, outer testa
spongy.
Vernacular name. Sarawakrawang (Malay).
Distribution. Burma, Peninsular Thailand, Sumatra, Peninsular Malaysia, Singapore, Borneo,
and the Philippines. Widespread in Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. Locally on hillsides and ridges in primary mixed dipterocarp forest and occasionally
in secondary forest and heath (kerangas) forest (in Sarawak), on low nutrient organic soils,
from near sea-level to 1400 m.
Uses. The timber, while not durable, has been used for building purposes (Peninsular
Malaysia).
3. Maclurodendron pubescens T. G. Hartley

(Latin, pubescens = hairy; the leaves and floral parts)
l.c. (1982) 11. Type: Patrick SAN 26359, Sabah, Sandakan district (holotype L; isotype K).
Small to medium-sized tree to 30 m tall and 35 cm diameter; bole to 18 m tall. Branchlets

hairy, becoming glabrous. Leaves obovate to oblanceolate or elliptic, 1123 x 511 cm,
thinly leathery, drying pale greenish brown to brown, pubescent below especially on the
midrib and veins, glabrous above; base narrow, apex acuminate to rounded; lateral veins 8
11 pairs; intercostal veins and reticulations faint; petioles 1.55 cm long. Inflorescences 3
11 cm long; peduncle axis and branches sparsely to densely hairy; pedicels hairy, 35 mm
long. Flowers 1.52 mm wide in bud; sepals c. 7 mm long, hairy; petals c. 2 mm long, hairy
below, glabrous to sparsely hairy above. Fruits 1012 mm in diameter, nearly round to
ovoid, shallowly 4-lobed, acute at the apex. Seeds 57 mm long, surface minutely
reticulate; outer testa spongy.
382
RUTACEAE (JONES)
383
Distribution. Endemic to Sabah, where it is found in Sandakan (especially common in

Leila and Sepilok-Kabili FR), Keningau, and Lamag districts.
Ecology. On ridges and hillsides in primary forest, from near sea-level to 140 m.
8. MELICOPE J.R. Forst. & G. Forst.

(Greek, meli = honey, kope = a cutting;
the emarginate lobes of the nectar-secreting disc)
serang (Malay)
Char. Gen. Pl. (1776) 55; Hooker f. l.c. (1875) 491; Kurz l.c. (1877) 181; King l.c. 212; Anderson l.c.
308; Hartley, Gard. Bull. Sing. 34 (1981) 91, Sandakania 4 (1994) 47; Hartley & Stone, Taxon 38
(1989) 199.
Shrubs or small to tall trees; dioecious in unisexual plants. Branches unarmed. Leaves
opposite or whorled, trifoliolate and/or unifoliolate, petioles wingless; leaflets leathery or
thinly so, margin mostly entire, articulated at the base. Inflorescences cymes or densely
flowered panicles or solitary, axillary or on branchlets below leaves, rarely terminal or on
stems. Flowers small, bisexual or unisexual, 4-merous; sepals 4, fused at base; petals 4,
free, valvate or imbricate, white or cream to greenish or yellowish, rarely pink; stamens 4 or
8, rarely 48, free, rudimentary in female flowers; disc cushion- to ring-like or cupular;
ovary 4-carpellate, carpels fused completely or only at base, rudimentary in male flowers,
ovules (1)2 per locule, styles united, rarely stylar branches divergent, stigma small to
capitate or peltate, often lobed, or 4-branched. Fruits of 14 basally fused follicles, erect or
spreading at maturity, grading to a 4-locular capsule; endocarp cartilaginous, remaining
attached in the dehisced follicle, fused to or separate from epicarp. Seeds 12 per locule, not
expelled from mature fruit at dehiscence; testa thick, hard, covered by a shiny, black
epidermis; endosperm abundant; embryo straight or nearly so; cotyledons flat.
Distribution. About 230 species ranging from Madagascar to India, S China, throughout
Malesia, Polynesia, the Hawaiian Islands, Australia and New Zealand. 14 species in Sabah
and Sarawak.
Taxonomy. In his revision of the genus Tetradium Lour., Hartley (l.c. 1981) redefined
Melicope and suggested transferring to it many of the species placed in the genus Euodia J.
R. Forst. & G. Forst. As characterised by Hartley, Melicope and Euodia are largely
differentiated on the basis of their seeds (smooth, shiny and remaining attached in the
dehisced follicle in Melicope, and dull, rough and discharged from the follicle in Euodia),
floral characteristics (bisexual or unisexual flowers with 4 or 8 stamens in Melicope, and
bisexual flowers with 4 stamens in Euodia), and geographical ranges (Euodia has a
narrower distribution, occurring in New Guinea, NE Australia, and east to Samoa and
Hawaii). As a consequence of this new generic delimination, there are no representatives of
the genus Euodia in Sabah or Sarawak.
384
RUTACEAE (JONES)
Key to Melicope species

1.
Stamens 8. Seeds attached to dehiscent fruit by a partially detached strip of pericarp

and/or raphe. Leaves trifoliolate and/or unifoliolate.........................................................2
Stamens 4. Seeds attached to dehiscent fruit by a funiculus. Leaves mostly trifoliolate,
or (in M. subunifoliolata) sometimes uniformly unifoliolate.......................................4
2.
Sepals shorter than 1 mm. Follicles 25 mm long. Leaves mostly trifoliolate..............

..........................................................................................................14. M. triphylla
Sepals 12 mm long. Follicles 1013 mm long. Leaves trifoliolate and/or unifoliolate.......3
3.
Terminal bud glabrous. Petiolules 12 mm long.......................................9. M. jugosa

Terminal bud hairy. Petiolules 315 mm long...........................................12. M. sororia
4.
Flowers bisexual; stigma not lobed or wavy. Inflorescences axillary and/or on

branchlets below leaves..............................................................................................5
Flowers usually unisexual; stigma lobed or wavy. Inflorescences axillary...................6
5.
Leaflet-margin entire. Pedicels 48 mm long. Petals densely hairy above. Follicles 46

mm long..........................................................................................2. M. bonwickii
Leaflets-margin entire or (in cultivated forms) lobed or wavy. Pedicels less than 3 mm
long. Petals glabrous. Follicles 23 mm long.......................................5. M. denhamii
6.
Trichomes mostly simple...........................................................................................7

Trichomes mostly in bundles or star-shaped.............................................................13
7.
Leaflet-base cordate to rounded; petiolules lacking...............................10. M. latifolia

Leaflet-base rounded or acute to narrowly tapered; petiolules lacking or up to 20 mm
long........................................................................................................................................8
8.
Petals 1.31.8 mm long. Leaflets 1.55.5 cm wide.......................................................9

Petals 1.53 mm long. Leaflets 2.521 cm wide.............................................................10
9.
Leaflets glabrous or midrib sparsely hairy below. Follicles 79 mm long; epicarp

subfleshy, becoming glabrous............................................................3. M. clemensiae
Leaflets glabrous above; midrib hairy to velvety below. Follicles 1010.5 mm long;
epicarp dry, hairy.....................................................................13. M. subunifoliolata
10. Petals silky-hairy above. Raphe of seeds contorted..................................4. M. confusa

Petals glabrous or sparsely hairy to hairy above, especially on lower half. Raphe of seeds
not contorted...........................................................................................................11
11. Terminal leaflets obovate to broadly obovate, 7.516.5 cm long; apex abruptly
acuminate; lateral veins 915 pairs. Follicles 34 mm long........................6. M. glabra
Terminal leaflets elliptic to obovate, 3.546 cm long; apex usually acuminate; lateral
veins 824 pairs. Follicles 312 mm long.................................................................12
385
12. Leaflets glabrous or nearly so, to 23 cm long; lateral veins raised above. Follicles 6
12 mm long..................................................................................11. M. lunu-ankenda
Leaflets nearly glabrous to hairy below, to 46 cm long; lateral veins usually depressed
above. Follicles 35 mm long...............................................................1. M. accedens
13. Leaflets nearly glabrous to hairy below. Staminal filaments sometimes hairy at base.
Epicarp subfleshy...................................................................................7. M. hookeri
Leaflets densely whitish-hairy below. Staminal filaments glabrous. Epicarp dry
................................................................................................................8. M. incana
1. Melicope accedens (Blume) T. G. Hartley
Fig. 8.
(Latin, accedere = approaching, probably referring to its resemblance to Euodia macrophylla)

l.c. (1994) 67. Basionym: Euodia accedens Blume l.c. (1825) 246. Type: Blume, s.n., Java (lectotype
L). Synonyms: Euodia macrophylla Blume l.c. (1825) 246; Zanthoxylum macrophyllum (Blume) Miq.
l.c. (1859) 670; Ampacus macrophylla (Blume) Kuntze l.c. 98; Evodia accedens (Blume) Miq. l.c.
(1859) 671; Ampacus accedens (Blume) Kuntze l.c. 98; E. nervosa Koord. & Valeton, Med. Lands Pl.
Tuin 17 (1896) 208, Merrill l.c. (1921) 314, l.c. (1929) 113, Masamune l.c. 358, Anderson l.c. 308.
Shrub or tree to 40 m tall. Young branchlets and terminal buds glabrous to hairy or velvety.
Trichomes mostly simple. Leaves opposite, trifoliolate (occasionally unifoliolate), petioles
2.524 cm long; leaflets elliptic to obovate, 9.546 x 421 cm; base rounded to narrowly
tapered and often asymmetric, apex usually acuminate; midrib sometimes velvety above and
below; lateral veins 1026 pairs, usually depressed above; petiolules lacking or to 15 mm
long. Inflorescences axillary, 324 cm long, glabrous to hairy or velvety; main branches
spreading, pedicels 0.53.5 mm long. Flowers unisexual or sometimes bisexual; sepals
ovate to triangular, 0.52 mm long, usually hairy below; petals ovate to elliptic, 1.52.8 mm
long, glabrous to hairy below, sparsely hairy to hairy, sometimes only on lower half, or
glabrous, above; stamens 4, filaments hairy at base; disc and ovary glabrous to densely
hairy, style hairy at base, stigma weakly 4-lobed. Fruits of round to ellipsoid or obovoid
follicles, 35 mm long; epicarp subfleshy, glabrous to hairy; endocarp glabrous. Seeds
round to ovoid or ellipsoid, sometimes compressed, 2.55 mm long; funiculus 0.31.5 mm
long.
Distribution. From Andaman Islands east to Vietnam and south to Peninsular Malaysia,
Borneo and Java. In Sabah and Sarawak, common in primary and secondary forest, swamp
forest, heath forest, peat swamp forest, and along forest edges and in open places, from near
sea-level to 1950 m; also reported from Kalimantan.
Taxonomy. Hartley (l.c. 1994) recognises two variants of this species, both of which occur
in Sabah and Sarawak. One variant, found throughout the range of the species, possesses
sepals which are nearly glabrous to sparsely hairy below, petals which are glabrous or
nearly so below, follicles 34 mm long with a glabrous or sparsely hairy epicarp, and
leaflets with 1324 pairs of lateral veins. The second variant, occurring in Peninsular
Malaysia, Sumatra and Borneo, differs in having hairy sepals, glabrous to hairy petals,
follicles 45 mm long with a hairy epicarp, and leaflets with 1016 pairs of lateral veins.
386
RUTACEAE (JONES)
387
2. Melicope bonwickii (F. Muell.) T.G. Hartley

(J. Bonwick, student of Australian geography)
l.c. (1994) 56. Basionym: Euodia bonwickii F. Muell., Fragm. 5 (1865) 56. Type: Dallachy, s.n.,
Australia, Queensland, Rockingham Bay (holotype MEL). Synonyms: Euodia speciosa Rchb. f. &
Zoll. ex Teijsm. & Binnen., Nat. Tijd. Ned. Ind. 29 (1867) 255; E. villamilii Merr., Philip. J. Sc. 9
(1914) Bot. 296.
Tree to 40 m tall. Young branchlets glabrous or rarely sparsely hairy; terminal bud
appressed hairy. Trichomes simple. Leaves opposite, trifoliolate, petioles 214.5 cm long;
leaflets elliptic to obovate, 1030 x 515 cm, glabrous or sparsely hairy on veins below;
base acute or narrowly tapered and sometimes asymmetric, apex acuminate; lateral veins
1424 pairs, sometimes raised above, sometimes with axillary cavities below; petiolules
lacking or to 4 mm long. Inflorescences on branchlets below leaves, rarely axillary,
glabrous to sparsely hairy, 3.510 cm long; pedicels 48 mm long. Flowers bisexual; sepals
nearly round, to 2 mm long, glabrous to sparsely hairy below; petals pink or rarely white,
ovate to elliptic or elliptic-oblong, 45.5 mm long, densely hairy above; stamens 4,
filaments glabrous; disc and ovary hairy to densely hairy, style glabrous, stigma capitate.
Fruits of nearly round to obovoid follicles, 46 mm long; epicarp dry, sparsely hairy to
almost glabrous; endocarp glabrous. Seeds nearly round to ellipsoid or nearly hemispherical,
34.5 mm long; funiculus 13 mm long.
Distribution. Java, Borneo, Philippines, and east to Papua New Guinea and Australia. In
Sabah, in primary and secondary forest, from near sea-level to 750 m; also occurs in
Kalimantan.
Uses. The bark is reported to be used in the treatment of leech bites (Indonesia).
3. Melicope clemensiae T.G. Hartley

(Mary S. Clemens, 18731968, plant collector of the Malesian region)
l.c. (1994) 64. Type: Aban Gibot SAN 60767, Sabah, Mt. Kinabalu Park (holotype L; isotype SAN).
Shrub or tree to 15 m tall. Young branchlets and terminal buds glabrous, or nearly so, to
velvety. Trichomes mostly simple. Leaves opposite, trifoliolate (rarely unifoliolate); petioles
1.58 cm long; leaflets elliptic to obovate or oblanceolate, 516 x 25.5 cm, glabrous or
midrib sparsely hairy below; base acute to narrowly tapered and often asymmetric, apex
acuminate; lateral veins 1118 pairs, raised above; petiolules 1.512 mm long. Inflorescences
axillary, laxly flowered, 2.516 cm long, glabrous to somewhat velvety; pedicels 25 mm
long. Flowers unisexual; sepals round to ovate-triangular, less than 1 mm long, hairy or
glabrous below; petals ovate to elliptic, c. 1.5 mm long, glabrous; stamens 4, filaments
glabrous; disc and ovary nearly glabrous to hairy, style glabrous, stigma weakly 4-lobed.
Fruits of round to obovoid follicles, 79 mm long; epicarp subfleshy, becoming glabrous;
endocarp glabrous. Seeds round, ovoid or ellipsoid, slightly compressed, 57 mm long;
funiculus 1.52 mm long.
Distribution. Endemic to Sabah and Sarawak; in primary and secondary forest, on ridges
and slopes, sometimes on limestone, from 650 to 1800 m.
388
RUTACEAE (JONES)
4. Melicope confusa (Merr.) Liu

(Latin, confusio = disorder; referring to this taxon's long confusion with Euodia glabra)
Ill. Native Introd. Lign. Pl. Taiwan 2 (1962) 876; Hartley l.c. (1994) 59. Basionym: Euodia confusa
Merr., Philip. J. Sc. 20 (1922) Bot. 391. Type: Ramos BS 15055, Philippines, Luzon, Laguna
Province (A).
Tree to 30 m tall. Young branchlets glabrous to sparsely hairy; terminal buds hairy to
velvety. Trichomes simple. Leaves opposite, trifoliolate; petioles 2.518.5 cm long; leaflets
elliptic to obovate or ovate, 1235 x 58 cm, glabrous to hairy below; base acute to
narrowly tapered, rarely asymmetric, apex acuminate; lateral veins 1124 pairs, raised to
depressed above; petiolules 315 mm long. Inflorescences axillary, 630 cm long, peduncle
glabrous to hairy, pedicels 13 mm long. Flowers unisexual; sepals nearly round or ovate,
to 1 mm long, hairy below; petals elliptic, 2.53 mm long, silky hairy above; stamens 4,
filaments sparsely hairy towards base or glabrous; disc and ovary hairy, style hairy at least
basally, stigma weakly 4-lobed. Fruits of round to ellipsoid follicles, 4.56 mm long, fused
basally and erect or diverging; epicarp subfleshy, sparsely hairy or glabrate; endocarp
glabrous. Seeds round to ovoid or ellipsoid, 3.54 mm long; funiculus 11.5 mm long,
raphe contorted.
Distribution. Borneo, Philippines, Celebes and Moluccas. In Sabah, found mainly in primary
and secondary forest, from near sea-level to 90 m; no record from Sarawak.
Uses. The bark is used medicinally for treating enlargement of the spleen (Philippines); a
decoction of the root or leafy shoot with liquor is ingested to treat hives (Taiwan).
5. Melicope denhamii (Seemen) T.G. Hartley

(H. M. Denham, the 19th century British sea captain)
l.c. (1994) 57. Basionym: Picrasma denhamii Seemen, Fl. Vit. (1865) 33. Type: McGillivray 46, New
Hebrides, Aneitum (holotype BM). Synonyms: Euodia tenuistyla Stapf, Trans. Linn. Soc. Bot. 4
(1894) 137, Merrill l.c. (1921) 314, Masamune l.c. 359; E. ridleyi Hochr., Icon Bog. 2 (1905) 151; E.
schullei var. ridleyi (Hochr.) Lauterb., Bot. Jahrb. 55 (1918) 230; E. suaveolens var. ridleyi (Hochr.)
Bakhuizen f., Blumea 6 (1950) 365.
Shrub or tree to 25 m tall. Young branchlets and terminal buds glabrous to densely hairy or
velvety. Trichomes simple, star-shaped or in bundles. Leaves opposite, trifoliolate
(occasionally unifoliolate); petioles 116 cm long; leaflets elliptic, 1019 cm long, sparsely
hairy to hairy on midrib and veins below; base cuneate to narrowly tapered, apex acuminate;
lateral veins 1522 pairs, raised or depressed above; petiolules lacking or up to 1 cm long.
Inflorescences axillary and/or on branchlets below leaves, 27 cm long, hairy, pedicels
0.22.3 mm long. Flowers bisexual; sepals ovate to triangular, to 0.8 mm long, glabrous to
hairy below; petals ovate-elliptic or elliptic, 1.22.2 mm long, glabrous; stamens 4, filaments
glabrous; disc glabrous; ovary hairy, style glabrous to sparsely hairy, stigma capitate. Fruits
of nearly round follicles, 23 mm long; epicarp dryish, glabrous to sparsely hairy;
endocarp glabrous. Seeds nearly round or hemispherical, 1.52.5 mm long; funiculus to 1.5
mm long.
389
Distribution. Borneo to S Philippines and throughout E Malesia to Fiji. In Sabah, in

primary and secondary forest and swamps, from near sea-level to 950 m; also known from
Kalimantan.
Taxonomy. Several cultigens of this species exist and are cultivated as ornamentals in
Borneo and elsewhere. They are all characterised by linear to elliptic or ovate leaflets
usually with lobed or wavy margins. The synonym Euodia ridleyi refers to one of these
cultigens.
6. Melicope glabra (Blume) T.G. Hartley

(Latin, glaber = hairless)
l.c. (1994) 60. Basionym: Fagara glabra Blume l.c. (1823) 40. Type: Blume, s.n., Java (lectotype
US). Synonyms: Euodia glabra (Blume) Blume l.c. (1825) 245, Anderson l.c. 308; Ampacus glabra
(Blume) Kuntze l.c. 98; E. kingii Engl. in Engler & Prantl, Pfl. Fam. 3, 4 (1896) 121; E. krukovii
Merr., Pap. Mich. Ac. Sc. 24 (1938) 75.
Tree to 40 m tall. Young branchlets glabrous to velvety; terminal buds sparsely hairy to
velvety. Trichomes simple. Leaves opposite, trifoliolate (occasionally unifoliolate); petioles
glabrous, 28.5 cm long; leaflets elliptic to broadly obovate, 7.516.5 x 412 cm, hairy on
midrib and veins below or glabrous; base acute to narrowly tapered, asymmetric in lateral
leaflets, apex abruptly acuminate or rounded or emarginate; lateral veins 915 pairs,
sometimes depressed above; petiolules 120 mm long. Inflorescences axillary, 730 cm
long, glabrous to velvety; pedicels 0.62 mm long. Flowers unisexual; sepals round to
ovate, less than 1 mm long, glabrous to hairy below; petals elliptic, 22.5 mm long, sparsely
hairy on lower half above or glabrous; stamens 4, filaments glabrous; disc and ovary
glabrous to hairy, style glabrous to hairy, stigma weakly 4-lobed. Fruits of round or broadly
elliptic follicles, 34 mm long; epicarp subfleshy, glabrous to sparsely hairy; endocarp
glabrous. Seeds round to broadly ellipsoid, 2.53 mm long; funiculus c. 1 mm long.
Distribution. Sumatra, Peninsular Malaysia, Singapore, Java, and Borneo. Found in
primary forest, mostly below 500 m but ascending as high as 1200 m. Reported once from
Sabah (Keith, N. Bor. For. Rec. 2 (1952) 373), but its occurrence there needs to be
confirmed.
7. Melicope hookeri T.G. Hartley

(Joseph D. Hooker, 18171911, eminent British botanist)
l.c. (1994) 70. Synonyms: Euodia robusta Hook. f. l.c. (1875) 488, Masamune l.c. 359, non E. robusta
A. Smith, J. Arn. Arb. 32 (1951) 253; Ampacus robusta (Hook.f.) Kuntze l.c. 98. Type: Maingay Kew
Distr. No. 278, Singapore (GH, lectotype).
Tree to 25 m tall. Young branchlets glabrous to velvety; terminal buds hairy to velvety.
Trichomes mostly in bundles or star-shaped. Leaves opposite, trifoliolate (occasionally
unifoliolate); petioles 413 cm long; leaflets ovate to elliptic or obovate, 7.518 x 49.5 cm,
390
RUTACEAE (JONES)
nearly glabrous to hairy below; base obtuse to narrowly tapered, apex acuminate, sometimes
abruptly so; lateral veins 1522 pairs, sometimes depressed above; petiolules 115 mm
long. Inflorescences axillary, 1026 cm long; pedicels 0.31.5 mm long. Flowers unisexual,
rarely bisexual; sepals round to ovate-triangular, less than 1 mm long, glabrous to hairy
below; petals ovate to elliptic, 1.52 mm long, glabrous; stamens 4, filaments sometimes
hairy at base; disc and ovary hairy, style hairy at base, stigma weakly 4-lobed. Fruits of
round to obovoid follicles, 3.54 mm long; epicarp subfleshy, glabrous or sparsely hairy;
endocarp glabrous. Seeds round to ellipsoid, 2.53.5 mm long; funiculus 11.2 mm long.
Distribution. Sumatra, Peninsular Malaysia, and Borneo. In Sabah and Sarawak, scattered
in primary and secondary forests and forest margins, to 1600 m. Also known from Brunei
and Kalimantan.
8. Melicope incana T. G. Hartley

(Latin, incanus = greyish-white; the leaflets)
l.c. (1994) 71. Type: Lobb, s.n., Borneo (holotype K). Synonyms: Euodia alba Hook. f., Trans.
Linn. Soc. 23 (1862) 166, Merrill l.c. (1921) 313, Masamune l.c. 358, Anderson l.c. 308, non E. alba
Laurterb., Nov. Guinea 14 (1924) 141; Ampacus alba (Hook.f.) Kuntze l.c. 98.
Tree to 35 m tall. Young branchlets and terminal buds hairy to velvety. Trichomes mostly
star-shaped. Leaves opposite, trifoliolate; petioles 320 cm long; leaflets elliptic to ovate or
obovate, 925 x 516 cm, densely whitish-hairy below, glabrous to hairy, or midrib densely
hairy above; base rounded, apex abruptly acuminate; lateral veins 2025 pairs, depressed
above; petiolules lacking or to 3 mm long. Inflorescences axillary, 923 cm long; pedicels
0.31 mm long. Flowers unisexual, rarely bisexual; sepals round to ovate-triangular, less
than 1 mm long, hairy below; petals elliptic, 1.72 mm long, sparsely hairy along median
line below or glabrous; stamens 4, filaments glabrous; disc and ovary hairy, style hairy,
stigma weakly 4-lobed. Fruits of ellipsoid to obovoid follicles, 34 mm long; epicarp dry,
very sparsely hairy; endocarp glabrous. Seeds round, c. 2 mm long; funiculus c. 1 mm long.
Distribution. Sumatra, Borneo and N Celebes. In Sabah and Sarawak, in primary and
secondary forest, alluvial forest and freshwater swamp forest, from near sea-level to 800 m.
Also known from Brunei and Kalimantan.
9. Melicope jugosa T.G. Hartley

(Latin, jugosus = mountainous; the habitat)
l.c. (1994) 51. Type: J. & M.S. Clemens 51184, British North Borneo, Mt. Kinabalu, Gurulau Spur
(holotype A; isotypes K, NY, UC).
Shrub or small tree to 3 m tall. Young branchlets somewhat corky, glabrous like the terminal
bud. Trichomes, if present, simple. Leaves opposite, trifoliolate and/or unifoliolate, glabrous;
petioles 25 cm long; leaflets of trifoliolate leaves elliptic-obovate or elliptic or obovate,
4.57 x 23 cm; base acute to narrowly tapered, margin toothed toward apex, apex rounded
or abruptly acuminate, lateral veins 711 pairs, sometimes raised above; petiolules 12 mm
long; leaflets of unifoliolate leaves elliptic or rarely elliptic-obovate, to 13.5 x 6 cm, base
rounded or rarely acute, lateral veins 913 pairs; petiolules lacking, or similar to that of
trifoliolate leaves. Inflorescences axillary, glabrous to sparsely hairy, 12 cm long; pedicels to
3.5 mm long. Flowers unisexual; sepals nearly round or ovate, to 2 mm long, sometimes
ciliolate; petals elliptic, to 4 mm long; stamens 8; stigma capitate, weakly 4-lobed. Fruits
glabrous, follicles ellipsoid, c. 10 mm long; epicarp dry. Seeds ellipsoid, c. 8 mm long.
391
Distribution. Endemic to Sabah, in forest from 2250 to 2400 m.
10. Melicope latifolia (DC.) T.G. Hartley

(Latin, latus = broad, folium = leaf)
l.c. (1994) 72. Basionym: Euodia latifolia DC. l.c. 724, Merrill l.c. (1921) 314, Masamune l.c. 358,
Anderson l.c. 308. Type: Doleschall 335, Moluccas, Ambon (neotype W).
Shrub or tree to 30 m tall. Young branchlets and terminal buds glabrous to softly hairy.
Trichomes simple. Leaves opposite, trifoliolate (occasionally unifoliolate); petioles 429
cm long; leaflets elliptic to ovate or obovate, 837 x 319 cm, glabrous to hairy above and
below; base cordate to rounded, sometimes asymmetric, apex acuminate; lateral veins 14
31 pairs, raised to slightly depressed above; petiolules lacking. Inflorescences axillary,
often densely flowered, 524 cm long, glabrous to softly hairy; pedicels lacking or to 3 mm
long. Flowers unisexual, sometimes bisexual; sepals ovate to triangular or rounded, to 1.5
mm long, softly hairy below; petals ovate to elliptic, 24 mm long, glabrous to hairy below;
stamens 4, filaments glabrous; disc more or less glabrous; ovary glabrous to hairy, style
glabrous, stigma 4-lobed or wavy. Fruits of elliptic follicles, 3.54.5 mm long, sometimes
fused up to full length; epicarp dry, glabrous to sparsely hairy; endocarp glabrous. Seeds
round to ellipsoid or hemispherical, 23 mm long; funiculus 0.51.5 mm long.
Distribution. Peninsular Malaysia, Java, Borneo, Phillipines, Papua New Guinea and east to
Samoa. In Sabah, in primary and secondary forest and open places, from near sea-level to
600 m. No record from Sarawak.
Uses. The leaves have been used for treating fever and cramps (Peninsular Malaysia,
Indonesia). Resin collected from the trunk has been used as a varnish and adhesive
(Indonesia).
11. Melicope lunu-ankenda (Gaertn.) T.G. Hartley

(the Sri Lankan name for this species)
l.c. (1994) 61. Basionym: Fagara lunu-ankenda Gaertn., Fruct. Sem. Pl. 1 (1788) 334. Type: Koenig,
s.n., Ceylon (holotype L). Synonyms: Euodia aromatica Blume l.c. (1825) 246; Zanthoxylon
aromaticum (Blume) Miq. l.c. (1859) 670; Ampacus aromatica (Blume) Kuntze l.c. 98; Zanthoxylum
roxburghianum Cham., Linnaea 5 (1830); E. roxburghiana (Cham.) Benth., Fl. Hongk. (1861) 59;
Ampacus roxburghiana (Cham.) Kuntze l.c. 98; E. lunu-ankenda (Gaertn.) Merr., Philip. J. Sc. 7
(1912) Bot. 378, Merrill l.c. (1921) 314, Anderson l.c. 308; E. arborea Elmer, Leafl. Philip. Bot. 8
(1915) 2806; E. malayana Ridl. l.c. (1922) 342, Masamune l.c. 358, Anderson l.c. 308; E. punctata
Merr., J. Str. Br. R. As. Soc. 86 (1922) 315, l.c. (1929) 113, Masamune l.c. 358, Anderson l.c. 308; E.
triphylla var. pubescens Ridl. l.c. (1930) 77, Masamune l.c. 359; E. concinna Ridl. l.c (1930) 78,
Masamune l.c. 358; E. obtusifolia Ridl. l.c. (1930) 78, Masamune l.c. 358.
392
RUTACEAE (JONES)
Shrub or tree to 30 m tall. Young branchlets glabrous to velvety; terminal buds sparsely
hairy to velvety. Trichomes mostly simple. Leaves opposite, trifoliolate (occasionally
unifoliolate); petioles 1.515 cm long; leaflets elliptic to obovate, 3.523 x 2.57 cm,
glabrous or nearly so; base rounded to narrowly tapered and often asymmetric, apex acute
to acuminate, rarely rounded or emarginate; lateral veins 817 pairs, sometimes raised
above; petiolules 0.515 mm long. Inflorescences axillary, 232 cm long with main
branches ascending, glabrous to velvety; pedicels 0.33 mm long. Flowers unisexual,
rarely bisexual; sepals round to ovate-triangular, 0.51.2 mm long, glabrous to hairy below;
petals ovate to elliptic, 1.53 mm long, glabrous to sparsely hairy below, hairy above on
lower half or glabrous; stamens 4, filaments hairy at base or glabrous; disc glabrous to
densely hairy or velvety; ovary nearly glabrous to densely hairy, style glabrous to hairy,
stigma weakly 4-lobed. Fruits of ellipsoid to obovoid follicles, 610 mm long (912 mm
long in montane plants); epicarp subfleshy, glabrous; endocarp glabrous. Seeds round to
ovoid or ellipsoid, sometimes compressed, 36 mm long; funiculus 0.53 mm long.
Distribution. Himalayas, Sri Lanka, Java, Borneo, SW Philippines, and Celebes. In Sabah
and Sarawak, common in primary and secondary well-drained or swampy forest and
montane shrubbery, from near sea-level to 2200 m. Also known from Brunei and
Kalimantan.
Taxonomy. Hartley (l.c. 1994) recognises lowland and montane variants of this species.
Plants from montane habitats have larger follicles and more leathery leaflets, usually with
rounded to abruptly tipped apices. Because the two intergrade, they are treated as a single
taxon.
Uses. The roots are reportedly used to treat colds and rheumatism (Taiwan), and the leaves
and flowers for menstrual disorders and fever (Peninsular Malaysia). Its timber is weak but
used in construction. The leaves are eaten as a condiment and have been used to flavour
food.
12. Melicope sororia T.G. Hartley

(Latin, sororius = sisterly; referring to its close relationship to Melicope jugosa)
l.c. (1994) 53. Type: J. & M.S. Clemens 29477, British North Borneo, Mt. Kinabalu, Tenompok
(holotype NY; isotypes A, B, BO, L, UC).
Shrub or small tree to 5 m tall. Young branchlets glabrous, rather corky; terminal bud
hairy. Trichomes mostly simple. Leaves opposite, trifoliolate and/or unifoliolate, rarely
bifoliolate, glabrous, petioles 110 cm long; leaflets of trifoliolate leaves elliptic to obovate,
6.514 x 36 cm, base acute to narrowly tapered and somewhat asymmetric, margin entire
or less commonly toothed toward apex, apex acuminate or rarely rounded, lateral veins 711
pairs, sometimes raised above, petiolules 315 mm long; leaflets of unifoliolate leaves to 16
x 8 cm, petiolule to 1 cm long, otherwise similar to leaflets of trifoliolate leaves.
Inflorescences axillary, glabrous, 2.510 cm long; pedicels to 3.5 mm long, sparsely hairy.
Flowers unisexual; sepals nearly round or ovate or elliptic, to 1.5 mm long, sometimes
ciliolate; petals elliptic, to 2.5 mm long; stamens 8; stigma capitate, weakly 4-lobed. Fruits
glabrous, follicles ellipsoid, 1013 mm long; epicarp dry. Seeds ellipsoid, 910 mm long.
393
Distribution. Endemic to Sabah, occurring in forest from 1500 to 2400 m.
13. Melicope subunifoliolata (Stapf) T.G.Hartley

(Latin, sub = nearly, unus = one, folium = leaf; the almost unifoliolate leaves)
l.c. (1994) 66. Basionym: Euodia subunifoliolata Stapf l.c. 138, Merrill l.c. (1921) 314, Masamune
l.c. 359. Type: Haviland 1193, British North Borneo, Mt. Kinabalu (holotype K).
Shrub or tree to 15 m tall. Young branchlets hairy to velvety; terminal buds velvety.
Trichomes mostly simple. Leaves opposite, trifoliolate and/or unifoliolate, glabrous above,
hairy to velvety on midrib (sparsely hairy on veins) below; petioles 19 cm long; leaflets of
trifoliolate leaves elliptic to obovate, 515.5 x 1.55.5 cm, base acute to narrowly tapered,
apex acuminate, lateral veins 1020 pairs, sometimes raised above, petiolules 0.510 mm
long; leaflets of unifoliolate leaves elliptic-obovate, to 8.5 x 3.5 cm, base acute, lateral veins
1114 pairs, otherwise similar to leaflets of trifoliolate leaves. Inflorescences axillary, laxly
flowered, 313 cm long, hairy to velvety; pedicels 12.5 mm long. Flowers unisexual;
sepals ovate to triangular, 0.71.5 mm long, hairy below; petals elliptic to ovate, 1.31.8
mm long, glabrous; stamens 4, filaments glabrous; ovary and top of disc hairy, style hairy at
base, stigma weakly 4-lobed. Fruits of ellipsoid follicles, 1010.5 mm long; epicarp dry,
hairy; endocarp glabrous. Seeds round to ovoid, sometimes compressed, 56.3 mm long;
funiculus 0.51.5 mm long.
Distribution. Endemic to Sabah, occurring in primary montane forest, often on ridges and
slopes, from 1200 to 2600 m, and sometimes descending to 180 m.
14. Melicope triphylla (Lam.) Merr.

(Greek, treis = three, phyllon = leaf; having trifoliolate leaves)
l.c. (1912) 375; Anderson l.c. 309; Hartley l.c. (1994) 54. Basionym: Fagara triphylla Lam., Encycl.
2 (1788) 375. Type: Sonnerat, s.n., Philippines (holotype P). Synonyms: Euodia triphylla (Lam.) DC.
l.c. 724; Zanthoxylum triphyllum (Lam.) G. Don, Gen. Hist. 1 (1831) 804; Ampacus triphylla (Lam.)
Kuntze l.c. 98.
Shrub or tree to 15 m tall. Young branchlets glabrous to hairy, sometimes glaucous; terminal
bud glabrous to velvety hairy. Trichomes, if present, simple. Leaves opposite, mostly
trifoliolate, petioles 311 cm long; leaflets elliptic to oblanceolate, 825 x 2.56 cm,
glabrous; base acute to narrowly tapered and sometimes asymmetric, apex acuminate;
lateral veins 719(25) pairs, sometimes raised above; petiolules 215 mm long. Inflorescences axillary and/or on branchlets below leaves, glabrous to hairy, 111 cm long, pedicels
lacking or to 4 mm long. Flowers unisexual; sepals ovate-triangular, to 0.8 mm long, apex
sometimes sparsely hairy; petals lanceolate to ovate or elliptic, to 3.5 mm long; stamens 8,
filaments sparsely hairy or glabrous; disc hairy or glabrous; ovary glabrous, stigma
capitate, 4-lobed. Fruits glabrous, follicles ellipsoid or nearly round, 2.55 mm long;
epicarp subfleshy. Seeds nearly round to ellipsoid, 2.44.5 mm long.
394
RUTACEAE (JONES)
Distribution. Taiwan and Ryukyu Islands, south to Borneo, and east to Papua New Guinea.
In Sabah and Sarawak, in primary and secondary forest, from near sea-level to 1300 m.
Also in Kalimantan.
9. MEROPE Roem.
(Merope of Greek mythology, one of the Heliades)
Syn. Hesper. 1 (1846) 44; Backer & Bakhuizen f. l.c. 106; Swingle l.c. (1967) 258; Stone l.c. (1972)
382, l.c. (1978) 115.
Shrubs or small trees. Branches armed with axillary, solitary or paired spines. Leaves
alternate, unifoliolate, leathery; petioles short, stout, wingless, articulated with the blade.
Inflorescences 1(2)-flowered, rarely few-flowered, axillary. Flowers bisexual, 45-merous;
calyx cup-like, 5-lobed; petals 5(6), free, imbricate in bud; stamens 10, free, equal; ovary
ovoid, 3(4)-carpellate, on a tall gynophore, ovules 24 per locule, style short, thick, stigma
flat. Fruit an ovoid, angular berry, with 34-flattened sides; pericarp thick, glandular, strongly
aromatic when crushed, filled with sticky clear fluid. Seeds few to many per fruit, oblong,
flattened, overlapping.
Distribution. One species, ranging from Burma to Indo-China and throughout Malesia.
Found in Sabah and Sarawak.
Ecology. A widely scattered, highly specialised genus which thrives in the saline soil of
tidal forests and mangrove swamps, often on the banks of streams. The tides aid in the
dispersal of the buoyant fruits.
Merope angulata (Willd.) Swingle
Fig. 9.
(Latin, angulatus = angled; the fruit)

J. Wash. Ac. Sc. 5 (1915) 420, l.c. (1967) 258; Backer & Bakhuizen f. l.c. 106; Stone l.c. (1972) 382,
l.c. (1978) 115. Basionym: Citrus angulata Willd., Sp. Pl. ed. 4, 3 (1800) 1426. Type: Limonellus
angulosus Rumph., Herb. Amboin. 2 (1741) 110. Synonyms: Sclerostylis spinosa Blume l.c. (1825)
134; Paramignya longispina Hook. f. l.c. (1875) 511; P. angulata (Willd.) Kurz, J. As. Soc. Beng. 44
(1875) 135.
Erect shrub or small tree to 3 m tall and 10 cm diameter, often with multiple stems arising
from a root crown. Branches sparse, with paired, rarely solitary, stout spines, to 5 cm long
on juvenile stems. Leaves oblong-ovate to obovate, 4.516 x 26 cm, thick, glabrous; base
rounded, margin subentire to faintly notched, apex acute or shortly acuminate; midrib
prominent below; lateral veins and reticulations inconspicuous; petioles 57 mm long.
Inflorescences 1(2)-flowered, glabrous; pedicels short. Flowers small, fragrant, buds
ovoid; sepals triangular, c. 1 mm long, glabrous; petals oblong-lanceolate, 79 mm long,
glabrous, white; filaments glabrous, white, anthers linear-oblong, yellow; ovary slender,
glabrous. Fruits ovoid to ellipsoid, 34.5 cm long, 34-sided in cross-section, apex bluntly
acuminate, green when ripe. Seeds 23 cm long, somewhat reniform, tapering to a blunt
point; testa rough; cotyledons green.
395
Vernacular names. Sabah and Sarawaklimau buaya, limau laut (Malay).

Distribution. Throughout the range of the genus. In Sabah and Sarawak, occurs in rather
small but dense populations in inland areas of tidal swamps and mangrove forest, often on
the edges of streams and on riverbanks, and usually in association with Nipa palm (Nypa
fruticans).
Uses. The fruits are used for the treatment of stomach disorders and colic (Peninsular
Malaysia).
10. MERRILLIA Swingle

(E. D. Merrill, 18761954, American botanist)
l.c. (1918) 337, l.c. (1967) 240; Ridley l.c. (1922) 353; Craib l.c. 231; Burkill l.c. (1966) 1481; Stone
l.c. (1972) 382; Corner l.c. 667; Stone & Jones l.c. (1988) 268.
Trees; crown rather bushy; trunk and main branches covered with pale fissured bark;
branches unarmed. Leaves alternate, pinnate; leaflets 513, lower ones small, higher ones
increasingly larger, terminal leaflet largest, thinly leathery, articulated at the base; rachis
flattened, narrowly winged. Inflorescences axillary, 12-flowered. Flowers bisexual, large,
slightly irregular, 5-merous; calyx cup-like, sepals 5, small; petals 5, free, long, imbricate;
stamens 10, free, unequal; ovary 5(6)-carpellate, bottle-shaped, on a well-developed
gynophore, ovules 810 per locule, style long, slender, stigma capitate. Fruit a subglobose
to oblong or ellipsoid berry; pericarp thick, resinous, 5-chambered, chambers with cartilaginous walls and filled with clear mucilage. Seeds numerous, flattened, densely scaly;
cotyledons plano-convex.
Distribution. One species; S Thailand, Sumatra, Peninsular Malaysia, and Borneo (Sabah).
Ecology. Scattered as solitary trees on stream banks and slopes of hills and ridges, in moist
primary and secondary forest, to 400 m.
Taxonomy. This anomalous genus contains a number of metabolites in common with
Murraya sect. Murraya (Kong et al., Biochem. Syst. Ecol. 16 (1988) 47). One of these
metabolites, yuehchukene, is of particular interest because of its anti-implantation effect in
rats. Swingle (l.c. 1967) suggested a possible origin of the genus from a Murraya-like
ancestor. Morphologically, there are a number of similarities between Merrillia and
396
RUTACEAE (JONES)
397
Murraya sect. Murraya. Both have yellowish stem- and root-bark, and some species of sect.
Murraya possess a winged leaf-rachis and densely villous seeds, characteristics found also in
Merrillia caloxylon. The two differ markedly, however, in the size and shape of their
flowers and fruits.
Merrillia caloxylon (Ridl.) Swingle
Fig. 10.
(Greek, kalos = beautiful, xylon = wood)

l.c. (1918) 338, l.c. (1967) 241; Ridley l.c. (1922) 354; Craib l.c. 231; Burkill l.c. (1966) 1481; Stone
l.c. (1972) 383; Corner l.c. 667; Stone & Jones l.c. 68; Ng l.c. 495. Basionym: Murraya caloxylon
Ridl., J. Str. Br. R. As. Soc. 50 (1908) 113. Type: Wray & Robinson 5548, Upper Perak, Kenering
(holotype BM; isotypes SING, US).
Small to medium-sized tree to 20 m tall and 25 cm diameter; bole short. Leaves to 20 cm

long; leaflets usually 79, alternate to subopposite, lowest leaflets stipule-like, larger
laterals elliptic, 7.59 x 4 cm; base triangular and slightly asymmetric, margin wavy to
slightly toothed, apex acuminate; petiolules nearly lacking. Inflorescences usually 1flowered. Flowers pendulous, trumpet-shaped; sepals triangular or ovate, to 2.5 mm long;
petals oblanceolate, gradually tapering to a narrow base, 2.55.5 cm long, greenish-white;
ovary and style hairy. Fruits c. 11 x 9 cm or larger; pericarp c. 13 mm thick, leathery,
warty, greenish becoming yellow. Seeds 810 per chamber, lens-shaped, c. 13 x 3 mm,
scales membranous, flattened, slightly fimbriate.
Distribution. Throughout the range of the genus. In Sabah, collected only once from the
bank of a stream in secondary forest at 20 m (Junaidi Payne, s.n., Sandakan, Ulu Gum
Gum; SAN).
Uses. In Peninsular Malaysia, the durable, handsome wood, which is yellow with dark
brown streaks, has been used to make walking sticks, smoking pipes, parang handles and
sheaths, and other small objects.
11. MICROMELUM Blume

(Greek, mikros = small, melon = apple; the shape of the fruits)
l.c. (1825) 137; Hooker f. l.c. (1875) 501; Kurz l.c. (1875) 136; King l.c. 218; Ridley l.c. (1922) 351;
Merrill l.c. (1929) 114; Craib l.c. 227; Masamune l.c. 360; Backer & Bakhuizen. f. l.c. 103; Burkill l.c.
(1966) 1492; Swingle l.c. (1967) 197; Stone l.c. (1972) 383; Anderson l.c. 309; Perry l.c. 366; Corner
l.c. 667; Whitmore, Tantra & Sutisna l.c. 307.
Shrubs or trees; branches unarmed. Leaves alternate, pinnate, rachis wingless; leaflets (1)
715(23), laterals alternate, subchartaceous to thinly leathery, usually obliquely asymmetric and articulated at the base. Inflorescences terminal, flat-topped paniculate corymbs,
often very large, flowers numerous. Flowers bisexual, 5-merous; calyx cup-like,
398
RUTACEAE (JONES)
399
sepals triangular, small; petals oblong-linear, free, valvate; stamens 10, alternately long and
short, filaments linear, glabrous, anthers ovate, sub-basifixed; disc annular; ovary 26carpellate, glabrous to densely hairy, radial walls often twisted, one oil-gland over each
locule, ovules 2 per locule, style slender, constricted at the base, detaching from ovary,
stigma flattened to subglobose. Fruit a subglobose or oblong berry, dryish; pericarp thin,
glandular, glabrate to hirsute. Seeds ellipsoid; cotyledons thin, folded.
Distribution. About 10 species; from W Pakistan, India and Sri Lanka to S China, and
south through Indo-China and Malesia to Australia, New Caledonia and S Pacific. One
species, M. minutum, in Sabah and Sarawak.
Micromelum minutum (G. Forst.) Wight & Arn.
Fig. 11.
(Latin, minutus = small; the flowers)

l.c. (1834) 93; Craib l.c. 227; Masamune l.c. 360; Backer & Bakhuizen f. l.c. 103; Burkill l.c. (1966)
1493; Swingle l.c. (1967) 203; Stone l.c. (1972) 383; Anderson l.c. 309; Perry l.c. 366; Corner l.c.
668; Whitmore, Tantra & Sutisna l.c. 307; Ng l.c. 496. Basionym: Limonia minuta G. Forst., Prodr.
(1786) 33. Type: Forster, s.n., Friendly Islands (BM). Synonyms: Micromelum pubescens Blume l.c.
(1825) 138, Merrill l.c. (1929) 114, Masamune l.c. 361, Anderson l.c. 309; M. glabrescens Benth. in
Hooker, Lond. J. Bot. 2 (1843) 212.
var. minutum
Shrub or small to medium-sized tree to 20 m tall and 15 cm diameter. Young branchlets
densely short-hairy, the trichomes greyish. Leaves to 30 cm long or more; leaflets 915,
ovate-lanceolate, 312 x 1.56 cm, thinly leathery, sometimes drying almost black, subglabrous to shortly hairy on veins below; base obtuse and asymmetric, margin entire or
wavy to shallowly dentate-crenate, apex attenuate-acuminate; lateral veins 58 pairs;
petiolules to 5 mm long. Inflorescences 1320 cm long, hairy; bracts short deltoid-linear;
pedicels to 5 mm long. Flowers small; sepals shortly hairy, sometimes glabrate; petals to 5
8 mm long, spreading, hairy below, pale green or yellowish-white; staminal filaments to 9
mm long, narrowed at apex, white; ovary cylindric, 5-carpellate, c. 1.5 mm long, hairy
becoming glabrous, style glabrous or sparingly hairy, stigma flattened to subcapitate. Fruits
ellipsoid-oblong, to 1 cm long, glabrate, yellow or red when ripe. Seeds with green, wrinkled
cotyledons.
Distribution. Sri Lanka, Indo-China, Peninsular Malaysia, Borneo, Philippines, Papua New
Guinea, NE Australia and the Pacific. A variable species with four varieties, the most
widespread of which, var. minutum, occurs in Sabah and Sarawak. The var. minutum is
scattered in Sabah and Sarawak, in primary and secondary forest on low undulating ground,
disturbed open sites, and forest margins, and frequently on limestone outcrops and sandy
coasts of the mainland and islands, from near sea-level to 600 m.
Uses. The roots and leaves are used in treating scabies, intermittant fever, headache, and
menstrual disorders (Indo-China, Peninsular Malaysia, Philippines). Its light, durable timber
is used for construction purposes (Peninsular Malaysia).
400
RUTACEAE (JONES)
401
12. MONANTHOCITRUS Tanaka

(Greek, monos = single, anthos = flower)
J. Arn. Arb. 9 (1928) 138; Swingle l.c. (1967) 252; Stone l.c. (1985) 216; Stone & Jones l.c. (1988)
268.
Erect shrubs to small trees. Branches slender, armed, spines axillary and paired or single,
unequal in length, rarely unarmed. Leaves alternate, simple; petioles short, wingless, not
articulated with the blade. Inflorescences axillary, usually solitary. Flowers small, bisexual, 5merous, buds ovoid or ellipsoid; calyx cup-like, slightly lobed, sepals 5, imbricate; petals 5,
ovate, imbricate; stamens 10, equal, free, anthers oblong-linear; disc annular; ovary 35carpellate, globose to oblong, glabrous to sparsely hairy, ovules 210 per locule, style stout,
stigma 35-lobed. Fruit a globose to ovoid or pyriform berry; pericarp somewhat leathery,
thin, glandular. Seeds 610 per fruit, tightly packed in the locules and embedded in
mucilage or scant pulp, flattened to concave or plano-convex, with or without thin
membranous margin; testa sometimes spotted.
Distribution. 4 species; Borneo, Irian Jaya, and Papua New Guinea. One species, M.
oblanceolata, in Sabah.
Taxonomy. Proposed by Tanaka in 1928, the genus remained monotypic until 1985, when
Stone described a new species, M. bispinosa, and transferred one other from Wenzelia
Merr., a related genus occurring in the Philippines, Irian Jaya, Papua New Guinea, and Fiji.
The two genera were originally distinguished chiefly on the character of the seeds (margin
laciniate-membranous and testa spotted in Monanthocitrus, and margin thin-membranous
and testa not spotted in Wenzelia). These differences do not hold up between the genera as
here understood, and their distinctness now remains somewhat problematical. The size of
the flowers (generally smaller in Monanthocitrus) and nature of the spines (generally paired
and subequal in Monanthocitrus) may be more significant characters, but conclusive
analysis will depend upon the availability of new material.
Monanthocitrus oblanceolata Stone & Jones
Fig. 12.
(Latin, ob = reversed, lancea = spear; the leaf shape)

l.c. 268. Type: Abu Bakar SAN 36203, Sabah, Sandakan, Labuk Road (holotype SAN; isotypes K, L).
Shrub or small tree to 5 m tall. Bark smooth, dark brown. Spines paired, 540 mm long or
more. Leafy stems zig-zag. Leaves oblong-oblanceolate or narrowly subelliptic, 420 x
1.57 cm, thinly leathery; base subcuneate or obtuse or subcordate, margin entire, apex
acuminate-caudate, tip to 26 mm long; midrib below prominent, sparsely hairy; lateral veins
clearly visible, 810 pairs; reticulations obscure; petioles 2.57 mm long. Inflorescences
1(2)-flowered; pedicels to 5 mm long. Flowers with calyx c. 2 mm wide, sepals ovatetriangular, c. 1 mm long, margin minutely ciliate; petals subelliptic, c. 5 mm long, glabrous,
whitish; staminal filaments c. 1.5 mm long, anthers with a minutely glandular-apiculate apex;
ovary narrowly bottle-shaped, 34-carpellate, sparsely hairy, style cylindrical,
402
403
404
RUTACEAE (JONES)
stigma minutely lobed. Fruits obovoid or pyriform, to 5 x 2.8 cm, beaked; pericarp yelloworange, smooth, on pedicels to 1 cm long. Seeds 810 per fruit, embedded in whitish, sweettasting flesh; testa thin; cotyledons plano-convex, thick, green, obscurely glandular.
Distribution. Endemic to Sabah and rather uncommon in Kota Belud, Beluran and
Sandakan districts, south to Lahad Datu and Tawau.
Ecology. Found as solitary trees or small populations in the understorey of lowland forests,
usually occurring on slopes and ridges of low hills, and sometimes bordering streams on flat
land, to 600 m.
Taxonomy. The species superficially resembles Merope angulata. However, the more
conspicuous leaf venation, smaller flowers, obovoid fruits, and different habitat conditions
readily distinguish this from Merope angulata which is associated with mangrove swamps.
The two species are very similar in size and habit.
13. MURRAYA Koenig ex L.

(J. A. Murray, 17401791, student of Linnaeus)
Mant. 2 (1771) 554, 563; Hooker f. l.c. (1875) 502; Kurz l.c. (1877) 190; King l.c. 219; Ridley l.c.
(1922) 353; Craib l.c. 230; Browne l.c. 315; Backer & Bakhuizen f. l.c. 103; Burkill l.c. (1966) 1531;
Swingle l.c. (1967) 231; Stone l.c. (1972) 384; Perry l.c. 367; Corner l.c. 668.
Shrubs or trees, deciduous or evergreen. Branches unarmed. Bark usually smooth, often
pale. Leaves alternate, pinnate or rarely unifoliolate, rachis wingless; leaflets usually 39
(to as many as 25), articulated at the base. Inflorescences axillary or terminal panicles,
cymes or corymbs. Flowers bisexual, medium-sized to large (over 1 cm long), 5-merous,
buds cylindrical or oblong; sepals ovate or lanceolate, fused at base; petals linear to
oblanceolate, imbricate; stamens 10, free, nearly equal or alternately long and short,
filaments slightly flattened, glabrous, anthers ovate to elliptic, nearly basifixed; disc annular to
cylindric; ovary 25-carpellate, ovoid to ellipsoid, glabrous or sometimes finely hairy, on a
gynophore, ovules (1)2 per locule, style slender, not persistent, stigma capitate. Fruit an
ovoid to round berry, pulp mucilaginous, peel thin, glandular. Seeds 1 to several per fruit,
smooth or hairy; testa thin; cotyledons green, plano-convex.
Distribution. About 15 species; from India to S China, Indo-China and Taiwan, and
throughout Malesia, eastward to NE Australia and New Caledonia. One native species
(Murraya paniculata) occurs in Sabah. Two additional species, M. exotica and M. koenigii,
are occasionally cultivated in Sabah and Sarawak.
Taxonomy. A study by Kong et al. (Biochem. Syst. Ecol. 14 (1986) 491) on the distribution
of the alkaloids yuehchukene and girinimbine in the roots of eight species of Murraya has
revealed a division within the genus between species producing one or the other of these
alkaloids. This division is supported by morphological differences which have long been
recognised. However, the most recent classification of the genus (Swingle l.c. 1967) does
not adequately reflect this information. One group (section Murraya) is characterised by
405
species containing yuehchukene (no girinimbine) and possessing yellowish stems and roots,
larger petals (12 cm long), and red, ovoid to ellipsoid fruits. Plants in the second group
(section Bergera) contain girinimbine (no yuehchukene) and possess brown stems and roots,
smaller petals (47 mm long), and purplish-black, globose to ellipsoid fruits. Of the species
occurring in Sabah and Sarawak, M. paniculata and M. exotica represent section Murraya,
and M. koenigii represents section Bergera.
Key to Murraya species

1.
Leaf stalks hairy. Inflorescences many-flowered corymbs. Fruits blackish when ripe.
Cultivated......................................................................................................................
M. koenigii (L.) Spreng.
l.c. (1825) 315. Basionym: Bergera koenigii L. l.c. 563. Synonym: Murraya foetidissima
Teijsm. & Binn., Nat. Tijd. Ned. Ind. 27 (1864) 41; Chalcas koenigii (L.) Kurz l.c. (1875)
132.
India, Sri Lanka, Burma, Indo-China, Hainan and S China. Commonly cultivated in
tropical countries, including in Sabah and Sarawak, for its leaves (Curry leaves;
daun kari).
Leaf stalks glabrate. Inflorescences few-flowered panicles or cymes. Fruits red when
ripe...................................................................................................................................2
2.
Leaflets obovate, less than 4 cm long, apex somewhat obtuse. Inflorescences terminal.
Fruits elliptic to subglobose. Cultivated..........................................................................
M. exotica L.
l.c. (1771) 563. Synonym: Murraya paniculata var. exotica (L.) Huang, Act. Phytotax.
Sinica 8 (1959) 100.
Native to China and Taiwan (?). Commonly cultivated throughtout the tropics.
Leaflets ovate or elliptic to rhomboid, 37 cm long, apex acuminate. Inflorescences
axillary. Fruits ovoid with acuminoid apex. Wild species...........................M. paniculata
Murraya paniculata (L.) Jack
Fig. 13.
(Latin, paniculatus = tufted; the inflorescence)

Mal. Misc. 1 (1820) 31; Ridley l.c. (1922) 353; Craib l.c. 230; Browne l.c. 315; Backer & Bakhuizen f.
l.c. 103; Burkill l.c. (1966) 1531; Swingle l.c. (1967) 232; Stone l.c. (1972) 384; Perry l.c. 367;
Corner l.c. 669; Ng l.c. 496. Basionym: Chalcas paniculata L., Mant. 1 (1767) 68. Type: India
(LINN); based partly on Camunium Rumph., Herb. Amboin. 6. Synonyms: Limonia lucida G. Forst.
l.c. (1786) 33; Murraya sumatrana Roxb., Fl. Ind. ed. 2, 2 (1832) 375; M. odorata Blanco l.c. (1845)
256.
var. paniculata
Shrub or small to medium-sized tree to 20 m tall, 25 cm diameter. Bark thin, pale to whitish.
Young shoots, twigs, sepals, petals, and ovary glabrous to slightly hairy. Leaves pinnate, to
17 cm long; leaflets 37, rarely unifoliolate, ovate or ovate-elliptic to rhomboid, 37 x
23.5 cm, chartaceous or thinly leathery, glossy and darker above, glabrous; base cuneate to
406
RUTACEAE (JONES)
rounded, margin entire or faintly crenulate, apex acuminate; lateral veins 58 pairs;
petiolules 26 mm long. Inflorescences axillary panicles or cymes, few-flowered; peduncle
14 cm long; pedicels 29 mm long. Flowers large; sepals (4)5, narrowly deltoid, c. 1 mm
long, glandular; petals (4)5, elliptic to oblong-obovate, to 1521 mm long or more, white;
stamens alternately long and short, the longer to 12 mm long, filaments dilated below; ovary
2-carpellate, glandular, style columnar, nearly 1 cm long, stigma enlarged, bilobed. Fruit an
ovoid berry, c. 12 mm long, apex acuminoid; peel shiny red, gland-dotted, glabrous. Seeds
densely hairy; cotyledons thick, fleshy.
Vernacular name. Sabahkemuning (Malay).
Distribution. Throughout the range of the genus. Of the four known varieties, only var.
paniculata occurs in Sabah, where it is occasionally found in lowland and hill forests,
usually on rocky soils or limestone, from near sea-level to 600 m.
Taxonomy. M. exotica has usually been placed in synonymy with this species or as a
variety of it. In this treatment, M. paniculata is restricted to the wild form found locally,
with comparatively larger leaflets and flowers. M. exotica, locally found only in cultivation,
differs in the shape of its leaflets (obovate to subelliptic) and fruits (ellipsoid to subglobose),
in having terminal inflorescences, and in its more restricted natural geographical range
(coastal areas of Hong Kong and China).
Uses. The leaves, fruits and bark of both M. paniculata and M. exotica have been used for
treating venereal disease, intestinal worms, skin disorders, and dysentery, among others
(China, Philippines, Peninsular Malaysia, Indonesia). Their heavy, yellow wood has been
used to make walking sticks, kris handles, and various small objects (Peninsular Malaysia,
Thailand). A face powder has been made from its bark and roots (Burma), and the fragrant
flowers are used in cosmetics (Java). The leaves contain an unidentified, water-soluble toxin
that has been shown to kill eggs and nymphs of the Asian citrus blackfly, Aleurocanthus
woglumi Ashby, with topical exposure (Dowell, Pan-Pacific Entomol. 65 (1989) 163).
14. PLEIOSPERMIUM (Engl.) Swingle

(Greek, pleio = few, sperma = seed)
J. Wash. Ac. Sc. 6 (1916) 426, l.c. (1939) 258, l.c. (1967) 290; Merrill l.c. (1929) 115; Masamune l.c.
361; Backer & Bakhuizen f. l.c. 104; Anderson l.c. 309.
Shrubs or small trees. Branches armed with axillary, solitary or paired spines, or unarmed
or nearly so. Young twigs angular, becoming terete. Leaves alternate, 13-foliolate; petioles
winged or wingless; leaflets thinly leathery, glabrous, margin entire, articulated at the base.
Inflorescences axillary or pseudoterminal panicles or racemes, often few-flowered, hairy.
Flowers small, bisexual, 45-merous, fragrant, buds cylindric; sepals triangular to linear-
407
408
RUTACEAE (JONES)
lanceolate, glabrous to finely hairy; petals linear-oblong, glabrous or finely hairy below,
white to greenish, imbricate in bud; stamens 8 or 10, free, alternately long and short,
filaments glabrous, anthers oblong to linear-oblong; disc annular to shortly cup-like; ovary
45-carpellate, cylindric or ovate, glabrous or hairy, on a short gynophore or nearly sessile,
ovules 2 per locule, style slender or stout, stigma subcapitate. Fruit a globose to oblong
berry, 1.52.5 cm in diameter; pericarp rough-glandular; pulp-vesicles slender, to 1 cm
long; juice oily-resinous. Seeds ovoid, flattened, to 1 cm long; testa smooth or wrinkled;
monoembryonic; cotyledons plano-convex, thick, green.
Distribution. 6 species; S India, Sri Lanka, Vietnam, Sumatra, Java, and Borneo. Two
species, P. latialatum and P. longisepalum, are found in Sabah. The former species also
occurs in Sarawak.
Key to Pleiospermium species

Petioles broadly winged, 1020 mm wide. Sepals 46 mm long, triangular........1. P. latialatum
Petioles narrowly winged, 35 mm wide. Sepals 912 mm long, linear.........2. P.
longisepalum
1. Pleiospermium latialatum Swingle

(Latin, latus = broad, ala = wing; the petiole)
l.c. (1939) 261, l.c. (1967) 293; Masamune l.c. 361; Anderson l.c. 309. Type: Elmer 21542, British
North Borneo, Tawau (holotype AA; isotypes BO, K, L, P, PNH, US).
Shrub or small to medium-sized tree to 25 m tall, 30 cm diameter; bole to 15 m tall; trunk,

bole, and main branches armed with paired spines to 3 cm long, generally unarmed on upper
branches. Leaves unifoliolate, oblong-elliptic or lanceolate, 915 x 3.57 cm; base rounded
or cuneate, apex acute-caudate, acumen 46 mm long and emarginate; lateral veins 1013
pairs; petioles 1.52.5 cm long, broadly winged, obcordate, top 12 cm wide, narrower at
base and pulvinoid. Inflorescences axillary clusters or pseudoterminal panicles. Flowers
with sepals triangular, (2)46 x 24 mm, glabrous, persistent and spreading at right
angles below the fruit; petals glabrous; ovary hairy. Fruits globose, 1.52.5 cm in diameter;
pericarp greenish yellow, with numerous raised oil-glands; pulp-vesicles slender, conical,
34.5 mm long, tips acute. Seeds 23 per fruit.
Vernacular names. Sarawaklimo antu (preferred common name), limo to (Kayan), limo
bali (Kenyah), para bileh (Berawan).
Distribution. Endemic to Borneo. Widespread and common in Sabah and Sarawak,
occurring in the lowlands and hills and along streams and rivers in primary mixed
dipterocarp and secondary forests on fertile clay-rich loam soils, and (in Sarawak)
on limestone slopes and ridges, from near sea-level to 300 m.
409
2. Pleiospermium longisepalum Swingle
Fig. 14.

l.c. (1939) 259, l.c. (1967) 293. Type: Castro & Melegrito 1348, British North Borneo, Banguey Is.
(holotype NA; isotypes K, US).
Shrub or small tree to 15 m tall, 40 cm diameter; trunk sometimes fluted. Branches unarmed
or nearly so. Leaves unifoliolate, elliptic, 7.511.5 x 35.5 cm; base rounded or cuneate,
apex acuminate, acumen rounded or retuse; lateral veins 810 pairs; petioles 11.5 cm long,
narrowly winged (wing 35 mm wide at top), tapering to a wingless, pulvinoid base.
Inflorescences axillary clusters or pseudoterminal panicles. Flowers with sepals linear,
912 x 23 mm, chartaceous, apex rounded or emarginate, persistent and reflexed below
the fruit. Fruits globose, tipped by the short persistent style, 22.5 cm in diameter; pericarp
1.52 mm thick, pimply from numerous raised oil-glands, ripening pale brownish yellow,
pulp-vesicles 46 mm long, tapering to a blunt point. Seeds 12 per fruit, c. 1 cm long.
Vernacular names. Sabahlimau hutan, limau limau (Malay).
Distribution. Endemic to Sabah where it is not uncommon in the foothills and low
mountainous areas of Kota Marudu, Ranau, and Tambunan districts, and occasionally on
off-shore islands (Pulau Banggi = Banguey Is., Pulau Bohayan).
Ecology. Usually found on the slopes of hills and adjacent to streams in primary and
secondary inland forests, and on high ground near the sea, to 1300 m.
15. SEVERINIA Tenore

(Severinus, 6th Archbishop of Rome, 640 A.D.)
Ind. Sem. Hort. Neap. 3 (1840); Masamune l.c. 357; Swingle l.c. (1967) 283; Stone l.c. (1978) 116;
Perry l.c. 371.
Shrubs or small trees. Branches spreading, armed with stout axillary spines, or unarmed.
Leaves alternate, simple, often rigid, conspicuously parallel-veined; petioles short, wingless,
not articulated with the blade. Inflorescences densely flowered, axillary or terminal panicles
or racemes, or few-flowered clusters. Flowers bisexual, small, 35(7)-merous; calyx cuplike, sepals imbricate, glabrous or hairy; petals valvate, glabrous or nearly so; stamens 515,
free, filaments dilated or flattened, anthers small, disc cup-like, enclosing base of ovary;
ovary subglobose or oblong, 15-carpellate, on a short gynophore, ovules 1 per locule, style
short, stigma ovoid. Fruit a small berry, round, juicy or semi-dry; pericarp dotted with oilglands, smooth; pulp-vesicles irregular in size and shape, lacking stalks. Seeds small,
ovoid, thick; testa thin; monoembryonic.
Distribution. 6 species; from S China, Taiwan, and Indo-China, south to Java, N Borneo,
the Philippines, Moluccas, and Irian Jaya. Two species (S. disticha and S. paniculata) occur
in Sabah.
410
RUTACEAE (JONES)
Taxonomy. The genus has been confused with Atalantia Corr. which resembles in it leaf
characters (shape, size, and venation) and habit. All species of Severinia have at one time
been placed in Atalantia. Atalantia differs, however, in having leaf-blades articulated with
the petioles, larger flowers, filaments more or less fused, well-formed pulp-vesicles, and
small, orange-like fruits. Its 11 species are distributed from India to W Malesia and is
represented in Peninsular Malaysia by two species. The relationship of Severinia to
Atalantia needs to be examined more critically.
Key to Severinia species

Leaves ovate-lanceolate, apex acute-acuminate. Inflorescences axillary or terminal,
racemose or paniculate. Ovary 2-carpellate............................................................1. S. disticha
Leaves oblong-ovate, apex rounded. Inflorescence terminal, paniculate. Ovary 4-carpellate
...............................................................................................................................2. S. paniculata
1. Severinia disticha (Blanco) Swingle
Fig. 15.
(Latin, distichus = of two rows; the leaf arrangement).

J. Wash. Ac. Sc. 28 (1938) 533, l.c. (1967) 287; Masamune l.c. 357; Stone l.c. (1978) 116. Basionym:
Limonia disticha Blanco l.c. (1837) 356. Type: Merrill, Sp. Blancoanae No. 594, Philippines, Luzon
(L). Synonyms: Limonia corymbosa Blanco l.c. (1845) 251; Atalantia nitida Oliv. l.c. 24; A. disticha
(Blanco) Merr., Bull. Gov. Lab. Philip. 27 (1905) 28, Masamune l.c. 357.
Shrub or small tree to 10 m tall, 15 cm diameter. Branches unarmed, occasionally with

paired spine-like paraphylls. Leaves 38(11) x (1.5)2.54 cm or more, ovate or ovatelanceolate, wavy in dried specimens; base cuneate to obtuse, margin subentire, sometimes
faintly crenulate, apex acute or acuminate, acumen usually blunt, often emarginate; lateral
veins parallel, straight, numerous, 1520 pairs, prominent below; petioles 39 mm long,
flattened above, glabrous to hairy. Inflorescences many-flowered, axillary or (rarely)
terminal panicles or racemes, 39 cm long; pedicles 35 mm long. Flowers 5-merous;
sepals 5, broadly rounded, margins ciliate; petals 5, oblong, minutely tipped, to 7 mm long,
white; stamens 10, alternately long and short, filaments to 5 mm long, flattened, narrowed at
apex and base, anthers ovate, apiculate-glandular; ovary 2-carpellate, oblong, glabrous to
sparsely hairy, style cylindrical, stout, stigma slightly lobed. Fruits subglobose, to 1.5 cm
wide, blackish-purple when ripe; pulp greenish, juicy. Seeds 12 per fruit, greenish white.
Distribution. Borneo (Sabah), E Java, Philippines, Moluccas, Flores, and Irian Jaya. In
Sabah, not uncommon in the understorey of beach strand vegetation, and in primary and
secondary forest on hills and ridges near the coast, frequently on offshore islands, usually on
sandy or rocky soil, rarely on limestone, from near sea-level to 90 m.
Uses. The ripe, juicy fruits are rather sweet and eaten fresh (Sabah, Indonesia). The roots are
said to be used medicinally, and its fruits made into glue (Philippines). The timber has been
used in construction (Indonesia, Philippines).
411
2. Severinia paniculata (Warb.) Swingle

(Latin, paniculatus = tufted; the inflorescence)
l.c. (1938) 533, l.c. (1967) 288; Stone l.c. (1978) 116. Basionym: Atalantia paniculata Warb., Bot.
Jahrb. 13 (1891) 340. Type: Warburg 20132, Moluccas, Ceram Laut (B). Synonyms: Atalantia
maritima Merr., l.c. (1914) 293; A. disticha (Blanco) Merr. var. paniculata (Warb.) Tanaka l.c. (1928)
141.
Shrub or small tree. Branches unarmed. Leaves oblong-ovate, 912 x 35 cm, or larger;
base gradually tapered, margin obscurely crenulate, apex rounded or emarginate; lateral
veins numerous, parallel, more or less prominent below; petioles c. 8 mm long, terete, hairy.
Inflorescences terminal panicles, 46 cm long or more, hairy; pedicels 24 mm long.
Flowers 5-merous; sepals 5, c. 1 mm long, tips blunt, margins ciliate; petals 5, 47 mm
long, lanceolate, tips rounded, subglabrous; stamens 10, alternately long and short, filaments
45 mm long, glabrous, anthers broadly cordate; ovary 4-carpellate, style deciduous. Fruits
globose, to 12 mm in diameter. Seeds 24 per fruit.
Distribution. Borneo (Sabah), S Philippines, Moluccas, and Sumbawa (the Lesser Sunda
Is.). In Sabah, less widespread than the related species S. disticha, and found in many of the
same habitats, particularly beach forest and offshore islands, at sea level.
Taxonomy. This species is very closely related to S. disticha and Tanaka (l.c. 1928)
considered it to be a variety. Many of their morphological characters appear to overlap.
Swingle (l.c. 1967) distinguished it from S. disticha based upon its larger leaves, smaller,
paniculate inflorescences, and ovaries with four carpels instead of two. A re-evaluation of
the taxonomic status of this species (and most other Severinia) would be useful.
16. TETRACTOMIA Hook. f.

(Greek, tetra = four, tome = a separation; the four carpels which split in fruit)
l.c. (1875) 490; King l.c. 211; Merrill l.c. (1921) 314; Ridley l.c. (1922) 345; Craib l.c. 215;
Masamune l.c. 361; Burkill l.c. (1966) 2182; Stone l.c. (1972) 385; Hartley, J. Arn. Arb. 60 (1979)
127; Anderson l.c. 309; Whitmore, Tantra & Sutisna l.c. 307. Synonym: Terminthodia Ridl., J. Fed.
Mal. St. Mus. 6 (1915) 141.
Shrubs or small to large trees. Branches unarmed. Leaves opposite, unifoliolate, margins
entire, pinnately veined; petioles wingless, articulated with the blade. Inflorescences
axillary panicles, sometimes 1- to few-flowered. Flowers bisexual, 4-merous; sepals 4,
fused at base, valvate or partially imbricate; petals 4, free, usually valvate, triangular,
spreading and becoming recurved; stamens 4, free, opposite the sepals, filaments flattened,
tapered, glabrous, elongating after anthesis, anthers ellipsoid, dorsifixed; staminodes 4,
alternating with the stamens, anthers minute, without pollen; disc broad, glabrous, rounded
to 4-angled; ovary 4-carpellate, carpels erect, free or fused at the base, joined near the apex
by a single style, locules 2-ovulate; style straight, composed of 4 elements twisted together,
stigma capitate. Fruits of 14 erect follicles, undeveloped carpels persistent in fruit, follicles
boat-shaped, free or fused at the base; exocarp and mesocarp leathery; endocarp
412
RUTACEAE (JONES)
413
cartilaginous, separating from the mesocarp. Seeds 12 per follicle, winged; testa papery;
endosperm fleshy; embryo straight.
Distribution. 6 species; S Thailand, Sumatra, Peninsular Malaysia, Borneo, Philippines,
Celebes, Papua New Guinea, and the Solomon Islands. The widely distributed T. tetrandrum
occurs in Sabah and Sarawak. The remaining species are localised endemics in other parts
of the range.
Ecology. Lowland to montane forests, peat swamp forests, heath forests, and subalpine
habitats, from near sea-level to 3300 m.
Tetractomia tetrandrum (Roxb.) Merr.
Fig. 16.
(Greek, tetra = four, andros = male; the number of stamens)

J. Str. Br. R. As. Soc. 76 (1917) 87, l.c. (1921) 314; Ridley l.c. (1922) 346; Craib l.c. 215; Masamune
l.c. 361; Burkill l.c. (1966) 2182; Stone l.c. (1972) 385; Hartley l.c. (1979) 132; Anderson l.c. 309;
Whitmore, Tantra & Sutisna l.c. 308. Basionym: Melicope tetrandra Roxb., Hort. Bengal. (1814) 88.
Type: Roxburgh, Icones 1411, Penang (K). Synonyms: Tetractomia beccarii Hook. f. l.c. (1875) 489,
Merrill l.c. (1921) 314, Masamune l.c. 361, Anderson l.c. 309; Terminthodia viridiflora Ridl. l.c.
(1915) 141; Tetractomia obovata Merr. l.c. (1917) 86, l.c. (1921) 314, Masamune l.c. 361;
Tetractomia holttumi Ridl., J. Bot. 62 (1924) 295; Tetractomia latifolia Ridl. l.c. (1930) 79,
Masamune l.c. 361, Anderson l.c. 309; Tetractomia montana Ridl. l.c. (1930) 79, Masamune l.c. 361,
Anderson l.c. 309; Tetractomia parviflora Ridl. l.c. (1930) 78, Masamune l.c. 361, Anderson l.c. 309.
Shrub or small to large tree to 30 m tall, 50 cm diameter; bole to 18 m tall. Branchlets

glabrous. Leaves sometimes clustered at branch ends, blades glabrous, leathery, obovate to
oblanceolate or elliptic, (2.5)7.522(32) x 114 cm; base rounded to attenuate, apex
emarginate or rounded to acuminate or mucronate; lateral veins 411 pairs; petioles 0.35
cm long, swollen at both ends. Inflorescences 1- to many-flowered, (0.8)416 cm long,
glabrous to sparsely hairy; pedicels 0.54 mm long. Flowers 2.510 mm wide; sepals
rounded to triangular, 0.41.5 mm long, glabrous to sparsely hairy, ciliolate when young;
petals green to yellowish, ovate-triangular, 14.5 mm long, usually glabrous; carpels free at
the base, glabrous or sparsely hairy, style glabrous. Fruits with free follicles, 411 mm
long, glabrous or nearly so.
Vernacular names. Sabah and Sarawakjampang, jampang rusa, medang rawang (preferred
common name), rawang mata, rawang paya (Malay).
Distribution. Throughout the range of the genus. In Sabah and Sarawak widespread; also in
Ecology. Common in secondary forests and open places on infertile organic soils, and also
mixed dipterocarp forest on humult ultisols, lowland peat swamp and heath forests, from
near sea-level to 1900 m.
Taxonomy. An extremely variable species in habit, leaf size and shape, and inflorescence,
flower and fruit size. In Sabah and Sarawak, this variation is characterised by a widespread,
414
RUTACEAE (JONES)
415
generalised form and several specialised forms or races which reflect the distribution of
certain habitats, i.e., lowland peat swamp forest, heath forest, lower montane forest, and
lowland dipterocarp forest. Hartley (l.c. 1979) describes these races and provides the names
of relevant previously described taxa assigned to these entities.
17. ZANTHOXYLUM L.
(Greek, xanthos = yellow, xulon = wood)
l.c. (1753) 270; Hooker f. l.c. (1875) 492; Kurz l.c. (1877) 180; King l.c. 213; Ridley l.c. (1922) 346;
Craib l.c. 218; Backer & Bakhuizen f. l.c. 96; Burkill l.c. (1966) 2326; Hartley, J. Arn. Arb. 47 (1966)
171, 51 (1970) 423; Stone l.c. (1972) 386; Anderson l.c. 309; Perry l.c. 369; Corner l.c. 670;
Whitmore, Tantra & Sutisna l.c. 308. Synonym: Fagara L., Syst. ed. 10 (1759) 897.
Scandent or erect shrubs or trees; dioecious or rarely monoecious, evergreen or deciduous.

Twigs and branches armed with spines or prickles. Leaves alternate, unifoliolate, trifoliolate or pinnate with up to 15 pairs of leaflets, with or without a terminal leaflet, leaflets
articulated at the base, leaf rachis sometimes winged. Inflorescences terminal or axillary
racemes, panicles or cymes. Flowers small, unisexual, rarely bisexual; sepals 45; petals
45; or perianth segments 68 and undifferentiated; stamens 46, opposite the sepals,
rudimentary in female flowers; disc flat or cushion-like; ovary 15-carpellate, rudimentary
in male flowers, carpels free or fused at base, ovules 2 per locule, styles fused to divergent,
stigma capitate. Fruits of 15 follicles, free or basally fused; outer wall glandular, red to
black; endocarp cartilaginous. Seeds ovoid to round, 1 per follicle, often hanging from the
dehisced follicle at maturity; testa black or reddish, glossy; endosperm white, fleshy.
Distribution. A large genus comprising approximately 200 or more species of mainly
pantropical distribution, with a few representatives in temperate eastern Asian and North
America. In Sabah and Sarawak, 4 species occur, two of which are trees or shrubs, and the
remaining two being scandent shrubs.
Ecology. Most of the species generally grow in rain forests and thickets at low and medium
elevations. The attractive seeds are dispersed by birds, accounting for the often wide and
sometimes discontinuous distribution of some of the species.
Uses. Various species are sources of spices and condiments, and local medicines. The
attractive wood of some is used for cabinetry and other fine work.
Key to Zanthoxylum species

1.
Erect shrubs or trees. Branchlets armed with straight prickles. Ovary 23-carpellate...2
Scandent or suberect shrubs. Branchlets and/or leaf axes armed with recurved prickles
or unarmed. Ovary 4-carpellate....................................................................................3
2.
Inflorescences paniculate. Leaflets 818 cm long. Carpels 3..........2. Z. myriacanthum

Inflorescences cymose. Leaflets 18 cm long. Carpels 2........................1. Z. avicennae
3.
Lateral leaflets subopposite to alternate. Midribs of leaflets not armed...........................

416
RUTACEAE (JONES)
Z. scandens Blume
l.c. (1825) 249; Backer & Bakhuizen f. l.c. 96; Hartley l.c. (1966) 177; Whitmore, Tantra &
Sutisna l.c. 308.
India, China, Taiwan, Ryukyu Islands, Sumatra, Java, Borneo; uncommon in Sabah
and Sarawak, known only from montane forest.
Climbing (occasionally suberect) shrub, dioecious. Leaves pinnate; leaflets 212
pairs, with margins finely glandular toothed. Inflorescences axillary, or axillary and
terminal panicles. Flowers 4-merous; petals occasionally purple-margined; ovary 4carpellate. Follicles in groups of 14, 45 mm in diameter.
Lateral leaflets opposite. Midribs of leaflet usually armed..............................................
Z. nitidum (Roxb.) DC.
l.c. (1824) 727; Ridley l.c. (1922) 1347; Burkill l.c. (1966) 2327; Hartley l.c. (1966) 180, l.c.
(1970) 423; Stone l.c. (1972) 386; Whitmore, Tantra & Sutisna l.c. 308.
India, Thailand, Vietnam, Malesia, Solomon Is., Australia. In Sabah (Ranau and
Pinangah districts), in forest to 1100 m.
Climbing (occasionally suberect) shrub, dioecious or (rarely) monoecious. Leaves
pinnate, leaflets 24 pairs, with margin entire to glandular toothed. Inflorescences
terminal and/or axillary, racemes or panicles. Flowers 4-merous; ovary 4carpellate. Follicles in groups of 14, 57 mm in diameter.
1. Zanthoxylum avicennae (Lam.) DC.

(Ibn Sina, or Avicenna, 9801037, Arabian medical writer and philosopher)
l.c. (1824) 726; Hartley l.c. (1966) 190, l.c. (1970) 423; Perry l.c. 370. Basionym: Fagara avicennae
Lam. l.c. 445. Type: dIncarville 179, China, Kwangtung (P). Synonyms: Zanthoxylum diversifolium
Warb. l.c. 339; Z. iwahigense Elmer, Leafl. Philip. Bot. 5 (1913) 1833.
Scandent or erect shrub or small tree to 15 m tall; dioecious, evergreen. Smaller branches
and twigs with straight or recurved prickles. Leaves pinnate, to 30 cm long, axis often
narrowly winged; leaflets 211 pairs, subopposite, ovate to elliptic-lanceolate, 18 x 13
cm, glabrous, thinly leathery; base rounded to cuneate, margin nearly entire to finely
glandular toothed, apex rounded to acuminate; lateral veins 411 pairs; petiolules 25 mm
long. Inflorescences terminal (occasionally axillary), flat-topped cymes, to 20 cm long, axis
glabrous. Flowers unisexual, 5-merous; sepals 5, triangular or rounded, under 1 mm long;
petals 5, elliptic, 12.5 mm long, white to greenish; stamens 5, rudimentary in female
flowers; disc flat or cushion-like; ovary 2-carpellate, rudimentary in male flowers, styles and
stigmas united, peltate. Fruits of 12 nearly round follicles, c. 4.5 mm in diameter, the
undeveloped carpels (if any) persistent in fruit.
Distribution. China, Vietnam, Thailand, Java, Borneo, Philippines, Celebes, Lesser Sunda
Islands and the Moluccas. In Sabah, reported only from Ranau district.
Ecology. Found locally on flat sites and hillsides in primary and secondary forest,
occasionally on ultramafic soil, to 1500 m.
Uses. The stems and bark are used medicinally as a tonic and for treating snake bite (IndoChina, Philippines).
417
418
2. Zanthoxylum myriacanthum Wall. ex Hook. f.
Fig. 17.
(Greek, myrios = numberless, akantha = thorn)

l.c. (1875) 496; King l.c. 214; Ridley l.c. (1922) 347; Burkill l.c. (1966) 2327; Hartley l.c. (1966) 185;
Stone l.c. (1972) 386; Anderson l.c. 309; Perry l.c. 371; Corner l.c. 671. Whitmore, Tantra, Sutisna l.c.
308. Type: Porter (Wallich Cat. No. 1214), Malacca (K). Synonyms: Fagara myriacantha (Wall. ex
Hook. f.) Engl. in Engler & Prantl l.c. (1896) 118.
Small to large tree to 30 m high, 35 cm diameter; dioecious, evergreen; bole to 15 m tall.

Stems with thick spines to 3 cm long; smaller branches and twigs with straight, hollow
prickles usually housing ants. Leaves pinnate, to 60 cm long; leaflets 411 pairs, opposite
or subopposite, elliptic, 818 x 38 cm, glabrous or shortly hairy below, leathery; base
blunt to nearly cordate and slightly asymmetric, margin finely glandular toothed, apex
acuminate; lateral veins 818 pairs; petiolules to 5 mm long. Inflorescences terminal and
axillary panicles, 1525 cm long, axes mostly glabrous. Flowers unisexual, 5-merous;
sepals 5, triangular, minute; petals 5, elliptic, 1.52.5 mm long, white or yellowish,
sometimes purplish; stamens 5; disc flat; ovary 3(4)-carpellate, rudimentary in male
flowers, styles and stigmas united, peltate. Fruits of 13 nearly round follicles, 36 mm in
diameter, the undeveloped carpels (if any) persistent in fruit.
Distribution. E India, N Vietnam, SW China, Sumatra, Peninsular Malaysia, Borneo,
Philippines. In Sabah, occurs scattered in Ranau, Tambunan and Keningau districts. In
Sarawak, collected once (Omar, s.n., Ulu Lawas).
Ecology. Locally found in primary and secondary forest in the hills and low mountains, to
1200 m.
Uses. The smoke of burning seeds is said to be inhaled for treating ulcerated syphilitic nose
(Peninsular Malaysia).
419
SIMAROUBACEAE
Julius Kulip & K.M. Wong
Sandakan, Malaysia
Merrill, EB (1921) 315; Ridley, FMP 1 (1922) 360; Masamune, EPB (1942) 361; Nooteboom, FM 1, 6
(1962) 193, FM 1, 6 (1972) 968, Blumea 11 (1962) 509; Keng, OFMSP (1969) 178; Kochummen,
TFM 2 (1972) 345; Cockburn, TS 1 (1976) 217; Anderson, CLTS (1980) 322; Corner, WSTM 2
(1988) 696; Whitmore, Tantra & Sutisna, CLK 2,1 (1990) 329.
Trees or shrubs (some sprawling), usually containing very bitter substances. Hairs mostly
simple and unicellular, sometimes with a glandular head. Leaves spirally arranged, simple
or pinnate, sometimes (in Ailanthus, Brucea and Soulamea) with pitted, concave, or flattish
glands on the lower surface; stipules usually absent except in Allantospermum, Irvingia and
Picrasma. Inflorescences usually compound, axillary, rarely terminal; plants monoecious or
dioecious. Flowers usually small, regular, unisexual or bisexual; sepals 35, almost always
partly united; petals 35, free; stamens inserted at the base of the disc, sometimes arranged
in two whorls, with the inner whorl alternate with the petals and the outer whorl (if present)
opposite the petals, anthers 2-celled, opening lengthwise; disc intrastaminal, sometimes
rather inconspicuous; ovary superior, 25-lobed, with 15 chambers, or with free carpels;
ovule one in each carpel, anatropous, placentation axile. Fruits usually not splitting, often
drupe-like, sometimes a samara or in Allantospermum septicidally splitting into 5 valves.
Seeds with scant or no endosperm.
Distribution. Some 30 genera and 200 species, distributed in the tropics and subtropics and
some in temperate Asia. In Sabah and Sarawak, represented by 9 genera with 11 species of
which only one (Harrisonia) includes scrambling shrubs.
Ecology. Simaroubaceae species are found mostly in lowland forest. Quassia indica shows
preference for temporarily inundated areas while Eurycoma longifolia has a distinct
preference for acidic, leached, well-drained soils. Soulamea amara occurs in the Barringtoniaformation of the coastal vegetation and prefers calcareous or rocky beaches. Pollination is
probably by insects as the flowers are often reported to be fragrant.
Uses. Ailanthus, Allantospermum and Irvingia are the only genera reaching timber size. All
the bitter-tasting genera are used locally in the preparation of traditional medicines,
especially as tonics, antidysenterics and antihelminthics, and the roots of Eurycoma
longifolia has been reported to contain biologically active compounds useful as anti-malaria
drugs (Chan et al., Planta Medica 52 (1986) 105).
Taxonomy. The Simaroubaceae are closely related to the Burseraceae, Meliaceae and
Rutaceae. While the genera are quite distinctly recognised, the limits of these families
overlap somewhat. Forman (Kew Bull. 19 (1965) 517) placed Irvingia in the Irvingiaceae
and Allanthospermum in the Ixonanthaceae but Nooteboom (l.c. (1972) 970) concluded that
421
the morphological, phytochemical and palynological evidence favoured including these

genera in the Simaroubaceae.
Key to genera
1.
Sprawling or scrambling spiny shrubs...........................................................................

Harrisonia R. Brown ex A. Juss.
Mem. Mus. Hist. Nat. Paris 12 (1825) 517; Masamune l.c. 362; Nooteboom, FM 1,6 (1962)
207.
34 species in the Old World Tropics. 1 species, H. perforata (Blanco) Merr.,

occurs in Sabah. Scandent prickly shrub; branches with thorns; leaf-stalk winged.
Lowlands to 400 m, also on Banggi island. Vernacular names: Sabahbogua
(Dusun Banggi), kait-kait (Tenom Murut), kukualang (Malay).
Shrubs or trees, unarmed.............................................................................................2
2.
Leaflets margin toothed.................................................................................3. Brucea

Leaves or leaflets margin entire...................................................................................3
3.
Leaves simple .............................................................................................................4

Leaves pinnate ............................................................................................................7
4.
Leaves spirally arranged, densely clustered at shoot tips, obovate, stalks 38 cm long
..................................................................................................................8. Soulamea
Leaves spiral to alternate and well-spaced along the shoots, elliptic, stalks 0.52 cm
long.............................................................................................................................5
5.
Leaves with pitted glands. Stipules absent.......................................7. Quassia (in part)

Leaves without such glands. Stipules either conspicuous conical structures or present
only in leaf buds on young shoots and falling off early.................................................6
6.
Twigs with annular stipule-scars, not swollen at leaf insertion. Leaf-stalks 1020 mm
long..............................................................................................................5. Irvingia
Twigs without annular stipule-scars, swollen at points of leaf insertion. Leaf-stalks
544 mm long................................................................................2. Allantospermum
7.
Leaflets sessile..........................................................................................4. Eurycoma

Leaflets distinctly stalked.............................................................................................8
8.
Leaflets very unequal at base. Fruits winged...............................................1. Ailanthus

Leaflets symmetric at base. Fruit a drupe.....................................................................9
9.
Veins sunken on both upper and lower leaflet surfaces; stipules absent..........................
.......................................................................................................7. Quassia (in part)
Veins prominent on both upper and lower leaflet surfaces; stipules present, suborbicular.....................................................................................................6. Picrasma
422
SIMAROUBACEAE (KULIP & WONG)
1. AILANTHUS Desf., nom. cons.

(from the Amboinese plant name aylanto)
Mem. Phys. Math. Ac. R. Sc. Paris (1786) 270, t. 8; Nooteboom, FM 1,6 (1962) 215; Kochummen l.c.
Dioecious trees with straight bole but without buttresses. Bark pale grey, smooth or
lenticellate; inner bark yellowish brown. Sapwood pale white. Twigs thick, with large leafscars. Leaves more or less tufted at the ends of twigs, pinnate, to 60 cm long, without a
terminal leaflet; leaflets 1013 pairs, 7.512.5 x 2.55 cm, hairy below, with scattered
glands in the forks of the veins on the underside; base very unequal, margins entire, apex
pointed; the leaves eventually drooping with bowed stalk. Flowers 56-merous; calyx small,
56-lobed, closed in bud and later irregularly splitting (often 2-lobed) to the base, rarely
cupular; petals 56, induplicate-valvate in bud, concave; stamens 10, in male flowers
inserted below the outer margin of the disc, in female flowers either of subnormal size (but
without pollen), or vestigial, or absent, anthers opening laterally or externally, the 2 cells
free in their lower half; ovary 25-carpellate, carpels free, flat, in the male flower vestigial
or absent; styles 25, free or united; ovule 1 in each carpel. Fruit a samara, elliptic or
oblong-lanceolate. Seeds flat, orbicular or obovate or somewhat triangular, without endosperm.
Distribution. 5 species in tropical and subtropical SE Asia from Turkestan and India to
China, through Malesia to Solomon Islands, Queenslands and northern New South Wales in
Australia. In Sabah and Sarawak, 2 species are known.
Ecology. In Sabah and Sarawak, both species are uncommon and found mainly in lowland
forests below 1000 m, in valleys, along streams, and open places.
Key to Ailanthus species

Leaflets with lower surface glabrous and with a pair of large glands..........1. A. integrifolia
Leaflets with lower surface hairy or sparsely hairy and with scattered small glands..............
......................................................................................................................2. A. triphysa
1. Ailanthus integrifolia Lam.

(Latin, integer = entire, folia = leaves or leaflets; the leaflet margins)
Dict. 3, 2 (1792) 417; Merrill, Interpr. Rumph. (1917) 299; Nooteboom, FM 1, 6 (1962) 218;
Kochummen l.c. 346; Cockburn l.c. 219; Whitmore, Tantra & Sutisna, l.c. 229. Type: Rumphius
Herb. Amb. 3:205, t. 132. Synonyms: A. blancoi Merr., Sp. Blanc. (1918) 205; A. peekelii Melch.,
Notizbl. Berl. Dahl. 10 1930) 893; Dysoxylum dasyphyllum Miq., Ann. Mus. Bot. Lugd. Bat. 4 (1868)
19.
Tree to 55 m tall, 65 cm diameter. Bark smooth, light brown or grey. Sapwood white,
yellow, pale brown or creamish, very soft; heartwood absent. Leaves 30200 cm long,
stalks 520 cm long; leaflets 29 pairs, 3.514 x 3.36.2 cm, glabrous on both surfaces;
base very oblique or sickle-shaped, apex blunt-acute; stalks 0.51.5 cm long; lateral veins
423
613 pairs; glands on lower surface large, black, flat, oblong, 0.55.0 mm diameter, mostly
paired near the base. Inflorescences to c. 40 cm long, glabrous, pedicels to c. 15 mm long.
Flowers with calyx more or less pubescent, closed in bud, rupturing and toothed irregularly,
rarely cupular, 14 mm high, rarely caducous; petals puberulous, acute or bluntish, to c. 9 x
3 mm; filaments with many long spreading hairs to glabrous, usually thickened downwards,
c. 0.5 mm in female flowers, to 4 mm long in male flowers; anthers c. 1 mm in female
flowers, to 2.5 mm long in male flowers; ovary 5, usually densely puberulous; styles 5,
connate at the base, including the long, stellately spreading stigmas, to c. 6 mm long. Fruits
of (1)35 samaras, each somewhat elliptic, 1122 x 2.55 cm, the vein reticulations
distinct on the outside, pale green; stalk 2.55 cm long. Seeds flat; testa thin; cotyledons 2.
Distribution. Malesia: all islands, except Java and the Lesser Sunda Islands, and Melanesia
(Bismarcks and Solomons). In Sabah, the species has been recorded from the Sandakan and
Beaufort districts. In Sarawak, once collected at Suai, Miri, 4th Div. (S. 39203). Also in
Kalimantan.
Ecology. In primary rain forest or rarely secondary forest, very rare.
Uses. In New Guinea and the Bismarcks, the timber is made into planks for house
construction (Nooteboom l.c.).
2. Ailanthus triphysa (Dennst.) Alston
Fig. 1.
(Greek, tri = 3-partite, phusis = in character; the calyx-tube)

Handb. Fl. Ceyl. 6, Suppl. (1931) 41; Nooteboom, FM 1,6 (1962) 219; Kochummen l.c. 346;
Cockburn l.c. 219; Whitmore, Tantra & Sutisna l.c. 329. Basionym: Adenanthera triphysa Dennst.,
Schluss. Hort. Mal. (1818) 32. Type: Dennst., Schluss. Hort. Mal. (1818) t. 32. Synonyms: A.
philippinensis Merr., Publ. Gov. Lab Philip. 35 (1906) 25; Hebonga obliqua Radlk., Philip. J. Sc. 6
(1911) Bot. 366.
Tree to 60 m tall and 50 cm diameter. Bark pale brown, smooth to lenticellate; inner bark
yellowish brown, mottled. Sapwood white. Twigs thick, reddish brown hairy. Leaves to 30
cm long, rachis slightly pubescent; leaflets almost sickle-shaped, c. 18 pairs, 7.511.7 x 3 4
cm, membranous, upper surface glabrous, lower surface densely hairy; base distinctly
unequal (one side sharply acute, the other side rounded), margin wavy, apex pointed; lateral
veins 820 pairs, more or less sunken above; glands small, scattered over the whole surface
especially on the midrib and veins. Inflorescences many-flowered, more or less pubescent,
c. 2060 cm long; bracts small, ovate to triangular, falling early; pedicels to c. 4 mm.
Flowers with calyx pubescent, less than 1 mm high, the triangular acute lobes as long as the
tube or a little longer; petals glabrous or nearly so, 35 x 11.5 mm; filaments twistedfolded in bud, filiform or sometimes attenuating from the base to the top; anthers c. 1.2 mm
long, 1 mm wide in male flowers, smaller in female flowers. Fruits of 13(4) samaras,
each somewhat obovate, 4.58 x 1.52.5 cm, ripening red; stalk 0.82 cm long.
424
425
426
Distribution. India, Sri Lanka, Burma, Thailand, Vietnam, through Malesia (except
Sumatra, Peninsular Malaysia, Lesser Sunda Is., and New Guinea) to Queenslands and the
northern parts of New South Wales. In Sabah uncommon, recorded from the interior parts
at Sook, Keningau and Tenom, and in the east coasts at Lahad Datu. Not yet recorded in
Sarawak. Also in Kalimantan.
Ecology. In primary lowland and hill forests.
Uses. The resin is tapped and used as incense and traditional medicine in India. In IndoChina the bark is burned as incense. The bark and the leaves are used for making a tonic,
especially for post-childbirth. They also possess febrifuge properties and are used for
treating dyspeptic complaints. The wood is used for making wooden shoes in Luzon (the
Philippines), for fishing floats, catamarans, sword-handles, and spear-sheath in India, and
for tea-boxes in Sri Lanka (Nooteboom l.c.).
2. ALLANTOSPERMUM Forman
(Greek, ala = wing, sperma = seed; the winged seeds)
Kew Bull. 19 (1965) 516; Nooteboom l.c. (1972) 968; Kochummen l.c. 347; Cockburn l.c. 217;
Whitmore, Tantra & Sutisna l.c. 329.
Trees. Twigs without annular stipular scars, swollen at points of leaf insertion. Leaves
simple, entire. Inflorescence a panicle. Flower bisexual, 5-merous; sepals 5; petals 5;
stamens 10, free, anthers versatile; disc 10-lobed, intrastaminal; ovary superior, shallowly 5lobed, stigma with a tiny papillose head. Fruit a capsule, broadly ellipsoidal, twisted after
splitting into 5 valves along the septa, leaving a central columella. Seeds cylindrical,
ellipsoid, shiny and waxy.
Distribution. 2 species, 1 in Malesia (Peninsular Malaysia and Borneo) and the other in
Madagascar (A. multicaule (Capuron) Noot.).
Allantospermum borneense Forman
Fig. 2.
(of Borneo)
l.c. 517, t. 1; Nooteboom l.c. 1972) 972; Kochummen l.c. 347; Cockburn l.c. 217; Whitmore, Tantra &
Sutisna l.c. 329. Type: Galau S. 15262, Sarawak, 1st Div., Semengoh Forest Reserve (holotype K;
isotypes L, SAN, SAR).
Tree, to 90 m tall and 60 cm diameter; bole fluted and often crooked; buttresses sharp and
spreading, to 3 m high. Bark pale brown with grey patches, smooth or with distant,
adherent, large, thinnish scales, minutely lenticellate; inner bark pink, mottled white.
Sapwood yellowish brown. Twigs brown, slender, more or less zig-zag. Leaves elliptic to
oblong, 6.515 x 1.73.3 cm, glossy above, dull beneath; base cuneate to broadly rounded,
apex blunt-acuminate; midrib and veins prominent on both surfaces; lateral veins 510 pairs,
intermediate veins extending about half-way to the margin; stalks blackish, channelled
427
428
above, 0.54.4 cm long. Inflorescences (1)3.35.9 cm long, bearing the scars of early
caducous bracts. Flowers with a stalk 79 cm long; sepals white, boat-shaped, 34 x 2 mm,
rounded at the apex, reflexed at anthesis; petals white, elliptic to obovate, 45 x 2.53 mm,
membranous, reflexed at anthesis, caducous; stamens to 6 mm long, anthers c. 1 mm long;
disc c. 1.5 mm diameter and 0.5 mm thick; ovary 5-lobed, c. 1.5 x 2 mm, style filiform, 34
mm long, purple, stigma knob-like. Fruits broadly ellipsoid, 5-lobed, 2.84.2 x 13.5 cm,
apical beak 210 mm long. Seeds cylindrical, often slightly curved, 22.5 x 46 mm.
Key to subspecies
Leafy branches conspicuously zig-zag. Infloresences 3.35.9 cm long, laxly branched.
Fruits 23.5 cm wide, apical beak 24 mm long...................................................................
subsp. borneense
Peninsular Malaysia, Borneo (Sabah, Sarawak, Brunei, Kalimantan). In primary mixed
dipterocarp forest on sandy humult ultisols; in Sabah also on ultramafic soils. Apparently
gregrariously flowering and fruiting at several year intervals.
Leafy branches only slightly zig-zag. Inflorescences 12 mm long, condensed. Fruits about
1.1 cm wide, apical beak 610 mm long...............................................................................
subsp. rostratrum Noot.
l.c. (1972) 972; Cockburn l.c. 217; Whitmore, Tantra & Sutisna l.c. 330. Type: Agama SAN
36068, Sabah, Lahad Datu, Pulau Sakar (holotype L; isotypes K, SAN).
Known only from Sabah (Sandakan and Lahad Datu areas).

Vernacular names. Sarawaknyalin (Iban). Bruneikayu tulang (Malay), tulang (Iban).
3. BRUCEA J.F. Mill., nom. cons.

(J. Bruce, 17301794, a Scottish scholar and explorer)
Icon. (1779) t. 25; Merrill l.c. (1921) 316; Ridley l.c. 361; Masamune l.c. 361; Corner l.c. 697;
Nooteboom, FM 1, 6 (1962) 209; Kochummen l.c. 348; Cockburn l.c. 219; Anderson l.c. 322.
Shrubs or small trees. Leaves pinnate, with a terminal leaflet; leaflets with toothed margins;
stipules none. Flowers unisexual, in axillary inflorescences; sepals 4, united at base; petals
4, free; disc thick with 4 lobes; stamens 4, with short filaments, vestigial or absent in female
flowers; ovary 4-carpellate, carpels free, styles free or united at base; ovules 1 in each
carpel, attached above the middle. Fruit a drupe, hardly fleshy, with a stone. Seeds with
very thin endosperm.
Distribution. 6 species in tropical Africa and Asia, including 2 in Malesia. In Sabah and
Sarawak one species.
Brucea javanica (L.) Merr.
Fig. 3.
(of Java)
J. Arn. Arb. 9 (1928) 3; Kochummen l.c. 348; Cockburn l.c. 219; Nooteboom, FM 1, 6 (1962) 210;
Anderson l.c. 322. Basionym: Rhus javanica L., Sp. Pl. (1753) 265. Type: Osbeck, s.n., Java (L).
429
430
Synonyms: Lussa radja Rumph., Herb. Amb. (Auct.) 7 (1755) 27, t. 15; Brucea sumatrana Roxb.,
Hort. Beng. (1814) 12; Brucea sumatrensis Spreng., Pl. Min. Cogn. 2 (1815) 90; Brucea amarissima
Desv. ex Gomes, Mem. Acad. Sc. Lisb. n.s. 4, 1 (1872) 30.
Shrub or small tree to 5 m high. Leaves 2040 cm long; leaflets ovate to oblong-lanceolate,
315, 4.511 x 1.54 cm, sparsely hairy above, more or less pubescent below, sometimes
glabrous; stalks 25 mm, the terminal one much longer. Flowers greenish white to greenish
red or purple. Fruits 14 together, 45 mm long.
Vernacular names. Sabahkuinin (Dusun/Kadazan Tambunan), mara (Maga), pait-pait
(Dusun/Kadazan Kinabatangan), payas (Dusun/Kadazan Ranau), tongkat ali (Papar Malay; in
common with Eurycoma). Sarawakjaloot (Murut).
Distribution. From Sri Lanka and the Deccan Peninsula through SE Asia to S China and S
Formosa, throughout Malesia and N Australia. In Sabah, it is found all over the state. In
Sarawak, it has been recorded from the 1st, 2nd, 4th and 7th Div. Also in Brunei.
Ecology. A common, light-tolerant plant, preferring open sites and secondary forest and
thickets, forest edges and ridges, even occurring in sunny places in sandy dunes and on
limestone rock. Flowering and fruiting throughout the year.
Uses. The roots and fruits contain bitter principles which possess medicinal value and used
as concoctions in the treatment of dysentery, diarrhoea and fever.
4. EURYCOMA Jack
(Greek, eurus = broad, kome = tuft or crust; the leaves crowded at the ends of branches)
Mal. Misc. 2 (1822) 45; Merrill l.c. (1921) 316; Ridley l.c. 361; Burkill, EPMP (1935) 984;
Masamune l.c. 361; Corner l.c. 698; Nooteboom, FM 1, 6 (1962) 203; Kochummen l.c. 349; Cockburn
l.c. 219; Anderson l.c. 323.
Trees, treelets or shrubs. Twigs stout with large leaf-scars. Leaves pinnate, with terminal
leaflet, crowded at branch ends; leaflets sessile, opposite or subopposite; base slightly
oblique, attached to the rachis with a prominent joint; lateral veins inconspicuous above and
below. Inflorescence a downturned axillary panicle; plants monoecious or dioecious.
Flowers unisexual, female always with large sterile stamens, males always with a pistillode; calyx small, 56-lobed; petals 56; stamens 56, on the sepals alternating with 56
small staminodes, stamens and staminodes sometimes united with the base of petals; disc
inconspicuous; carpels 56, each with 1 ovule, free, the styles slightly united, stigma peltate,
56-lobed. Fruit a nut, to 5 per flower, each on a short stalk c. 3 mm long, ellipsoid or
ovoid. Seeds without endosperm.
Distribution. 3 species in tropical SE Asia, Sumatra, Borneo and S Philippines. In Sabah
and Sarawak, only 1 species.
431
432
Eurycoma longifolia Jack
Fig. 4.
(Latin, longus = long, folium = leaves)

l.c. 45; Ridley l.c. 362; Merrill l.c. (1921) 316, PEB (1929) 116; Masamune l.c. 361; Burkill l.c. 984;
Corner l.c. 604; Nooteboom, FM 1, 6 (1962) 205; Kochummen l.c. 349; Cockburn l.c. 219; Anderson
l.c. 323. Type: Jack s.n., Sumatra (holotype K). Synonyms: E. merguensis Planch. in Hooker, Lond.
J. Bot. 5 (1846) 584; Picroxylon siamense Warb., Fedde Rep. 16 (1919) 256; Manotes asiatica Gagn.,
Bull. Soc. Bot. Fr. 98 (1951) 207.
Spindly unbranched tree or shrub, to 8 m tall and 15 cm diameter, or with a few upright
branches, each crowned by an umbrella-like rosette of leaves. Bark greyish brown, smooth.
Leaves to 100 cm long; leaflets lanceolate to obovate-lanceolate, rarely oblong, 520 x
1.56 cm; base oblique, apex blunt to slightly acuminate; midrib raised on both surfaces;
lateral veins inconspicuous above and sunken below. Flowers reddish; petals hairy on both
sides, c. 4.55.5 x 1.52.5 mm; styles rather long, stigma c. 1 mm above the ovaries. Fruits
1017(20) x 512 mm.
Vernacular names. Sabahionadiandau, nuad-mandau (Runggus), tombuid (Dusun/Kadazan
Tambunan), tongkat ali (Malay), tongkat langit (Kuala Penyu Malay). Sarawakbedara
(Semantan Malay), sengkanyat (Iban), sengkayap (Iban), tongkat ali (Santubong & Miri),
tungkat ali (Lundu).
Distribution. Lower Burma, Thailand, Laos, Cambodia, Indo-China; Malesia: Sumatra,
Peninsular Malaysia, and Borneo. Common throughout both Sabah and Sarawak; also in
Brunei and Kalimantan
Ecology. Abundant on well-drained sandy soils below 1200 m, in primary and secondary
mixed dipterocarp, heath and submontane forests.
Uses. In Sabah the roots are mixed with other medicinal plants, e.g., Cinnamomum species,
and used to prepare a health tonic. In Brunei, the bark is used as a blood coagulant in
complication during childbirth. The young leaves can be eaten raw to cure stomach-aches.
In Peninsular Malaysia, Chan et al. (Planta Medica 52 (1986) 105) reported that methanolextracts of roots contained biologically active compounds showing a strong antiplasmodial
activity against a multi-drug resistant K1 strain of Plasmodium falciparum from Thailand.
The putative aphrodisiac properties of the roots have, as yet, been substantiated by rigorous
experiment.
5. IRVINGIA Hook.f.
(E.G. Irving, 18161855, Scottish botanist)
pauh kijang
Trans. Linn. Soc. 23 (1860) 167; Ridley l.c. 363; Corner l.c. 699; Nooteboom, FM 1, 6 (1962) 223;
Kochummen l.c. 350; Cockburn l.c. 221; Anderson l.c. 323; Whitmore, Tantra & Sutisna l.c. 330.
Large trees; buttresses steep, to 6 m high. Bark fawn, smooth with distant loose scales,
minutely lenticellate; inner bark mottled, cream-yellow. Sapwood orange-brown. Twigs
with stipules forming a narrow, conical cap surrounding the terminal buds, soon falling,
leaving conspicuous annular scars. Leaves simple, glabrous, entire. Inflorescences axillary
433
and terminal panicles. Flowers (4)5-merous, bisexual; sepals connate (united) at the base;
petals overlapping in buds; stamens twice as many as petals, inserted beneath the large,
cushion-shaped, intrastaminal disc; ovary 2-chambered, conical or somewhat flattened,
sessile; style 1, stigma inconspicuous; ovules solitary. Fruit a drupe, large, 12-seeded,
resembling a mango.
Distribution. 3 species in tropical Africa and 1 species in tropical SE Asia and W Malesia.
Ecology. Frequent in lowland forests.
Uses. The fruit of all species is edible, but usually only the seeds are eaten.
Irvingia malayana Oliv. ex A.W. Benn.
Fig. 5.
(of Malaya)
in Hooker f., Fl. Brit. Ind. 1 (1875) 522; Ridley l.c. 364; Corner l.c. 699; Nooteboom, FM 1, 6 (1962)
223; Kochummen l.c. 350; Cockburn l.c. 221; Burgess TBS (1966) 455; Anderson l.c. 323; Whitmore,
Tantra & Sutisna l.c. 330. Type: Maingay 298, Malacca (holotype K). Synonyms: Irvingia oliveri
Pierre, Fl. For. Coch. 4 (1892) t. 263 B; Irvingella malayana van Tiegh., Ann. Sc. Nat. 9, 1 (1905)
276; Irvingella oliveri (Pierre) van Tiegh. and Irvingella harmandiana van Tiegh. l.c. 279; Irvingia
harmandiana (van Tiegh.) Pierre ex Lecomte, Fl. Gen. I.-C. 1 (1911) 701; Irvingia longipedicellata
Gagnep., Fl. Gen. I.-C. Suppl. 1 (1946) 670.
Medium-sized to large tree reaching 50 m tall and 50 cm diameter, with big limbs;
buttresses steep, plank-like and spreading, to 3 m high. Bark greyish to whitish, scaly to
flaky, sometimes smooth, minutely lenticellate. Leaves elliptic-oblong to lanceolate, 820 x
2.59 cm, upper surface shiny, lower surface slightly glaucous especially when fresh; base
often rounded, apex usually pointed; midrib raised above; lateral veins 1016 pairs, looping
and joining at margin, prominent on both surfaces; stipule-cap 34 cm long. Flowers
greenish white or yellowish, small. Fruits ellipsoid, c. 6 x 4 cm, slightly glaucous. Seedling
with first two leaves opposite; germination epigeal.
Vernacular names. Sabahmengkudu (Dusun/Kadazan Tuaran/Ranau; doubtful, as this
name normally refers to Morinda in the Rubiaceae), pauh kijang (Malay), selangan tandok
(Malay), tenghilan (Dusun/Kadazan Tuaran). Sarawakpatok entilit (Iban).
Distribution. Thailand, Indo-China, and Malesia (Sumatra, Peninsular Malaysia, Borneo
and Bawean). Widespread in Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. Scattered in mixed dipterocarp forest on clay-rich soils, to 300 m.
Timber. Burgess (l.c. 455) summarises the timber properties of this species. Pauh kijang
produces a very strong and springy timber, hard to saw and work, due to the high density. It
takes a very fine finish, and requires very little filling, and turns very well. In Sandakan,
furniture of this timber glued with synthetic resin glues has tended to fail at the glue-line.
434
Uses. The yellow wood is too hard to work with and not very durable. In Peninsular
Malaysia, it has been used for making kris-handles and handles of tapping knives
(Nooteboom, l.c.). The seeds contain a creamy yellow, nice-smelling fat known as "dika"
fat in Europe, used for making soap, wax, and candles. The seeds can also be eaten
(Nooteboom, l.c.).
6. PICRASMA Blume
(Greek, pikros = bitter, osme = smell or taste; the bark and other parts)
Bijdr. 5 (1825) 247; Ridley l.c. 361; Burkill l.c. 1723; Nooteboom, FM 1, 6 (1962) 212; Kochummen
l.c. 351; Anderson l.c. 323; Whitmore, Tantra & Sutisna l.c. 330.
Trees or shrubs. Leaves pinnate, with terminal leaflet, stalk base and rachis nodes usually
swollen; leaflets opposite or subopposite, entire, veins prominent on both upper and lower
surfaces; stipules present, suborbicular and falling off early. Inflorescences axillary, longpeduncled, compound-cyme, unisexual (plants monoecious or dioecious). Flowers 45merous, female usually twice as large as male; sepals small, free to united half way up;
petals persistent in female, much longer than the sepals; stamens 45; disc thick; carpels up
to 7, free, each with 1 ovule, vestigial or absent in male; styles united except at base,
sometimes 1 or 2, free; stalks jointed in the lower half. Fruit 14, drupe-like; exocarp thin,
fleshy, wrinkled when dry; endocarp hard. Seeds without endosperm.
Distribution. About 8 species, 6 in tropical America, 2 species in Asia including 1 in
Malesia.
Uses. Picrasma species contain alkaloids, the source of quassia chips used in insecticide.
Picrasma javanica Blume
Fig. 6.
(of Java)
l.c. 248; Ridley l.c. 361; Burkill l.c. 1723; Nooteboom, FM 1, 6 (1962) 213; Kochummen l.c. 351;
Anderson l.c. 323; Whitmore, Tantra & Sutisna l.c. 330. Type: Blume, s.n., Java (holotype L).
Synonyms: P. nepalensis A.W. Benn., Pl. Jav. Rar. (1844) 201; P. andamanica Kurz ex A.W. Benn.
in Hooker f. l.c. (1875) 520; P. philippinensis Elmer, Leafl. Philip. Bot. 5 (1913) 1837.
Tree to 24 m tall and 25 cm diameter; bole fluted. Bark dark, smooth, brittle; inner bark dull
yellow. Sapwood with clearly visible vessels. Leaves with 57 leaflets, stalk 26 cm long;
leaflets entire, 420 x 110 cm; base wedge-shaped, margin wavy or wrinkled, apex
acuminate; lateral veins 38 pairs, petiolules to 7 mm long; stipules leafy, nearly rounded,
725 x 520 mm, usually falling off early leaving a large scar. Inflorescences to 20 cm
long. Flowers 4-merous, white to yellow or green; sepals glabrous to puberulous, triangular
to ovate, c. 1 mm; petals ovate-oblong or oblong, often acute-acuminate to mucronate,
glabrous, or sparsely hairy, with a conspicuous midrib; stamens usually longer than petals in
male flowers, shorter than petals in female flowers, filaments gradually thinner towards the
top, hairy at base, 0.52 mm long in female flowers and 15 mm in male flowers, anthers
12 mm long in male and to 1 mm and empty in female flowers; ovary 4-lobed, 4-carpellate,
glabrous to hairy, styles 11.5 mm long, stigmas c. 2 mm. Fruits green to red or blue, ovoid
to depressed globlose, 910 x 712 mm. Seeds with a broad hilum; testa rather thick and
hard.
435
Vernacular names. Sabahbalimbing, panguban (Dusun/Kadazan Tambunan). Sarawak

kayu pahit (Malay).
Distribution. Tropical SE Asia (from Sikkim, Assam, Burma and Tonkin southward to
Malesia). In Borneo, recorded in Sarawak only at about 400 m on the Gunung (Mt.) Api
limestone, at Mulu, and in Sabah on the west coasts.
Ecology. Uncommon, usually scattered in rainforests from near sea-level to 1500 m.
Uses. The bark contains quassin, which gives its bitter taste. In Java, the leaves have been
applied for treating sores (Nooteboom l.c.). The trunk is too small for timber and the wood
is not durable (Burkill l.c.).
7. QUASSIA L.
(named after a slave in Surinam who reported
the medicinal properties of the wood to Dalberg, friend of Linnaeus)
Sp. Pl. ed. 2 (1762) 553, l.c. (1763) 1679; Merrill l.c. (1921) 315 (as Samadera); Ridley l.c. 363;
Burkill l.c. 1945; Masamune l.c. 362 (as Samadera); Nooteboom, FM 1, 6 (1962) 198, Blumea 11
(1982) 514; Kochummen l.c. 352; Cockburn l.c. 217.
Trees or shrubs. Leaves pinnate or simple, with pitted glands on the upper surface along the
margin and especially at the apex; stipules and scars absent. Inflorescence a simple or
branched raceme, a panicle, or an umbel. Flowers 46-merous, unisexual or bisexual, or
polygamous; petals imbricate or contorted in bud, longer than the calyx, sometimes very
long; stamens hairy, adaxial scale with a shorter or longer free apex; disc cylindrical or
subglobose; carpels free, more or less puberulous; style 1, with a terminal, inconspicuous
stigma. Fruits 16 per flower, drupaceous or woody, often compressed laterally, with a
narrow, unilateral, sharp-edged thinner part in the apical half. Seeds with a thin testa,
without endosperm.
Distribution. Pantropical, c. 25 species in tropical and subtropical America, 510 species in
Africa, 2 species in lower Burma and Cambodia (one of which is found almost throughout
Malesia), 1 species endemic to Borneo and Sumatra, and 2 species in Queenslands.
Ecology. In lowland rain forests.
Key to Quassia species

Leaves pinnately compound. Flowers in panicles. Fruits 15 from each flower, drupaceous,
slightly flattened, ellipsoid.........................................................................1. Q. borneensis
436
437
438
Leaves simple. Flowers in umbel-like clusters. Fruits 14 from each flower, strongly
flattened, with straight inner and semicircular outer margin..............................2. Q. indica
1. Quassia borneensis Noot.

(of Borneo)
FM 1, 6 (1962) 203, Blumea 11 (1962) 518; Cockburn l.c. 219; Anderson l.c. 323; Whitmore, Tantra
& Sutisna l.c. 330. Type: Meijer SAN 20499, Sabah (holotype L; isotypes K, SAN).
Tree to 25 m tall and 25 cm diameter; buttresses low. Bark pale yellow to greyish brown,
densely fissured-corky. Sapwood white. Leaves pinnate, spirally arranged; leaflets 24
pairs, elliptic to obovate-oblong, 812 x 44.5 cm, glabrous, upper surface shiny, lower
surface dull; with small pitted glands along the margins and in the acumen on the upper
surface; base cuneate, apex shortly rounded to acuminate; lateral veins sunken on both
surfaces, obscure, ending in a marginal vein; stalk c. 5 cm long, rachis terete, petiolules
11.5 cm long, articulated at the base. Inflorescences puberulous all over, shorter than the
leaves. Flowers (male) 45-merous, pedicels to 7 mm long; calyx c. 1 mm high; petals
contorted or imbricate in bud, glabrous, elliptic to ovate-oblong, 34 x 2 mm; stamens
slightly shorter than the petals, anthers oblong; disc c. 0.5 mm high, at the base c. 2 mm
wide and at the apex c. 1 mm wide, the upper half distinct from the lower half and folded
around the barren ovaries. Fruits 15 in each flower, drupaceous, prune-shaped, dark
purple-red when ripe, slightly flattened-ellipsoid, with a faint dorsal and ventral ridge, 23 x
1.5 cm; pericarp thin but hard. Seeds with a thin testa; cotyledons large, green, planoconvex.
Vernacular names. Sabahmamungal (Malay), pait-pait (Malay). Sarawakmedang pahit
(Malay).
Distribution. Malesia: Sumatra (Indragiri), Borneo (Sabah and Sarawak). Uncommon.
Ecology. Primary mixed dipterocarp forest on humult ultisols; also, rarely, in peat swamp
and kerangas (heath) forests.
2. Quassia indica (Gaertn.) Noot.
Fig. 7.
(of the Indies)

FM 1, 6 (1962) 199, Blumea 11 (1962) 517; Kochummen l.c. 352; Cockburn l.c. 217; Anderson l.c.
323; Whitmore, Tantra & Sutisna l.c. 330. Basionym: Samadera indica Gaertn., Fruct. 2 (1791) 352,
t. 156, f. 3. Type: Gaertner, Fruct. 2 (1791) 352, t. 156, f. 3. Synonyms: Manungala pendula Blanco,
Fl. Filip. (1837) 306; Samadera brevipetala Scheff., Nat. Tijd. Ned. Ind. 32 (1871) 410.
Tree or shrub, to 20 m and 8 cm diameter. Bark brownish green, smooth; inner bark
pinkish. Sapwood pale yellow; cambium yellow. Branchlets with small pith, with several
stiff persistent scales at the base of each shoot. Leaves simple, elliptic-oblong to lanceolate,
1213 x 412 cm; base acute or sometimes rounded, or subcordate, apex blunt or acuminate
or sometimes rounded; midrib, lateral and intercostal veins prominent on both
439
440
surfaces; stalks 12.5 cm long. Inflorescences jointed at the lower half, 0.52.5 cm,
growing during anthesis; bracts minute. Flowers to 20 or more; calyx 4-lobed, 23 mm
long, lobes about as long as or longer than the tube, puberulous outside; petals 4, free,
dorsally puberulous, obtuse, usually narrowed to the base, creamy green to violet, to 3 x 1
cm; filaments puberulous, hairy except toward the apex, to 2.5 cm long, inserted at the base
of the disc, anthers lanceolate to oblong, c. 4 x 2 mm; styles to 2 mm long. Fruits 14
together, flattened, with straight inner and semicircular outer margin, which is sharp and
thinner in the upper half, the apex more or less overtopping the subapical stylar scar, 49 x
2.5 cm.
Vernacular names. Sabahkacang-kacang (Malay), kelapahit (Sook Murut). Sarawak
manuggal (Iban).
Distribution. Madagascar, Sri Lanka, S Concan, Malabar, Lower Burma (Martaban,
Tenasserim), Andamans, and Cochinchina, through Malesia to the Bismarcks and
Solomons. In Sabah, it is found mainly in the east coast (Sandakan & Lahad Datu districts),
with only a few records from the west coast (Sipitang district). In Sarawak, it is found
throughout the state.
Ecology. Locally abundant in tidal swamp forests below 150 m, sometimes in localities
which are periodically inundated by fresh or salt water, for example on the edge of
mangroves. Occasional in freshwater swamp forest; also occurs in mixed dipterocarp forest.
Uses. In Sarawak, the wood is used for making knife-handles. The seeds are given as an
emetic and purgative, and sometimes in bilious fevers (Nooteboom l.c.).
8. SOULAMEA Lam.
(soelamoe, a Ternatean name for the plant)
Dict. Enc. Meth. 1 (1783) 449; Masamune l.c. 362; Nooteboom, FM 1,6 (1962) 221; Cockburn l.c.
217.
Shrubs or small trees. Leaves simple, obovate, spirally arranged, densely clustered at shoot
tips, sometimes with few glands underneath. Flowers in axillary racemes or narrow thyrses,
3(45)-merous, bisexual; floral parts persistent; sepals more or less connate at the base,
slightly imbricate in bud; petals longer than sepals; stamens twice as many as petals, in 2
distinct rows, inserted under the lower outer margin of the disc, anthers versatile; disc
3(45)-lobed, each lobe forked; ovary (1)23-carpellate, styles horizontally adnate to
their carpels, stigmas small; ovules sessile. Fruits dry, (1)2(3)-celled, indehiscent, flattened,
distinctly winged, more or less emarginate, rarely flattened, ovoid, acute. Seeds attached
adaxially nearly halfway down; testa thin; cotyledons plano-convex.
Distribution. 9 species. One species is endemic to the Seychelles, 6 species occur in New
Caledonia, and one species in Fiji. One species is widely distributed in Malesia and Polynesia,
and it occurs in Sabah and Sarawak.
441
Soulamea amara Lam.
Fig. 8.
(Latin, amarus = bitter; the taste of the tissues)

l.c. (1783) 449; Miquel, Fl. Ind. Bat. 1, 2 (1859) 129; Masamune l.c. 362; Nooteboom, FM 1, 6 (1962)
221; Cockburn l.c. 217. Type: Rumphius Herb. Amb. t. 415 (L). Synonyms: Rex amaroris Rumph.,
Herb. Amb. 2 (1743) 129, t. 41; Cardiocarpus amarus Reinw., Syll. Ratisb. 2 (1826) 14; Cardiophora
hindsii Benth. & Hook. f., Lond. J. Bot. 2 (1843) 216.
Shrub or small tree to 5(15) m tall; young shoots and buds rusty tomentose. Leaves
crowded at the apex of the branchlets, on dropping leaving large scars; blade obovateoblong, 1035 x 412 cm; base cuneate, apex blunt but sometimes mucronate; midrib and
veins hairy below; midrib slightly immersed or inconspicuous above, strongly prominent
beneath; lateral veins straight, parallel, ending in an intramarginal looped vein, sulcate,
slightly prominent or inconspicuous above; intercostal veins finely dense-reticulate beneath;
stalks pithy, shrunken at the base when dry, sometimes also at apex, hairy, 38 cm long.
Inflorescences erect, shorter than the leaves, 312 cm long. Flowers c. 2 mm long; pedicels
to 5 mm long; sepals puberulous, erect, appressed, 0.51.0 mm long; petals concave,
spreading, finally reflexed, sparsely hairy to glabrous, accrescent, to 2.5 x 1 mm; stamens
with glabrous filaments to 1 mm long, anthers c. 0.75 mm long; ovary 23-carpellate, never
with more than 2 carpels fertile, carpels connate except at the top. Fruits obcordate, to 2 x
2.5 cm, strongly emarginate; pericarp hard and corky; wings often nearly touching near the
inward curved style-bases. Seeds round, 0.51 cm across.
Distribution. From Borneo eastwards to Micronesia (West and East Carolines and
Marshalls) and Melanesia (New Britain, Solomons, New Hebrides); in Malesia: Borneo
(Sabah, Sarawak, Karimata Is.), Moluccas, and New Guinea.
Ecology. A typical constituent of the Barringtonia-formation of the beach vegetation, but
much rarer than most of the species belonging to that formation, though locally common on
the sandy beaches and behind coral reefs.
Uses. None known in Sabah and Sarawak. The roots and the fruits of this very bitter plant
have been used to treat cholera, pleurisy, and other fevers; a beverage prepared from
powdered leaves is taken against colic and cough, and the fruits have been used to induce
vomitting in treating snake bites (Nooteboom l.c.).
442
SONNERATIACEAE
Othman Bojo
Universiti Malaysia Sarawak,
Kota Samarahan, Malaysia
Merrill, EB (1921) 418; Holthuis & Lam, Blumea 1, 5 (1942) 216; Backer & van Steenis, FM 1, 4
(1951) 280, FM 1, 5 (1958) 557, FM 1, 6 (1972) 973; Whitmore, TFM 1 (1973) 442; Cockburn, TS 1
(1976) 223; Anderson, CLTS (1980) 323; Ashton, MNDTS 2 (1988) 374; Whitmore, Tantra &
Sutisna, CLK 2, 1 (1990) 331.
Trees. Bark cream to grey, smooth or shallowly irregularly flaky; inner bark soft, brown.
Sapwood soft, pale yellow. Leaves simple, opposite, biseriate, entire, coriaceous, shortstalked, without stipules. Inflorescences terminal corymbs or 13-flowered fascicles.
Flowers bisexual, pedicelled, rather large, regular; sepals persistent, thickly leathery,
connate at base to form a tube or cup with 48 triangular lobes, segments valvate in bud,
often reddish inside; petals absent or as many as sepals, broad and wrinkled or very narrow
and smooth, alternating with the sepals; stamens 12 to many, inserted on the sepals,
1many seriate, inflexed in bud, filamens filiform to subulate; anthers 2-celled, kidneyshaped or oblong, opening lengthwise; ovary superior, sessile and with a broad base,
enclosed by the calyx-base until it expands, 420-celled, septa thin; style 1, long, stout,
stigma 1, capitate or slightly lobed; ovules anatropous many, placentation axile. Fruits
many-seeded berries (in Sonneratia) or dehiscent capsules (in Duabanga), subtended by the
persistent calyx-tube. Seeds small, without endosperm; embryo straight.
Distribution. 2 genera with 7 species distributed in the Old World Tropics from East Africa
to the Pacific islands. In Sabah and Sarawak, 2 genera with 4 species.
Ecology. Whereas Sonneratia is a mangrove genus found mainly on sandy and muddy tidal
flats, estuaries, brackish streams and coral terraces at tide level, Duabanga is a component
of lowland and hill forests, and being light-demanding, is frequently found in secondary
forests, forest-edges and river-banks. The flowers produce a mild sour to musty odour,
expand at sunset, last for only one night and are pollinated by nectarivorous bats. The
fruits/seeds are dispersed either by water (Sonneratia) or by wind (Duabanga).
Uses. The soft pale wood of the Sonneratiaceae is not durable and its only known use is as
firewood. However, timber of Duabanga is occasionally used for boat-building. Fruits of
some species (S. caseolaris and S. ovata) are edible, and the leaves and pneumatophores of
S. caseolaris are reported to have medicinal value.
Taxonomy. Previous authors such as Ridley (FMP 1 (1922) 819), Watson (MFR 6 (1928)
50), Browne (FTSB (1955) 248), Hsuan Keng (OFMSP (1978) 155), Corner (WSTM 1
(1988) 470) and Brummitt (Vasc. Pl. Fam. & Gen.Dicot. (1992) 607) included Duabanga
443
444
SONNERATIACEAE (OTHMAN)
and Sonneratia in the Lythraceae. This contention seems to be supported by anatomical

(Metcalfe & Chalk, Anat. Dicot. 1 (1964) 6), embryological (Venkateswarlu, Proc. Indian
Acad. Sci. Ser. B, 5 (1937) 206), and palynological evidence (Erdtman, Poll. Morph. & Pl.
Tax. 1 (1952) 412; Thanikaimoni & Jayaweera, Tran. Sect. Sci. & Techn., Inst. Pranc.
Pondichery 5, 2 (1966) 1; Muller, Pollen et Spores 2 (1969) 223; Germeraad, Hopping &
Muller, Rev. Palaeobot. & Palyn. 6 (1968) 189). However, accounts by Backer & van
Steenis (l.c.), Cockburn (l.c.), Ashton (l.c.), Mabberley (PB (1987) 546), and van Steenis
(Checkl. Gen. Names Males. Bot., Sperm. (1987) 115) include the two genera in the
Sonneratiaceae, a view which is adopted here. For anatomical information on the family, the
work of Reinders-Gouwentak (FM 1, 4 (1951) 513) should be consulted.
Key to genera
Monopodial trees of inland forests, without breathing roots; branches drooping. Leaves
coriaceous, glaucous below, with many prominent lateral veins and intra-marginal veins,
base heart-shaped. Flowers in terminal corymbs; petals broad, wrinkled. Fruits 48-valved
capsules. Seeds tailed at each end by the protracted testa..................................1. Duabanga
Sympodial trees of mangroves and sea-coasts, with breathing roots; branches not drooping.
Leaves succulent, not glaucous, venation obscure; base not heart-shaped. Flowers in clusters
of 13 at the ends of the branchlets; petals absent or very narrow. Fruits indehiscent berries.
Seeds not tailed................................................................2. Sonneratia
1. DUABANGA Buch.-Ham.
(from duyabangga, an Indian vernacular name)
Trans. Linn. Soc. Lond. 17 (1893) 177; Ridley, FMP 1 (1922) 824 (under Lythraceae); Burkill, EPMP
(1935) 868; Backer & van Steenis l.c. 288; Jayaweera, J. Arn. Arb. 48 (1967) 89; Ashton l.c. 375;
Kochummen & Wyatt-Smith, MFR 17 (1964) 271; Cockburn l.c. 223; Corner, WSTM (1988) 476
(under Lythraceae); Whitmore, Tantra & Sutisna l.c. 331.
Medium-sized to big trees; crown monopodial, rather open, ultimate branches drooping;
bole with insignificant concave buttresses at base. Bark smooth, pale ochreous-cream or
grey; inner bark pale yellow, soft, fibrous. Sapwood white, soft. Twigs 4-angled or winged,
often with an interpetiolar ridge. Leaves oblong-ovate, base heart-shaped, glaucous
beneath, coriaceous; lateral veins prominent beneath, looped at the margin forming intramarginal veins; petioles short. Flowers 5many, in terminal corymbs, 48-merous; calyx
fleshy, green, cup-shaped with triangular segments; petals shortly clawed, white or
yellowish, broad, wrinkled; stamens 12 or many, 1-seriate; anther recurved over one end of
the connective; ovary 48-celled; stigma stout, lobed. Fruits 48-valved, loculicidally
dehiscent. Seeds tailed at both ends by the extended testa.
Distribution. 2 species, from eastern Himalaya to New Guinea. Only 1 species (D.
moluccana) is found in Sabah and Sarawak and is common.
445
Ecology. Mainly in mixed evergreen and deciduous forests at 151200 m. Usually

occurring as scattered individuals on brownish yellow sandy clay loam soils or yellow
podsols. The seeds are readily dispersed by wind.
Taxonomy. Based on the revision of Sonneratiaceae by Backer & van Steenis (l.c. 288),
Duabanga consists of two species, D. grandiflora and D. moluccana. In 1967, Jayaweera
(l.c. 89), added one new species, D. taylorii Jayaweera. However, detailed study of D.
taylorii by Geesink (Blumea 2, 18 (1970) 454) reveals that this species is a primary hybrid
between D. grandiflora and D. moluccana which is believed to have originated in the Bogor
Botanic Garden in the 1850s.
Uses. In Sabah and Sarawak, its non-durable and soft wood has no commercial value.
However, in other regions, the wood is used for house and boat-building and also for
firewood.
Duabanga moluccana Blume
Fig. 1.
(of the Moluccas islands)

Mus. Bot. Lugd. Bat. 1 (1849) 109; Merrill, PEB (1929) 212; Holthuis & Lam l.c. 216; Backer & van
Steenis l.c. 288; Jayaweera l.c. 91; Cockburn l.c. 223; Anderson l.c. 323; Ashton l.c. 375; Corner l.c.
476; Whitmore, Tantra & Sutisna l.c. 331. Type: Forstein, s.n., Moluccas, Ambon (holotype L).
Synonym: D. borneensis R. Knuth, Fedde Rep. 38 (1935) 121.
Tree to 45 m tall and 100 cm diameter, with or without buttresses; trunk columnar. Bark
smooth becoming fissured or scaly with age, lenticellate, grey or brownish; inner bark
yellowish or pale brown, soft and fibrous. Sapwood cream, darkening purplish on exposure.
Twigs 38 mm diameter, slender, 4-angled, becoming terete with age, hairy when young,
glabrescent. Leaves ovate, oblong or oblanceolate, 1026 x 413 cm, dark green above,
paler beneath, leathery; base shallowly heart-shaped, apex acuminate; midrib prominent
beneath; lateral veins 1126 pairs, joined near the margin to form an intra-marginal vein;
petioles 510 mm long. Inflorescences few to many-flowered, dense or rather lax,
pubescent at first, becoming glabrous. Flower-buds 2 x 1.5 cm, ovoid, ribbed, acute.
Flowers 4-merous, with musty odour; calyx-tube c. 2 cm across at anthesis, lobes 4, c. 12
cm deep; petals 4, c. 2 x 1.2 cm, yellowish or cream, falling off early; stamens 12, anthers
yellow turning to pale brown; style c. 40 mm long, dark red, stigma dark green. Fruits
ovoid or oblong, 4-valved, 1.53 x 12 cm. Seeds c. 0.7 mm long.
Vernacular names. Sabahmagas (Kadazan). Sarawakbenung kasung (Bidayuh), sawih
(Iban).
Distribution. Java, Borneo (Sabah, Sarawak, Brunei, Kalimantan), Celebes, Moluccas, and
New Guinea.
Ecology. In Sabah and Sarawak, widely distributed at 15750 m, on damp clay-rich fertile
soils, especially in high light intensity areas such as river-banks, forest-edges, logged-over
forests, road-sides, abandoned cultivation sites and also on limestone hills.
Uses. The timber is little used at present. Recorded uses include temporary light
construction, floaters for extracting timber logs, fish-net floats and dug-out boats. It grows
446
fast and may have potential to be planted for pulping materials or for core-stock in plywood
manufacturing.
2. SONNERATIA L. f., nom. cons.

(Pierre Sonnerat, 18th century French explorer-naturalist)
Suppl. Pl. Syst. Veg. (1781) 38; Merril l.c. (1921) 416; Ridley l.c. 825; Watson l.c. 121; Burkill l.c.
2051; Backer & van Steenis l.c. 282; Wyatt-Smith, MF 16 (1953); Kochummen & Wyatt-Smith l.c.
315; Whitmore l.c. 444; Cockburn l.c. 224; Anderson l.c. 323; Duke & Jackes, Blumea 32 (1987) 277;
Ashton l.c. 375; Corner l.c. 476 (under Lythraceae); Whitmore, Tantra & Sutisna l.c. 331. Synonyms:
Blatti Adans., Fam. 2 (1763) 88; Pagapate Sonnerat, Voy. Nouv. Guin. (1776) 16.
Small to medium-sized glabrous trees, with sympodial crown and low branching; bole fluted
at base, surrounded by spindle-shaped breathing roots (pneumatophores), buttresses
absent. Bark smooth, becoming scaly or finely fissued with age, lenticellate, pale greygreen to dark brownish grey; inner bark pinkish or brownish, rather watery. Young twigs
(branchlets) drooping or erect, distinctly jointed above the nodes, 4-angled, becoming terete
with age. Flowers in terminal clusters of 13, 48-merous; calyx cup-shaped, 48-lobed,
segments ovate or triangular, often reddish within; petals absent or 48, narrow, and
falling off early; stamens many, falling off early; ovary 10- to many-celled; style slender,
sinuous in bud. Fruits indehiscent berries, depressed-globose, tipped by the persistent style
and seated on the persistent star-like calyx-cup, pericarp leathery. Seeds many, embedded in
foul-smelling pulp, not tailed at both ends; embryo straight.
Distribution. 5 species distributed along sea-shores of tropical East Africa, Asia, Malesia,
Australia and Pacific Islands. 2 species occur in Sabah and three in Sarawak.
Ecology. Mangrove trees, found on sandy and muddy tidal flats, estuaries, brackish watercourses, river-banks and creeks and coral-terraces at tide level. The flowers are ephemeral
and pollinated by nectar-feeding bats and birds. The fruits are water-dispersed. Hybrids
have been recorded between S. alba x S. caseolaris and S. alba x S. ovata along the Brunei
coasts, and their status has been substantiated by palynological (Muller, Ancient Pacific
Floras (1964) 33), morphological and cytological (Muller & Hou-Liu, Blumea 14 (1966)
337) evidence.
Uses. The timber is classified as moderately hard to hard and moderately heavy to heavy.
However, it is only locally used for firewood and occasionally for house-posts. Burgess
(TBS (1966) 461) claimed that the wood of S. caseolaris could be pulped by the sulphate
process to give a pulp with similar strength properties to commercial eucalypt pulp. The
fruits of some species are edible. In Sarawak, fruits of S. caseolaris and S. ovata are edible.
Key to Sonneratia species

1.
Leaves elliptic to elliptic-ovate. Young twigs drooping. Petals linear-lanceolate, dark

red; filaments bright pink at base, upper part white................................2. S. caseolaris
Leaves obovate, broadly ovate to suborbicular. Young twigs erect. Petals (if present)
linear, white; filaments white ......................................................................................2
447
448
2.
Leaves obovate, base cuneate. Petals linear, white...........................................1. S. alba

Leaves broadly ovate to suborbicular, base rounded or subcordate. Petals absent............
.....................................................................................................................3. S. ovata
1. Sonneratia alba J. Smith

(Latin, albus = white; the flowers)
in Rees, Cycl. 33 (1819) 2; Beccari, Nelle For. di Borneo (1902) 579; Merrill, Interp. Rumph. Herb.
Amb. (1917) 383, l.c. (1921) 418; Backer & van Steenis l.c. 285; Wyatt-Smith l.c. 214; Browne l.c.
323; Duke & Jackes l.c. 286; Ashton l.c. 379; Corner l.c. 476; Whitmore, Tantra & Sutisna l.c. 331.
Type: Rumph. Herb. Amb. 3 (1754) 111, t. 73, Ambon. Synonyms: Rhizophora caseolaris L. in
Stickman, Herb. Amb. (1754) 13, in part; Blatti alba (J. Smith) Kuntze, Rev. Gen. 1 (1891) 238.
Tree to 40 m tall and 70 cm diameter; stems usually crooked; crown rather open, more or
less broadly oblong, low-branching. Young twigs erect. Bark slightly cracked to fissured,
grey to brownish above normal tide mark, lenticels usually present; inner bark not
laminated, mottled cream and pinkish brown. Leaves fleshy, broadly obovate to rarely
ovate, 3.510 x 27.5 cm; base cuneate, apex broadly rounded; lateral veins 79 pairs, thin,
inconspicuous; petioles 713 mm, stout. Flower-buds oblong, 22.5 x 1.5 cm. Flowers
solitary or in terminal clusters of 23, 67(8)-merous; calyx 1.52.5 cm long, cup
obconical, ribbed or angular, angles as many as segments and alternating with them,
segments 67, ovate-oblong, 1.32 cm long, inner surface red, especially at base; petals
67, linear, inconspicous and almost resemble the filaments, white, pink at base; filaments
white; ovary 1418-celled. Fruits 45 cm long, 34 cm across, dark green, subtended by
the ribbed calyx-cup.
Vernacular names. Sabahpedada (Malay, Kadazan). Sarawakperepat (Malay, Iban).
Distribution. Tropical East Africa, N Madagascar, Seychelles, Sri Lanka, Andamans,
continental SE Asia, throughout Malesia to Australia and the W Pacific Islands. In Sabah, a
common component of mangroves in Kudat, Sandakan, Lahat Datu and Semporna. In
Sarawak, very common in Buntal Bay from Santubong to Bako National Park and elsewhere
such as the Lawas mangroves. Also in Brunei and Kalimantan.
Ecology. A pioneer mangrove species found on muddy and shallow parts of calm sea-shores
and along the mouths of tidal creeks, forming dense pure stands on intertidal sites. Rarely
found far from river mouths and sandy beaches.
Uses. In Sabah and Sarawak, the wood is used as firewood. Elsewhere, such as in Minahasa
(NE Celebes), the wood is valued for ship and house-building.
2. Sonneratia caseolaris (L.) Engl.

(Latin, caseus = cheese; the fruit which resembles a small, depressed rounded piece of
cheese)
in Engler & Prantl, Nachtr. (1897) 261; Merrill l.c. (1917) 383, l.c. (1921) 418; Backer & van Steenis
l.c. 283; Wyatt-Smith l.c. 213; Browne l.c. 248; Whitmore l.c. 445; Cockburn l.c. 225; Anderson l.c.
323; Duke & Jackes l.c. 289; Ashton l.c. 379; Corner l.c. 476; Whitmore, Tantra & Sutisna l.c. 331.
Basionym: Rhizophora caseolaris L., p.p. in Stickman, Herb. Amb. (1754) 13. Type: Rumph. Herb.
449
Amb. 3 (1754) 112, t. 7475, Ambon. Synonyms: S. acida L., Suppl. 252; Aubletia caseolaris (L.)
Gaertn., Fruct. (1788) 379; Blatti caseolaris (L.) Kuntze, l.c. 238; Blatti acide (L.) Lam., Enc. 1
(1789) 429; S. pagatpat Blanco, Fl. Filip. (1837) 424; Blatti pagatpat (Blanco) Niedenzu in Engler &
Prantl, Pfl. Fam. 3, 7 (1891) 21; S. ovalis Korth., Ned. Kruidk. Arch. 1 (1846) 198; S. evenia Blume,
Mus. Bot. Ind. Bat. 1 (1851) 337; S. lanceolata Blume l.c. (1851) 337; S. obovata Blume l.c. (1851)
337; S. neglecta Blume l.c. (1851) 338.
Tree to 20 m tall and 30 cm diameter; stems frequently leaning and crooked; crown rather
open. Bark smooth, greenish grey and lenticellate when young, becoming irregularly
fissured and pale brown with age; inner bark cream or pinkish brown, faintly laminated;
exudate watery. Young twigs drooping, yellowish brown turning to greenish brown with
age. Leaves elliptic to elliptic-ovate, 3.89.5 x 28 cm; base cuneate, apex blunt or
subacuminate; lateral veins 812 pairs, thin and inconspicuous; petiole 711 mm long,
reddish. Flower buds broadly ovoid or oblong to subglobose, 22.5 x 11.5 cm. Flowers
68-merous; calyx at anthesis shallowly cup-shaped, smooth, segments 68, usually longer
than the tube, cream or pale yellow inside, base crimson, forming a crimson ring around the
fruit; petals linear-lanceolate, 68, 2025 x 1.52 mm, dark red; filaments white, bright
pink at base; ovary 1621-celled. Fruits c. 4 cm long, 7.5 cm across, dark green, crowned
with persistent conically thickened style base, subtended by the flattened, green calyx-tube
with horizontally spreading lobes.
Vernacular names. Sabahperepat (Malay, Kadazan). Sarawakpedada (Iban, Malay).
Distribution. Sri Lanka, tropical SE Asia to New Guinea, Northern Australia, the Solomon
Island, and New Hebrides. In Sabah and Kalimantan, very common in mangrove front. In
Sarawak and Brunei, common along tidal river-banks and mangroves, especially on deep
muddy river-banks, and from the limit of brackish water down to the lower end of the Nypazone near the river-mouths.
Ecology. Individual trees of S. caseolaris are usually found scattered among other
mangrove species (e.g., Avicennia species) rather than forming a pure stand.
Uses. The young, sour-tasting berries are edible and are also used in traditional medicine,
chiefly externally but also internally (Burkill l.c. 2088). Burkill & Haniff (Gard. Bull. S.S. 6
(1930) 204) reported that leaves pounded with rice were applied for small-pox. Pectin is
locally extracted from the fruits. The wood is used as fuel, and the pneumatophores, after
boiling in water, furnish an inferior substitute for cork.
3. Sonneratia ovata Backer
Fig. 2.
(Latin, ovatus = oval; the leaf shape)

Bull. Jard. Bot. Btzg. 3, 2 (1920) 329; van Steenis, Bull. Jard. Bot. Btzg. 3, 12 (1931) 162; Backer &
van Steenis l.c. 162; Wyatt-Smith l.c. 214; Whitmore l.c. 445; Anderson l.c. 323; Duke & Jackes l.c.
299; Ashton l.c. 380; Corner l.c. 476; Whitmore, Tantra & Sutisna l.c. 331. Type: Backer 21422, Java
(lectotype L).
Tree to 20 m tall and 20 cm diameter; stems short and usually twisted. Bark grey, smooth to
slightly fissured, lenticellate; inner bark pale brown to reddish, faintly laminated. Young
twigs erect, greyish brown. Leaves usually ovate to orbicular or broadly ovate, 2.57.5 x
450
39 cm; base rounded or subcordate, apex broadly rounded; adaxial surface slightly
corrugated by the fine but conspicuous 910 pairs of lateral veins; petioles 411 mm long.
Flowers usually in terminal clusters of 23(4), occasionally solitary; buds broadly ovoid,
apex rounded or obtuse, finely verruculose, 1.52 x 11.5 cm; calyx cup-shaped, tapering
abruptly into a stalk-like base, ribbed, segments usually 6, ovate-triangular, 1315 mm
long, inner surface tinged reddish at base; petals absent; filaments white; ovary 1315celled. Fruits c. 4 cm long, 7 cm across, dark green when young, turning to yellowish green
when ripe.
Vernacular name. Sarawakrogam (Malay).
Distribution. China, Thailand and throughout Malesia. In Sarawak, Brunei and Kalimantan
very local. In Sabah, has not been recorded with certainty.
Ecology. Occasionally found on banks of tidal rivers and muddy soils inundated only by
spring tides.
Uses. In Sarawak, the mature fruit is edible and eaten as ulam (a salad) by the Malays.
451
STAPHYLEACEAE
J. T. Pereira
Sandakan, Malaysia
Merrill, EB (1921) 355; Ridley, FMP 1 (1922) 511; van Steenis, Nova Guinea 10,2 (1959) 211; van
der Linden, FM 1, 6 (1960) 49; Airy-Shaw, Kew Bull. 18, 2 (1965) 252; Backer & Bakhuizen f., FJ 2
(1965) 145; Sponberg, Journ. Arn. Arb. 52 (1971) 196; Whitmore, TFM 1 (1972) 446; Anderson,
CLTS (1980) 324; Dickison, Journ. Arn. Arb. (1984) 149; Whitmore, Tantra & Sutisna, CLK 2, 1
(1990) 332; Webster, Ann. Missouri Bot. Gard. 81 (1994) 54.
Trees or shrubs, non-laticiferous. Leaves simple to compound (imparipinnate to paripinnate), oppposite or alternate, leaflet margin toothed; stipules usually falling off early.
Inflorescences terminal, subterminal or axillary panicles or racemes. Flowers bisexual,
sometimes unisexual, actinomorphic (with regular symmetry); sepals 5, imbricate; petals 5,
imbricate; stamens 5, alternating with the petals outside the disc; anthers 2-celled,
longitudinally dehiscent, dorsifixed; disc present or absent; ovary superior, 24-celled,
styles 24, free or fused together, stigmas capitate or coherent, forming a peltate structure;
ovules one or more per cell, placentation axile or basal. Fruit an indehiscent berry, follicle
or inflated capsule. Seeds one to several, arillate or exarillate; embryo straight; cotyledons
flat; endosperm fleshy or horny.
Distribution. 5 genera with about 60 species. Staphylea (11 species), Tapiscia (1 species)
and Euscaphis (1 species) are found in the north temperate region. Huertea (4 species)
occurs from West Indies to Peru. Turpinia has 3040 species distributed from the Indomalesian region to Japan and tropical America. In Sabah and Sarawak, Turpinia is
represented by 6 species all of which occur in Sabah, although only 2 species are found in
Sarawak.
Uses. Some species of the temperate Staphylea and Euscaphis are cultivated as ornamental
plants. Timbers of a few species of Turpinia are occasionally utilised locally. The fruit of
Euscaphis japonica, a common tree or shrub in Japan and Central China, is used for
preparing a drug.
Taxonomy. In early publications, the family was considered closely affiliated to the
Rhamnaceae. Later authors (cf. van Steenis l.c. 211, van der Linden l.c. 49, Spongberg l.c.
196) placed it near to the Aceraceae, Celastraceae, Cunoniaceae, and Sapindaceae. Its
affinity to the Cunoniaceae was based on the evidence derived from morphology, anatomy
and embryology (Hallier, 1908; Whitmore l.c. 446; Thorne, 1976; Dickison l.c. 149).
Anatomically, there are major differences between the Staphyleaceae and the Sapindaceae
(Metcalfe & Chalk, Anat. Dicot. 1 (1957) 443) although the Staphyleaceae have some
similarities to the woody Saxifragaceae in terms of nodal and leaf anatomy (Dickinson, Bot.
453
Gaz. 148,4 (1987) 475). Whitmore (l.c. 446) included Bischofia under the Staphyleaceae,
which had been initially placed in the Euphorbiaceae by Decaisne (1842) and later isolated
as a separate family, the Bischofiaceae, by Airy-Shaw (1965). However, evidence from
embryology and anatomy appears to support its inclusion in the Euphorbiaceae (Webster,
1994). The Indochinese Tricaphis was also included in the Staphyleaceae by Gagnepain
(Not. Syst. 13 (1948) 190) but van Steenis (1960) excluded it, and suggested that it
belonged to either the Anacardiaceae or Sapindaceae. Hence, the taxonomic status of this
family, particularly the position of certain genera, remains controversial.
TURPINIA Vent., nom. cons.

(P.J.Fr. Turpin, 17751840, an eminent French botanical artist)
Choix (1803) 31, t. 31; Ridley l.c. (1922) 511; Merrill & Perry, Journ. Arn. Arb. 22 (1941) 543;
Metcalfe & Chalk l.c. 443; van Steenis l.c. 211; van der Linden l.c. 49; Backer & Bakhuizen f. l.c.
145; Burkill, EPMP 2 (1966) 2234; Grey-Wilson, Kew Bull. 26, 1 (1972) 141; Whitmore l.c. 446;
Anderson l.c. 324; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 332; Pereira, Sandakania 5 (1994)
15. Synonyms: Triceros (non Griff.) Lour., Fl. Coch. 1 (1790) 184; Dalrympelea Roxb., Pl. Corom. 3
(1820) 76, t. 279; Ochranthe Lindl., Bot. Reg. 8 (1836) t. 1819; Hasskarlia Meisn., Pl. Vasc. Gen. 2
(1843) 348; Kaernbachia (non O. Kuntz.) Schltr., Bot. Jahrb. 52 (1914) 151.
Trees or shrubs. Twigs smooth, terete, pithy, glabrous or hairy. Stipules interpetiolar, entire
or bifid at apex, falling off early (except in T. stipulacea), leaving prominent scars. Leaves
opposite-decussate, unifoliolate, trifoliolate to pinnate (imparipinnate or paripinnate),
petioles wrinkled at base; glands present at base of leaflets and at the insertion of petiolules
to the rachis; leaflets chartaceous to coriaceous, margin serrate to crenate, midrib prominent.
Inflorescences axillary, terminal to subterminal panicles or racemes. Flowers bisexual;
outer sepals broader than the inner ones, ovate, margin ciliate; petals of equal size, longer
than sepals, margin ciliate; stamens of equal size, broad at the base, glabrous, anthers oval,
apex sometimes apiculate; disc lobed, fleshy; ovary superior, (2)3(4)-celled; styles closely
adnate together forming a peltate stigma; ovules one to numerous in each cell, in axile
plancentation in 2 rows. Fruit a globose to subglobose or trilobed, indehiscent, fleshy berry,
typically crowned by three radial lines or horn-like structures (remains of the styles) on top;
fruit wall thin to thick, fleshy. Seeds mostly angular, one to a few in each cell, pale to dark
brown; cotyledon flat, roundish.
Distribution. About 3040 species from Sri Lanka to southern Japan, southward and
eastward to Malesia, west Pacific and the central and south American tropics (West Indies,
Columbia, Ecuador, Peru). In Sabah and Sarawak, the genus is represented by 6 species, 2
of which are endemic to Sabah, viz., Turpinia nitida and Turpinia stipulacea.
Ecology. Evergreen understorey trees or shrubs of lowland tropical to subtropical rain
forests to montane zones reaching c. 2400 m in New Guinea and Mt. Kinabalu and to 3000
m in the Himalayas and Yunnan. Fruits are probably dispersed by birds and other animals
and the hard-coated seeds can probably withstand acidity and digestion in intestines during
dispersal by animals.
454
STAPHYLEACEAE (PEREIRA)
Uses. Turpinia species produce timber of a low durability, hence usage in construction is
exceptional. The wood is used for packaging cases in Indonesia (Burkill l.c. 2234). Turpinia
nepalensis, common in western China, produces a useful tough wood. Most Turpinia
species are grown as pioneer trees for reafforestation due to their rapid growth on eroded
mountain slopes in central Java.
Key to Turpinia species

(based mainly on leaf characters)
1.
Stipules 1425 mm long, conspicuous and persistent on apices of leafy twigs................

.............................................................................................................6. T. stipulacea
Stipules not exceeding 10 mm in length, not very conspicuous, falling off early from
leafy twigs...................................................................................................................2
2.
Lower surface of leaflets sparsely to densely short-hairy on the midrib, lateral veins and
blade..................................................................................................3. T. grandis
Lower surface of leaflets glabrous................................................................................3
3.
Intercostal veins pseudoscalariform (with many almost straight cross-veins between the
lateral veins).....................................................................5. T. sphaerocarpa (in part)
Intercostal veins reticulate (with wide areoles, or obscurely or distinctly tessellate, with
no or very few straight cross-veins between the lateral veins).......................................4
4.
Leaflets thickly coriaceous, margin strongly recurved when dry...................4. T. nitida

Leaflets chartaceous or coriaceous, margins not strongly recurved when dry................5
5.
Leaflets narrowly acuminate to caudate, acumen usually 12 cm.........1. T. borneensis

Leaflets shortly acuminate to cuspidate, acumen to c. 1 cm long, often shorter.............6
6.
Leaves typically 2046 cm long (rachis 1018 cm long)....5. T. sphaerocarpa (in part)
Leaves often less than 20 cm long (rachis less than 10 cm long)..................................7
7.
Stipules 2.53.5 x 22.5 mm. Leaflets coriaceous and slightly shiny on lower or both
surfaces. Fruit globose to subglobose, without short apical horns, but with radial lines at
the apex. (Plants restricted to limestone habitats.)..............................2. T. calciphila
Stipules 57 x 34 mm. Leaflets chartaceous and not shiny. Fruit slightly trilobed at the
apex, with short apical horns. (Plant of non-limestone habitats.)..............................
.........................................................................................5. T. sphaerocarpa (in part)
Key to Turpinia species

(based mainly on fruits and flowers)
1.
Fruits slightly to distinctly trilobed at the apex, typically with short apical horns.........2
Fruits globose to subglobose, without short apical horns..............................................3
2.
Petals 24 x 1.52 mm; anthers 0.81 mm long; pistils 2.53.5 mm long. Seeds 58
mm across. Leaflets oblong, elliptic-oblong to lanceolate or ovate, typically shiny
above, coriaceous, margin strongly recurved................................................4. T. nitida
455
Petals 1.82 x 11.1 mm; anthers 0.5 mm long; pistils 22.2 mm long. Seeds 24 mm
across. Leaflets ovate-elliptic to elliptic, not shiny above, chartaceous, margin not
recurved.................................................................................5. T. sphaerocarpa (in part)
3.
Fruit-wall brittle, 0.10.3 mm thick (Plants restricted to limestone habitats)..................

............................................................................................................2.. T. calciphila
Fruit-wall not brittle, thicker than 0.5 mm (Plants not restricted to limestone
habitats).......................................................................................................................4
4.
Ovules (2)34 per cell...............................................................................................5

Ovules (4)68(9) per cell.........................................................................................6
5.
Petals 2.54.5 mm long; stamens 23 mm long; pistils 23 mm long, ovary 11.5 mm

across. Stipules 1425 mm long, conspicuous and persistent on apices of leafy
twigs.....................................................................................................6. T. stipulacea
Petals 1.22 mm long; stamens 0.81.8 mm long; pistils 12 mm long, ovary 0.61 mm
across. Stipules 38 mm long, not very conspicuous, falling off early from leafy
twigs....................................................................................................1. T. borneensis
6.
Seeds 514 per fruit. Lower surface of leaflets sparsely to densely short-hairy on the
midrib, lateral veins and blade. Leaf rachises 38 cm long............................................
.................................................................................................................3. T. grandis
Seeds 17 per fruit. Lower surface of leaflets completely glabrous. Leaf rachises 1018
cm long.............................................................................5. T. sphaerocarpa (in part)
1. Turpinia borneensis (Merr. & Perry) B.L. Linden

(of Borneo)
l.c. 49; Anderson l.c. 324; Whitmore, Tantra & Sutisna l.c. 332. Basionym: Turpinia montana var.
borneensis Merr. & Perry l.c. 549. Type: J. & M. S. Clemens 29391 bis, British North Borneo,
Kinabalu, Tenompok (holotype UC; isotypes BO, K).
Tree to 20 m tall and 20 cm diameter. Bark shallowly fissured, pale brown; inner bark
greyish. Sapwood brownish. Stipules 38 x 24 mm, not very conspicuous, falling off early
from leafy twigs, glabrous, margin sparsely hairy to entire. Leaves 35(7)-foliolate, (13)
1635(40) cm long; petioles 311 cm long, 12 mm thick, glabrous; leaf-rachis 4.513 cm
long, glabrous; petiolules of lateral leaflets 215 mm long, glabrous; leaflets drying pale
green above, chartaceous, glabrous, not shiny, ovate-elliptic to lanceolate, 518 x 27 cm;
base cuneate, sometimes rounded or oblique, margin finely serrate, not recurved, apex
narrowly acuminate to caudate, acumen (0.5)12 cm long; lateral veins 711 pairs,
distinctly looping at the margin, prominent on lower surface; intercostal veins reticulate or
distinctly tessellate and prominent, with no or very few straight cross-veins between the
lateral veins on both surfaces. Inflorescences axillary panicles, to 30 cm long, rachis
subglabrous. Flowers cream to pale yellow, sepals ovate, thin, 1.22 x 11.8 mm; petals
obovate, 1.22 x 0.81.5 mm; stamens 0.81.8 mm long, glabrous, filaments 0.51.5 mm
long, glabrous, anthers 0.40.5 mm long, apiculate or not; pistils 12 mm long, styles 0.6
1.8 mm long; ovary 0.61 mm across, 3(4)-celled, each cell with 34 ovules. Fruits
globose, 818 mm diameter, without short apical horns but with three radial lines at the
456
apex, wrinkled; wall thin, not brittle, 0.51 mm thick. Seeds pale to dark brown, 410 x 6
10 mm, 14 in each fruit.
Distribution. Borneo (Kalimantan and Sabah) and the Philippines; not yet recorded in
Sarawak.
Ecology. Primary and secondary mixed dipterocarp forest to 1800 m. Flowering has been
recorded from November to March whereas fruiting has been noted from May to October.
2. Turpinia calciphila J. T. Pereira

(Latin, calx = lime; Greek, philos = loving; referring to the habitat)
Sandakania 5 (1994) 18. Type: Anderson S. 4710, Sarawak, Baram, Gunung Api (holotype SAR;
isotypes BO, K, L, SING).
Tree to 25 m tall and 50 cm diameter. Bark creamy yellow, slightly flaky. Stipules 2.53.5 x
22.5 mm, not very conspicuous, falling off early from leafy twigs, glabrous, margin hairy.
Leaves 13-foliolate, 916(25) cm long, petioles 1.54(7.5) cm long, 12 mm thick,
glabrous; leaf-rachis 1.54 cm long, glabrous; petiolules of the lateral leaflets 514 mm
long, glabrous; leaflets drying pale green to brown above, coriaceous, glabrous, slightly
shiny on lower or both surfaces, ovate to ovate-elliptic, 6.515 x 3.56.5 cm; base rounded
to slightly cuneate, sometimes oblique, margin shallowly serrate, sometimes slightly
undulate, not recurved, apex shortly acuminate, acumen 0.41 cm long; midrib and lateral
veins raised and distinct on lower surface; lateral veins 68 pairs, ascending, parallel, open
or looping towards the margin; intercostal veins reticulate or distinctly tessellate beneath,
with no or very few straight cross-veins between the lateral veins. Inflorescences axillary
panicles to 25 cm long, rachis densely hairy at the distal end. Flowers not known in mature
state. Fruits globose to subglobose, 510 mm diameter, without short apical horns but with
three radial lines at the apex, smooth; wall 0.10.3 mm thick, brittle. Seeds dark brown to
reddish brown, 48 x 58 mm, 16 in each fruit.
Vernacular name. Sarawaklaba (Iban).
Distribution. Endemic to Sarawak (4th Division, Baram, Gunung Api and Gunung Buda;
1st Division, near Bau, Bt. Buan and Bt. Gebung).
Ecology. So far only known from the lower, mesic slopes of limestone screes, to 900 m.
3. Turpinia grandis B.L. Linden

(Latin, grandis = large, big; the leaves)
l.c. 49; Whitmore, Tantra & Sutisna l.c. 332. Type: Endert 4669, E Borneo, W Koetai, Kiau River,
700 m alt. (holotype L; isotypes BO, K).
Tree to 20 m tall and 30 cm diameter. Bark smooth, greyish; inner bark yellowish.
Sapwood pale white. Stipules 4 x 35 mm, not very conspicuous, falling off early from
457
458
leafy twigs, densely hairy, margin hairy. Leaves 1-3-foliolate, 2039 cm long, petioles 2.59
cm long, 24 mm thick, minutely hairy; leaf-rachis 38 cm long, short-hairy; petiolules of
lateral leaflets 515 mm long, minutely hairy; leaflets drying dark brown above, ovate to
ovate-oblong, 1425 x 713 cm, thickly chartaceous, sparsely to densely short-hairy
beneath; base narrowly rounded to cuneate, sometimes asymmetric, margin sharply serrate,
not recurved, apex shortly acuminate, acumen 0.51 cm; midrib sparsely to densely shorthairy; lateral veins 712 pairs, ascending, parallel, open or sometimes looping towards the
margin, distinct beneath, sparsely to densely short-hairy; intercostal veins pseudoscalariform (with many almost straight cross-veins between the lateral veins).
Inflorescences axillary or terminal panicles to 35 cm long, rachis densely to sparsely hairy.
Flowers pale yellowish; sepals ovate, thin, 1.82.5 x 12 mm; petals linear obovate, 23 x
11.5 mm; stamens 2.53 mm long, glabrous, filaments 23 mm long, anthers 0.50.6 mm
long, not apiculate; pistils 23 mm long, styles 1.52 mm long, ovary 11.1 mm across, 3celled, each cell with (5)68(9) ovules. Fruits globose, 1020 mm diameter, without
short apical horns but with three radial lines at the apex, wrinkled, wall not brittle, 1.52
mm thick. Seeds pale brown, 58 x 58 mm, 514 in each fruit.
Distribution. Endemic to Borneo (Kalimantan, Sabah, Sarawak). In Sabah, collected from
Tambunan (SAN 113560 and SAN 113536) and Gunung Lotung, Lamag (SAN 83167) and in
Sarawak from the 4th Division, Baram, Kelabit Highlands (S. 35520).
Ecology. Scattered on ridges, forested river-banks and deep ravines. Flowers recorded
around May and fruits from October to November.
4. Turpinia nitida Merr. & Perry

(Latin, nitidus = shiny or polished; the leaf surface)
l.c. 543; B.L. Linden l.c. 49; Whitmore, Tantra & Sutisna l.c. 33. Type: J. & M. S. Clemens 30840,
British North Borneo, Mt. Kinabalu, Penibukan (holotype UC; isotypes BO, K).
Tree to 15 m tall and 40 cm diameter. Bark fissured or lenticellate, greyish brown to black;
inner bark reddish brown. Sapwood pale yellow. Stipules 3.510 x 25 mm, not very
conspicuous, falling off early from leafy twigs, glabrous, margin entire to sparsely hairy.
Leaves 1357-foliolate, (15)1835(39) cm long; petioles 313 cm long, 24(6) mm
thick, glabrous; leaf-rachis 218 cm long, glabrous; petiolules of lateral leaflets 125 mm
long, glabrous; leaflets, drying pale green, sometimes pale brown, oblong, elliptic-oblong to
lanceolate or ovate, 621 x 311 cm, thickly coriaceous, glabrous, typically shiny on upper
surface; base rounded to acute, sometimes asymmetric, margin sharply to shallowly serrate,
strongly recurved, apex acuminate to cuspidate, acumen 0.51.5(2) cm long; lateral veins
611 pairs, ascending arcuately half way, parallel, open or sometimes looping towards the
margin, obscure on upper surface; intercostal veins reticulate or distinctly tessellate, with no
or very few straight cross-veins between the lateral veins on lower surface. Inflorescences
axillary or terminal panicles, to 45 cm long, rachis glabrous or sparsely hairy. Flowers
yellowish to pink-tinged, sepals oblong to ovate, 1.53.5 x 1.52.5 mm, thin; petals
spathulate, 24 x 1.52 mm; stamens 1.84 mm long, filaments 1.58 mm long, anthers 0.8
1 mm long, slightly to distinctly apiculate; pistils 2.53.5 mm long, styles 23 mm long;
ovary 12 mm across, (2)3(4)-celled, each cell with (3)47(8) ovules. Fruits 3-lobed,
459
typically with three short apical horns at the apex, smooth, woody, 820 mm diameter, wall
not brittle, 1.55 mm thick. Seeds pale brown, 58 mm across, 17 in each fruit.
Distribution. Endemic to Sabah (Ranau, Sandakan, Kudat and Lahad Datu), not yet known
in Sarawak.
Ecology. Montane forest to 2400 m and also in primary and secondary forest on hills near
streams. Frequently found on ultramafic soils. Flowers documented around November to
May, fruits mostly during July to September.
5. Turpinia sphaerocarpa Hassk.
Fig. 1.
(Greek, sphaero = globose, spherical, karpos = fruit; the fruit shape)

Flora 25, 2 (1842) Beibl. 1, 42; Ridley, J. Str. Br. R. As. Soc. 82 (1920) 179, J. Mal. Br. R. As. Soc. 1
(1923) 58; Merrill & Perry l.c. 543; B.L. Linden l.c. 49; Whitmore l.c. 448; Anderson l.c. 324;
Whitmore, Tantra & Sutisna l.c. 332; Pereira l.c. 21. Type: s. coll., s.n., Herbarium Reinwardtianum
in Acad. Lugduno-Batavo (holotype L). Synonyms: Dalrymplea javanica Hassk., Pl. Jav. Rar. (1848)
439; Turpinia latifolia Wall. ex Ridl. l.c. (1920) 178; T. laxiflora Ridl. l.c. (1920) 179.
Tree to 35 m tall and 90 cm in diameter; buttresses to 6 m high. Bark slightly fissured, flaky
to smooth, greyish brown, yellowish brown to pale brown; inner bark yellowish to brown.
Sapwood yellow to white. Stipules 47 x 35 mm, not very conspicuous, falling off early
from leafy twigs, glabrous to minutely hairy, margin entire to sparsely hairy. Leaves 357
9-foliolate, 946 cm long; petioles (2)411 cm long, 15 mm thick, glabrous; leaf-rachis 2
18 cm long, glabrous; petiolules of the lateral leaflets 115 mm long, glabrous; leaflets
drying greenish brown to dark purplish brown, ovate-elliptic, elliptic, ovate, sometimes
lanceolate to oblanceolate, 519 x 22.511 cm, chartaceous to coriaceous, glabrous, not shiny
to shiny above; base rounded to cuneate sometimes asymmetric, margin shallowly to deeply
serrate, not recurved, apex shortly acuminate to cuspidate, acumen 0.41 cm long; midrib
raised and distinct on lower surface; lateral veins 510 pairs, ascending, parallel, open or
looping towards the margin; intercostal veins pseudoscalariform (with many almost straight
cross-veins between the lateral veins) to reticulate with wide areoles, obscurely or distinctly
tessellate (with no or very few straight cross-veins between the lateral veins).
Inflorescences axillary to terminal panicles, 1740 cm long, rachis sparsely to densely hairy
at the distal end. Flowers with sepals ovate, thin, 12 x 12 mm; petals spathulate to linear
obovate, 1.73.5(4) x 0.81.5 mm, thin; stamens 1.53.5 mm long, filaments 13 mm
long, anthers 0.40.6 mm long, apiculate or not; pistils 1.53.5 mm long, styles 13 mm
long, ovary 0.81.2(1.5) mm across, (2) 3-celled, each cell with (4)57(9) ovules. Fruits
518 mm diameter, slightly trilobed at the apex and with three short apical horns, or
globose and without short apical horns but sometimes with three radial lines at the apex,
smooth to wrinkled upon drying; wall not brittle, 0.53 mm thick (rarely 0.10.3 mm).
Seeds pale brown to dark brown, 29 x 29 mm, 17 in each fruit.
460
Key to varieties
Leaf-rachises 1018 cm long. Leaves typically 2046 cm long. Leaflets (7)1019 x (3)5
11 cm. Fruits globose, without short apical horns...............................................................
var. sphaerocarpa
Synonym: Dalrymplea javanica Hassk., l.c. 439; Turpinia latifolia Wall. ex Ridl. l.c. (1920)
178; Turpinia laxiflora Ridl. l.c. (1920) 179.
Sumatra, Peninsular Malaysia, Singapore, Borneo, Java, Philippines, Lesser Sunda

Islands & Moluccas. Common in Sabah and Sarawak and scattered in primary and
secondary lowland to submontane forests to 1500 m, near river banks, also occurring
in limestone forest. Found on various types of high nutrient soils. Vernacular names:
Sarawakmaba (Iban), entrang (Sg. Sinjan, Kuching), tait berak (Kelabit).
Leaf-rachises 28 cm long. Leaves 918(20) cm long. Leaflets 510 x 2.55 cm. Fruits
slightly trilobed at the apex, with three short horns...............................................................
var. microcerotis J. T. Pereira
Sandakania 5 (1994) 21. Type: J. & M. S. Clemens 28707, British North Borneo, Ranau,
Kinabalu, Tenompok (holotype SING; isotypes BO, K).
Borneo (Sabah and Kalimantan). In Sabah collected from Tenompok and Mt. Kinabalu,
Ranau. Not yet recorded for Sarawak. Lowlands, on sandy loam soils (Kalimantan) to
montane forest to c. 1500 m (Sabah). Vernacular name: Sabahtapong-tapong
(Dusun).
6. Turpinia stipulacea B.L. Linden

(Latin, stipula = stipule; the conspicuous stipules)
l.c. 49; Whitmore, Tantra & Sutisna l.c. 332. Type: Carr SFN 27516, British North Borneo, Mt.
Kinabalu, near Tibabah River (holotype SING).
Tree to 10 m tall and 10 cm diameter. Bark thin, greyish, slightly fissured; inner bark dull
pale brownish. Sapwood pale yellow. Stipules 1425 x 415 mm, conspicuous and persistent
on apices of leafy twigs, woody, glabrous, margin entire. Leaves 7911-foliolate, 2043
cm long; petioles 58 cm long, stout, glabrous; leaf-rachis 2030 cm long; petiolules of
lateral leaflets 311 cm, stout, glabrous; leaflets drying reddish brown to greenish brown
above, ovate-elliptic, 714 x 27 cm, chartaceous to thinly coriaceous, glabrous; base
cuneate to rounded, margin shallowly undulate to deeply serrate, apex shortly acuminate,
acumen 0.50.8 cm long; lateral veins 811 pairs, ascending, parallel, or open towards the
margin, distinct on lower surface; intercostal veins pseudoscalariform (with many almost
straight cross-veins between the lateral veins). Inflorescences axillary panicles, to 35 cm
long, rachis subglabrous. Flowers with sepals oblong to ovate, 23.5 x 1.52.5 mm, thick;
petals linear to elliptic, 2.54.5 x 11.5 mm, thin; stamens 23 mm long, glabrous, filaments
1.82.8 mm, anthers 0.40.7 mm long, distinctly apiculate; pistils 23 mm long, styles c. 2
mm long; ovary 11.5 mm across, 3-celled, each cell with (2)34 ovules. Fruits globose,
(6)1016 mm diameter, without short apical horns but with three radial lines at the apex,
wrinkled, wall not brittle, 0.51.5 mm thick. Seeds dark brown, 610 x 610 mm, 14 in
each fruit.
461
Distribution. Endemic to Sabah (Ranau only).

Ecology. Occurs along streams or ridges on montane forest to 2100 m.This species
superficially resembles Turpinia sphaerocarpa in some vegetative characters. It can be
distinguished by the larger and more persistent stipules and smaller number of ovules.
462
STYRACACEAE
R. Kiew
Universiti Pertanian Malaysia,
Serdang, Malaysia
Merrill, EB (1921) 485; Backer & Bakhuizen f., FJ 2 (1965) 203; van Steenis, FM 1, 4 (1949) 49, FM
1, 6 (1972) 976; Cockburn, TS 2 (1980) 103; Anderson, CLTS (1980) 327; Whitmore, Tantra &
Sutisna, CLK 2, 1 (1990) 340.
Trees or shrubs. Leaves simple, spirally arranged or pseudo-alternate, margin entire or

toothed, mostly with stellate or lepidote indumentum, without stipules. Inflorescences
racemes or panicles, axillary or terminal. Flowers regular; calyx tubular more or less adnate
to the ovary, lobes if present valvate; corolla mostly joined into a basal tube (rarely of free
petals), lobes 47, valvate or imbricate; stamens equal and alternate or double the number of
petals, mostly adnate to the tube; ovary superior (rarely semi-inferior), 35-celled, style 1,
stigma punctiform to 35-lobed. Fruits capsules (rarely berries), dehiscent or not, pericarp
often thick and woody or corky, calyx persistent. Seeds 1 to many per locule, with
endosperm.
Distribution. 12 genera and c. 190 species, mostly in the northern hemisphere, especially E
Asia (absent from Australia and the Central Pacific). In Malesia, represented by 2 genera,
Bruinsmia and Styrax, with the greatest diversity in Sumatra and no species as yet being
found in the Philippines, C and E Java and the Lesser Sunda Islands. Two species are
known from Borneo, one, Styrax agreste, occupies the southern half of the island and has
not yet been collected from Sabah and Sarawak, although it may be expected; the other,
Bruinsmia styracoides, is less collected from Sarawak than from Sabah.
Ecology. Lowland and submontane forests to 1600 m.
Uses. Styrax benzoin is the source of benzoin resin (kemenyan), which is used for medicinal
and a variety of other purposes.
Taxonomy. Symplocos is treated as a separate family as it does not have the stellate hairs,
nor fasciculate stamens, linear anthers and the half or wholly superior ovary characteristic of
the Styracaceae.
BRUINSMIA Boerl. & Koord.

(A.E.J. Bruinsma, 18521943, Dutch East India Forest Service)
Nat. Tijd. Ned. Ind. 53(1893) 68; Perkins, Pfl. R. 30 (1907) 14, 88; van Steenis, Bull. Jard. Bot. Btzg.
3, 12 (1932) 215, l.c. (1949) 49; van Steenis & Bakhuizen f., Bot. Jahrb. 86 (1967) 393; Cockburn, TS
2 (1980) 103.
Glabrate trees with flattened-angular twigs. Leaf margin entire to coarsely crenate.
Inflorescences compound, dichotomous, terminal or axillary panicles; pedicels articulated
463
464
STYRACACEAE (KIEW)
at apex. Flowers dull green or white; corolla-lobes free without a staminal tube; ovary
glabrous, 35(6)-loculed. Fruits fleshy, indehiscent, style base persistent. Seeds minute,
1many per locule.
Distribution. 2 species, one in Borneo; the other, B. polysperma (Clarke) Steenis, is known
from India (Assam), Burma and Thailand (Chiangmai).
Taxonomy. Bruinsmia is distinguished from Styrax by the leaves, which are glabrescent or
sparsely stellate hairy, by the compound dichotomous inflorescences (as opposed to
racemose-paniculate), the articulate flower-stalks, the 3- to 5-celled ovary and fruit, and by
the many seeds in the fruit.
Bruinsmia styracoides Boerl. & Koord.
Fig. 1.
(resembling the classic Greek tree, Styrax officinalis, which yields the resin storax)
I.c. 68; Perkins l.c. 14, 88; van Steenis l.c. (1932) 215, l.c. (1949) 49; Cockburn l.c. 103; Anderson l.c.
327; Whitmore, Tantra & K. Sutisna l.c. 340. Type: Koorders 8529 and 8530 (1893), Java
(syntypes BO, K, L).
Tree to 45 m tall and 30200 cm diameter; clear bole 23 m; crown large and spreading,
flowering at 5 m; trees over 25 m with buttresses c. 65100 cm high, 57 cm thick and 65
80 cm out. Bark smooth becoming slightly fissured with age, grey to grey-brown; inner
bark soft and stringy, pale brown becoming reddish brown on exposure with a jelly-like
exudate that reddens on exposure. Sapwood white. Twigs flattened-angular at nodes.
Indumentum of young twigs, lower leaf surface and panicle dense, comprising stellate hairs,
persisting only on inflorescences. Leaves ovate to oblong, 7.519 x 310 cm, chartaceous;
base usually rounded, margin serrate, apex acute to acuminate; lateral veins 610 pairs;
stalk grooved, 0.51.5 cm long. Panicles terminal and 4.525 cm long or axillary and 2.57
cm long, with dense yellow-brown pubescence; pedicels c. 1 mm, lengthening to 23 mm in
fruit, articulated; bracteoles narrow, 12.5 mm long. Flowers: calyx broadly cup-shaped,
entire or shortly 5-toothed, 1.23 mm long and 5 mm wide; corolla lobes 5(6), free, ovateoblong, 910 x 44.5 mm, imbricate, pale yellow, silvery hairy on both sides; stamens
yellow-orange, 10(12), 5 longer (66.5 mm) alternate, 5 shorter (55.5 mm) epipetalous, or
10 subequal (3.54 mm), joined to the base of corolla; ovary 23 x 2.53.5 mm, imperfectly
5(6)-locular; style 37 mm long, stigma capitate; ovules many. Fruits dark green, on dishlike calyx 7 mm wide, globular to pear-shaped, 610 x 69 mm, attenuated to persistent
style base 0.76 mm long; fruit-wall indehiscent, thick and pulpy. Seeds many per locule, c.
1.5 mm long, black, 4-angular and very finely corrugated.
Vernacular names. Sabahtingo-tingo (Dusun Kundasan). Sarawakempudu (? Iban).
Distribution. Sumatra, W Java, Borneo, Philippines (Mindanao), Celebes, Moluccas (Bacan
Is.) and New Guinea. In Sabah and Sarawak, widely distributed but more common in Sabah.
Ecology. On steep hill sides or ridges in primary, occasionally in disturbed forest, from the
lowland at 450 m to montane forest at 2000 m. Most flowering and fruiting occurs between
May and November, buds and mature fruits develop simultaneously.
465
TRIGONIACEAE
K.M. Wong
Sandakan, Malaysia
Merrill, PEB (1929) 132; Masamune, EPB (1942) 378; van Steenis, FM 1, 4 (1949) 59; Kochummen,
TFM 1 (1972) 449; Anderson, CLTS (1980) 340; Ashton, MNDTS (1988) 469; Whitmore, Tantra &
Sutisna, CLK 2, 1 (1990) 361.
Trees or shrubs. Leaves simple, entire; stipules small, caducous. Inflorescences racemose
or paniculate, axillary and terminal. Flowers bisexual, bilaterally symmetrical; sepals 5,
free or joined at the base, imbricate, unequal; petals 35, free, imbricate, unequal; stamens
312, grouped to one side opposite the anterior petals, filaments fused at the base, anthers
2-celled and dehiscing lengthwise; ovary superior, 3-locular, woolly; style simple; ovules
1several per locule, placentation axile. Fruit a 3-lobed structure splitting into 3 flat,
winged fruit portions (samaras). Seeds compressed, hairy, endosperm none.
Distribution. 3 genera and 44 species, in tropical South America and Malesia. In Malesia,
only the monotypic genus Trigoniastrum is found.
Taxonomy. Trigoniaceae is generally accepted as most closely related to the Polygalaceae.
TRIGONIASTRUM Miq.
(Latin, trigonus = 3-angled, aster = star; the fruit form)
Fl. Ind. Bat. Suppl. (1860) 394; Merrill l.c. 132; Masamune l.c. 378; van Steenis l.c. 59; Kochummen
l.c. 449; Anderson l.c. 340; Ashton l.c. 469; Whitmore, Tantra & Sutisna l.c. 361.
Trees. Leaves alternate and distichous on branches. Flowers with 5 petals, the posterior
one largest and saccate and with a concave gland at the mouth of the sac, the lateral ones
spathulate and spreading, the anterior two closer together and obliquely held; stamens 6;
ovules solitary in each locule.
Distribution. Monotypic, confined to Sumatra, Peninsular Malaysia and Borneo.
Trigoniastrum hypoleucum Miq.
Fig. 1.
(Greek, hypo = lower side, leuco = white or glaucous; the leaf surface)
l.c. 394; Merrill l.c. 132; Masamune l.c. 378; van Steenis l.c. 59; Kochummen l.c. 449; Anderson l.c.
340; Ashton l.c. 469; Whitmore, Tantra & Sutisna l.c. 361. Type: Teijsmann, s.n., Sumatra,
467
468
TRIGONIACEAE (WONG)
Lampong, Mengala (holotype L; isotype K). Synonym: Isopterys penangiana Wall. ex A.W. Benn. in
Hooker f., Fl. Br. Ind. 1 (1872) 208.
Small to medium-sized tree, to 30 m tall and 50 cm diameter, sometimes with buttresses up

to 1 m high. Bark pale greenish grey to brown, smooth to lenticellate; inner bark pale
greenish yellow to yellowish brown, exuding a yellowish sticky sap which turns reddish on
exposure. Sapwood yellowish ochre, hard. Leaves elliptic, 520 x 1.58 cm, chartaceous to
thickly coriaceous, glaucous beneath, glabrous or yellow-hairy on the veins below and
midrib on upper surface; base cuneate to rounded, apex acute to cuspidate; midrib sunken to
flat on upper side, prominent on lower surface, drying pale brown to dark purplish; lateral
veins 58 pairs, distinct, looping coarsely towards the margin; intercostal veins reticulate,
prominent on the lower leaf-surface and sometimes the upper surface; stalks 59 mm long.
Inflorescences 715 cm long, paniculate, the axes scantily to densely yellow short-hairy all
over. Flowers fragrant; sepals ovate-acute, c. 2.5 x 1.5 mm, pale green, yellow short-hairy;
posterior petal white with a yellowish basal sac-like part c. 2 mm across and a reflexed
distal part c. 2.5 mm long and 2 mm wide, the concave gland at the mouth of the sac to 0.8
mm across; lateral petals spathulate, 3.54 x 1.5 mm, creamy yellow; anterior petals oblong,
oblique, 45 mm long, white; staminal tube c. 1 mm long, anthers c. 0.5 mm long; ovary
pale yellow woolly, to 0.5 mm across, style c. 1.5 mm long and subglabrous. Fruit a 3lobed structure; each lobe a samara, ovate, flattened and wing-like, the inner basal portion
straight and fused with the other 2 lobes, the whole lobe to 6 cm long and 2.5 cm wide,
yellowish green ripening pale brown.
Vernacular names. Sabahnone generally, but once (SAN 32418) documented as miapa
(Dusun Bundu Tuhan). Sarawakatap (Kenyah), lia (Iban), ngilis (Iban), nyalin (Iban: in
common with Xanthophyllum which it resembles), nyalin bintek (Iban), tulang kwe
(Kenyah). Bruneimengilas babi (Iban).
Distribution. As for genus. In Sabah and Sarawak in all districts. Also in Brunei and
Kalimantan.
Ecology. Although widespread in Borneo, the species is nowhere common. It has a wide
ecological amplitude and has been collected from peat swamp, freshwater swamp, heath
(kerangas), alluvial and mixed dipterocarp forests in the lowlands and hills, to submontane
forest at about 1200 m. Ashton (l.c.) points out there may be two ecotypes, one a small tree
with thick leaves found in peat swamp, heath and mixed dipterocarp forests on poor sandy
soils, and the other a larger tree with thinner, narrower leaves in mixed dipterocarp forest on
richer sites. Van Steenis (l.c.) records flowering and fruiting specimens collected in every
month of the year; it is possible, though, that there are local variations in the phenology of
the species. The winged fruits are wind-dispersed.
Uses. None recorded, but the wood has been tested and is listed as being hard or very hard
and heavy or very heavy, with a moderately fine texture (Wong (1982) DMT).
The leafy twigs alone are easily confused with those of Xanthophyllum (Polygalaceae),
although the flowers and fruits readily distinguish it. In many Xanthophyllum species, the
leaf blades have distinct glands and the ultimate leafy twigs are green, and so are different.
469
ABBREVIATIONS OF
FREQUENTLY CITED REFERENCES
Anderson, J.A.R. 1980. A Checklist of the Trees of Sarawak. Forestry Dept., Sarawak.
CLTS
Ashton, P.S. 1988. Manual of the Non-Dipterocarp Trees of Sarawak. Forestry Dept.,
Sarawak.
MNDTS
Backer, C.A. & R.C. Bakhuizen v.d. Brink, Jr. 1965. Flora of Java. Noordhoff, Groningen,
the Netherlands.
FJ
Browne, F.G. 1955. Forest Trees of Sarawak and Brunei. Govt. Press, Sarawak.
FTSB
Burgess, P.F. 1966. Timbers of Sabah. Forestry Dept., Sabah.
TBS
Burkill, I.H. 1966. A Dictionary of the Economic Products of the Malay Peninsula. Kuala
Lumpur.
EPMP
Cockburn, P.F. 1976 & 1980. Trees of Sabah, Vols. 1 & 2. Forestry Dept., Sabah.
TS
Corner, E.J.H. 1988. Wayside Trees of Malaya. 2 Vols. Malayan Nature Society, Kuala
Lumpur.
WSTM
Flora Malesiana
FM
Keith, H.G. 1937. The Timbers of North Borneo. Govt. Col. N. Borneo, Sandakan.
TNB
Keng, H. 1969. Orders and Families of Malayan Seed Plants. University of Malaya Press,
Kuala Lumpur.
OFMSP
Mabberley, D.J. 1987. The Plant Book. Cambridge University Press, Cambridge.
PB
Masamune, G. 1942. Enumeratio Phanerogamarum Bornearum. Taihoku.
EPB
471
Soc., Sp. No.
EB
Merrill, E.D. 1929. Plantae Elmerianae Borneenses. University of California Publications
in Botany, Vol. 15. University of California Press, Berkeley.
PEB
Ng, F.S.P. (ed.). 1978 & 1989. Tree Flora of Malaya, Vols. 3 & 4. Longman, Malaysia.
TFM
Ng, F.S.P. 1991 & 1992. Manual of Forest Fruits, Seeds and Seedlings. Malaysian Forest
Records, No. 34. Vols. 1 & 2. FRIM, Malaysia.
MFFSS
Perry, L.M. 1980. Medicinal Plants of East and Southeast Asia. Cambridge.
MPESA
Plant Resources of South East Asia. Wageningen.(Various authors and volumes).
(PROSEA, to be listed without editor or author names, but to include volume and page
numbers).
Ridley, H.N. 1923-1925. The Flora of the Malay Peninsula, Vols. 1-5. Reeves, London.
FMP
Smythies, B.E. 1965. Common Sarawak Trees. Borneo Literature Bureau, Sarawak.
CST
Stapf, O. 1894. On the Flora of Mount Kinabalu, in North Borneo. Transactions of the
Linnean Society, London, Ser. 2, 4, pp. 69-263.
FMK
Whitmore, T.C. (ed.). 1972 & 1973. Tree Flora of Malaya, Vols. 1 & 2. Longman,
Malaysia.
TFM
Whitmore, T.C. & I.G.M. Tantra. 1986. Tree Flora of Indonesia. Check List for Sumatra.
Forest Research & Development Centre, Bogor.
CLS
Whitmore, T.C., I.G.M.Tantra & U. Sutisna. 1990. Tree Flora of Indonesia. Check List
for Kalimantan, Part 1 & Parts 2.1-2.2. Forest Research & Development Centre, Bogor.
CLK
Willis, J.C. 1973. A Dictionary of the Flowering Plants & Ferns, revised by H.K. Airy
Shaw. Cambridge University Press, Cambridge.
DFPF
472
Wong, T.M. 1982. A Dictionary of Malaysian Timbers. Malayan Forest Records No. 30.
Forest Research Institute, Kepong, Kuala Lumpur.
DMT
COMMONLY USED
ABBREVIATIONS FOR LOCALITIES
English
Word
Malay
Abbreviation
Word
Abbreviation
Central
Bukit
Bt.
Division
Div.
Gunung
G.
East
Kampung
Kg.
Forest Reserve
FR
Sungai
Sg.
Island
Is.
Tanjung
Tg.
Mount
Mt.
National Park
NP
North-East
NE
North-West
NW
River
R.
South
South-East
SE
South-West
SW
West
473
GLOSSARY
abaxial
accrescent
achene
acrid
acrodromous
acroscopic
actinomorphic
acumen
acuminate
acute
adaxial
adherent
adnate
alate
anastomosing
anatropous
androdioecious
androecium
androgynophore
anisophylly
annular
annulus
anterior
anther
anthesis
antidysenteric
antiemetic
antihelminthic
apetalous
apiculate
arachnoid
arboreous
arcuate
areolation
areole
aril
arillate
the side of an organ facing away from the distal portion of the
axis which bears it
increasing in size with age, e.g., the calyx of some plants after
flowering
a small, hard, dry, non-splitting fruit
sharp, irritating to the taste
with veins converging and uniting at the apex of the leaf
towards the apex or distal part of an organ; e.g., flowers in an
inflorescence
(flowers) radially symmetrical
a tapering point or tip
drawn out into a long point; having a gradually tapering point;
pointed; e.g., leaf apex
ending in a sharp tapering point; e.g., leaf apex
the side of an organ facing the distal portion of the axis which
bears it
in contact but not fused together; e.g., floral parts
attached to some other organ
winged; as a stem or petiole
joining of veins to form a network
(ovules) with hilum and micropyle close together and chalaza
at the other end
plants with male or hermaphrodite flowers
male reproductive organs of a plant
common structure or stalk that supports androecium and
gynoecium
the occurrence of leaves with distinct form and size
ring-like
ring
frontal
the part of a stamen which contains pollen
(flowers) that period between the opening of the bud and the
withering of the stigma and/or stamen
against dysentery
preventing vomiting
working against worms
without petals
with an abrupt small tip, as a leaf
like a cobweb
with the habit of a tree; tree-like
curved or shaped like a bow; arc-shaped
of fine veins anastomosing and enclosing tiny spaces
a tiny space marked on a surface; e.g., on leaves
a fleshy expansion of the funicle, arising from the placenta, and
enveloping a seed
having / bearing an aril
474
arillode
arilloid
articulated
attenuate
auricle
auriculate
axile
axillar/axillary
ballistic
basal
basifixed
berry
bifid
bifoliolate
bilocular
biseriate
bisexual
bistipulate
bitegmic
bract
bracteate
bracteolate
bracteole
bud
bullate
caducous
calcareous
calyptra
calyx
campanulate
camptotropous
canaliculate
capitate
capsular
capsule
carinate
carminative
carpel
carpellate
cartilaginous
caruncle
a false aril
resembling an aril
jointed
gradually narrowing to a tip or base; e.g., leaf-blade
ear-like lobe or appendage
with auricles; e.g., leaf with expanded bases surrounding stem
(ovule placentation) with ovules attached to central axis within
the ovary
(inflorescence; bud) borne in the axil, i.e. the junction between
leaf-stalk and stem
(seed dispersal) referring to the fruit which discharges its seeds
elastically
at or near the base of an organ
(stamens) having filament attached to anther base
non-spliting pulpy fruit with many seeds
forked; divided nearly to middle line
with two leaflets
having two cavities or chambers; eg., anther or ovary
in two series
having both male and female reproductive organs; hermaphrodite
with two stipules
(ovules) with two layers of integuments
a modified leaf in whose axil a flower or branch axis arises
having bracts
with bracteoles
a secondary bract as one on a pedicel of a flower, usually
smaller than bract
a rudimentary state of a stem or branch; an unexpanded flower
blistered like a savoy-cabbage leaf
fall off very early; deciduous; e.g., stipules
growing on soil derived from decomposition of calcareous
rocks; limy
a cap-shaped structure
the outer whorl of floral envelope; collective name for sepals
bell-shaped
(ovule) curved or bent like a horse-shoe
longitudinally channelled or grooved
enlarged or swollen at tip; arranged into a mass at apex, as
some inflorescences
possessing fruit of capsule-type
a dry, dehiscent fruit
having a ridge or a keel
relieving flatulence
a simple pistil formed by a single leaf-like, ovule-bearing
structure; a basic ovule-bearing chamber of the ovary
with carpels
firm and tough but elastic like cartilage
a wart or protuberance near the hilum of a seed
475
cataphyll
catkin
caudate
cauliflorous
chalaza
chartaceous
ciliate
ciliolate
circumscissile
coherent
colic
collateral
colleter
columella
columnar
commissure
concave
concolorous
concrescent
cone
conical/ coniform
connate
connective
connivent
contorted
convex
convolute
cordate
coriaceous
corolla
corymb
corymbiform/corymbose
crassinucellate
crenate
crenulate
crescent
the early leaf-forms of a plant or shoot, as cotyledons, budscales, rhizome scales, etc.
a deciduous spike, consisting of unisexual apetalous flowers
with a slender tail-like appendage
having flowers on the main stem
that part of an ovule or a seed where the nucellus joins the
integuments
papery
fringed with hairs
ciliate but hairs minute
dehising in transverse circular line, the top separating like the
lid of a pill box; e.g., in fruits
the incorporation of one part with another, as the petals to form
a tubular corolla
a severe spasmodic abdominal pain
positioned side by side
mucilaginous hairs on the buds of many flowering plants
which secrete gum
a persistent central axis around which the carpels of some fruits
are arranged
having a form of a column; pillar-like
the face by which one carpel joins another as in the
Umbelliferae (cabbage family)
having an outline or surface curved like the interior of a circle
or sphere
uniform in tint
growing together
a dry multiple fruit- or seed-bearing structure; e.g., in pine tree
cone-shaped
united to similar structures
the portion of a stamen distinct from the filament which
connects the two halves of an anther
coming in contact; converging
twisted
having a more or less rounded surface
rolled around
heart-shaped
leathery in texture
the inner series of floral envelope, composed of petals;
collective name for petals
a flat-topped or convex and open flower-cluster of the
indeterminate or centripetal order
arranged in corymbs
(ovule) having a thick nucellus
toothed with the teeth rounded at apex
crenate with small teeth
a structure having a shape like the moon as seen in the first or
the last quarter of the month
476
cristate
crustaceous
crypts
cryptocotylar
cucullate
cuneate
cupular
cupuliform
cuspidate
cyme
cymose
cymules
deciduous
decurrent
decussate
dehisce
dehiscent
deltate/deltoid
dentate
denticulate
dichotomous
didynamous
dimorphic
dioecious
discoid
dispersal
distichous
distylous
diuretic
divaricate
divergent
domatia
dorsifixed
drupaceous
drupe
druplet
dyspeptic
eglandular
crested; having a tesselled margin

brittle in texture
stomatal pits; also applied to sunken glands, receptacles for
secretions in a plant with dotted leaves
(seed) with hidden cotyledons during germination
hooded or hood-shaped
wedge-shaped
furnished with, or subtended by a cup-like structure
cup-shaped
tipped with a sharp, rigid point
a flower-cluster of determinate or centrifugal type, often
convex or flat-topped
bearing cymes; cyme-like
a small cyme or portion of one
falling off, as petals fall after flowering or leaves in autumn or
dry season
running down, as when leaves are prolonged beyond their
insertion, and thus run down the stem
in pairs that alternate at right angle, thus in four rows
gape or burst open spontaneously when ripe, as seed capsules
or anthers
opening by definite pores or slits at maturity to release the
content; as in fruits or anthers
shaped like a Greek letter Delta; equilaterally triangular
toothed
minutely toothed
two-forked, the branches equal or nearly so
(flower) having four stamens arranged in pairs, two long and
two short
in two forms
(plant) having unisexual flowers, with the staminate and
pistillate flowers borne on separate individuals
resembling a disk
the various ways by which seeds or fruits are scattered away
from the mother-trees
disposed in two vertical ranks, as in leaves
having two styles
promoting the secretion of urine
extremely divergent
separated or turning in different directions
dome-like, usually hairy projections sheltering parasites
(stamen) having filament attached to the back of an anther
resembling a drupe
one-seeded, indehiscent fruit with the pericarp fleshy or
leathery and the seed-coat hard and stony
a diminutive drupe
stomach discomfort; e.g., indigestion
without glands
477
ejaculatory apparatus
ellipsoid
elliptic
emarginate
emetic
endocarp
endocarpid
endosperm
entire
eophyll
ephemeral
epicarp
epigeal
epipetalous
epiphyllous
epiphytic
episepalous
epitropous
estipulate
exarillate
excentric
excrescence
exfoliate
exocarp
exserted
exstipulate
extra
extrastaminal
extrorse
exudate
falcate
fascicle
fasciculate
febrifuge
ferruginous
filament
filamentous
filiform
fimbriate
fissured
flavedo
foliaceous
a structure that forcibly throws out endogenously formed

content; e.g., in fruit
a tridimensional structure elliptic in outline; e.g., seeds
shaped like an ellipse, oblong with regularly rounded ends;
e.g., leaf-blade
having a notch cut out, usually at the extremity
that causes vomiting
the inner layer of a pericarp
indehiscent kernel or pyrene embedded in certain fruits
albumen deposited within the embryo-sac of a seed
having an even margin, without lobes or teeth; e.g., leaf-blade
early form of leaf
lasting for a day or less
the external layer of a pericarp
(seed germination) with the cotyledons lifted above the ground
surface by an elongating axis
(stamens) borne upon the petals
borne upon the leaves
relating to epiphytes
borne upon the sepals
denotes an anatropous ovule with its raphe averse when
ascending, adverse when suspended
without stipules
without aril
out of the centre; one-sided
growing in an unnatural way, as a wart or other outgrowth
to come away in scales or flakes
the outer layer of a pericarp
projecting beyond the surrounding organ, as stamens beyond
the tube of the corolla
without stipules
outside or beyond
arising or situated outside the stamens
directed outwards, as the dehiscence of an anther
discharge from incision or pore; e.g., gum, latex, moisture,
resin, etc.
sickle-shaped
a close bundle or cluster of flowers, leaves, etc.
connected or drawn into a fascicle
a medicine that reduces fever
rust-coloured
the stalk of an anther
having a form of a filament
thread-shaped
with the margin bordered by long slender processes
deeply grooved or furrowed
yellowness
having the texture or shape of a leaf
478
follicle
follicular
fulvous
funicle/funiculus
furfuraceous
fusiform
gametophyte
germicidal
germicide
gibbous
glabrate/ glabrous
glabrescent
glandulous
glaucous
globose/globular
granular
gynobasic
gynoecium
gynophore
herbaceous
hermaphrodite
hesperidium
heterostylous
hilum
hirsute
hirsutulous
homostylous
hyaline
hypanthium
hypocotyl
hypogeal/hypogeous
hypogynous
idioblast
imbricate
imparipinnate
a fruit of one carpel, opening by a ventral suture to which the

seeds are attached
shaped like a follicle
dull yellow; tawny
the cord or thread which connects the ovule or seed to the
placenta
scurfy; having soft scales
spindle-shaped
the generation of a plant that bears the sexual organs,
producing gametes, in turn giving rise to the sporophyte
ability to destroy germs
a substance that destroys germs
enlarged, humped or swollen on one side
smooth; without pubescence
becoming glabrous, or slightly so
possessing glands
bluish green
rounded or spherical in shape
like grains or granules
denotes a style that adheres by its base to a prolongation
upwards of the torus between carpels
an organ in which female cells are formed
a stalk supporting the ovary/pistil/carpel
having the texture, colour and properties of a herb
(flower) with both stamens and pistils; having the
characteristics of both sexes
many-celled, few-seeded indehiscent fruit, having the epicarp
and mesocarp joined together, and the endocarp projecting into
the interior as membranous partitions which divide the pulp
into chambers; such as the orange
(flowers) with different types (length) of styles
the scar left on a seed where formerly attached to the funicle or
placenta
coarse-hairy
minutely hirsute
with uniform styles
colourless or translucent
an enlargement or elongation of the torus or floral axis below
the calyx
the portion of the axis below cotyledons in an embryo
(seed germination) with cotyledons not lifted above the ground
surface by an elongating axis
free from but inserted below the gynoecium or pistil; e.g.,
petals and sepals
a special cell in a tissue that markedly differs from the rest in
form, size, or contents
overlapping
pinnate with an odd terminal leaflet
479
incipient bract
indehiscent
indumentum
induplicate
indusium
inferior
inflexed
inflorescence
infructescence
insecticidal
intercostal
interpetiolar
intervenium
intra
intramarginal
intrapetiolar
intrastaminal
introrse
invaginating
juvenile
lacerate
laciniate
lamellate
lamina
laminate
lanate
lanceolate
laticiferous
latrorse
laxative
lenticel
lenticellate
lenticular
lepidote
ligulate
ligule
linear
lobulate
locular/loculate
locule/ loculus
bract in an initial stage of development

not opening by valves or along regular lines
hairy covering
with the margins bent inwards, and the external face of these
edges, applied to each other, without twisting
an epidermal outgrowth covering and protecting another
structure; a ring of collecting hairs below the stigma
(ovary) arising or growing at a level below the insertion of
other floral parts
turned abruptly or bent inwards
the deposition of the flowers on the floral axis; flower cluster
the inflorescence in a fruiting stage; collective fruits
ability to destroy insects
between veins of a leaf
between the petioles
a portion of parenchyma between the veins of a leaf
within
placed within the margin near the edge
within the petiole or between it and the stem
within the stamens, as the disc of Anacardiaceae (the mango
family)
turned inwards, towards the axis
involuting or drawn into a sheath
young stage of growth
torn or irregularly cleft
incised or slashed into narrow lobes
made up of thin plates
the blade of a leaf
consisting of plates or layers
clothed with woolly and intergrown hairs
lance-shaped; narrow and tapering to each end
latex-bearing
turned or directed towards the side of an organ; e.g., anthers of
a flower
tending to stimulate or facilitate evacuation of the bowel
lenticular corky spots on young bark, corresponding to
epidermal stomata
having lenticels
like a doubly convex lens
beset with small scurfy scales
furnished with a ligule
a strap-shaped body/structure, such as the limb of the ray
florets in Compositae (sun-flower family)
narrow, several times longer than wide
having small lobes
divided into locules or cavities
the cell or cavity of an ovary or an anther
480
loculicidal
lysigenous
megasporophyll
membranous
merism
mesocarp
micropyle
microsporangium
microsporophyll
monoecious
monoembryonic
monogeneric
monopodial
monotypic
mucilaginous
mucro
mucronate
mucronulate
multiflorous
nectariferous
nectarivorous
nocturnal
nucellus
obcordate
oblanceolate
oblong
obovate
obpyriform
obturator
obtuse
ochraceous
olivaceous
orbicular
orthotropic
orthotropous
ovary
ovate
ovoid
dehiscence of a fruit down the center of a compartment or a

locule
a state when a cavity is formed by a degeneration or dissolving
of cells
seed- or ovule-bearing leaf-like structure
thin and semi-transparent
repetition of parts to form a symmetry or pattern
the middle layer of a pericarp
aperture at the apex of an ovule through which pollen-tube
enters the embryo-sac
a microspore- or pollen-bearing structure or organ
a leaf-like structure bearing microsporangium
(plant) having stamens and pistils in separate flowers, but
borne on the same plant
with only one embryo
a family having a single genus
(tree) with the branches or appendages arising from a simple
axis
a genus with a single species
slimy, composed of mucilage
a sharp point abruptly terminating an organ
abruptly terminated by a short and straight point
tipped by a diminutive mucro
many-flowered
nectar-bearing
nectar-feeding
(flowers) night-blooming
the body of the ovule containing the embryo-sac; the kernel of
an ovule
inversely heart-shaped
inversely lanceolate; tapering towards the base more than
towards the apex
much longer than broad, with nearly parallel sides
inversely egg-shaped, with narrow end attached to stalk
inversely pear-shaped
a wart-like protuberance of the placenta covering the
micropyle
with blunt end
yellow with a tinge of red
olive-coloured
of a flat body with a circular outline
assuming a vertical position
(ovule) having a straight axis, the chalaza being at the insertion
and the micropyle at the opposite end
that part of the pistil which contains the ovules
egg-shaped (two dimensional), the broad end attached to the
stalk
somewhat egg-shaped (three dimensional)
481
ovule
palmate
palmatifid
palynological
panicle
paniculate
papilla
papillate/papillose
paraphyll
parenchyma
parenchymatous
parietal
paripinnate
patelliform
pedicel
peduncle
pedunculate
pellucid
peltate
pendulous
percurrent
perennial
perianth
pericarp
perigynous
petal
petiole
petiolule
phanerocotylar
pilose
pinnate
pistil
pistillode
placenta
placentation
plagiotropic
pleurisy
plicate
the female gamete- or egg-cell-bearing tissue of seed-bearing

plants; the organ which after fertilization develops into a seed
(leaf-blade) divided into lobes which arise from a common
centre
(leaf-blade) cut in a palmate fashion nearly to the petiole
pertaining to pollen and spores
a loose flower-cluster, as a branched raceme or corymb
having flowers clustered in a panicle
soft superficial glands or protuberances
having papillae
leaf-like expansion produced near the leaves
soft, succulent tissue in plant composed of more or less
isodiametric, thin-walled cells
consisting of parenchyma
(ovules) borne on or belonging to the inner wall of an ovary
pinnate, with an equal number of leaflets, that is without a
terminal one
shaped like a small flat dish
the ultimate stalk of a single flower in an inflorescence
the stalk of a solitary flower or an inflorescence
borne upon a peduncle
wholly or partially transparent
shaped like a small shield; fastened to stalk at a point within
margin
hanging or pendent
extending throughout the entire length
persisting throughout the year or for a number of years
the floral envelopes of which calyx and corolla cannot be
distinguished
fruit-wall
(flowers) having sepals, petals and stamens adnate with the
lower part of the pistil
one of the leafy expansions in the floral whorl constituting the
corolla
leaf-stalk
stalk of a leaflet in a compound leaf
(germination) with exposed cotyledons
having soft distinct hairs
compound leaf with the leaflets arranged on each side of the
midrib or rachis
the female organ of a flower, consisting of ovary, style and
stigma
rudimentary pistil
the organ which bears ovules in an ovary
the manner by which ovules are attached to the ovary
having oblique or horizontal direction of growth
inflammation of the pleura
folded into plaits, usually lengthwise
482
plumose
pneumatophore
polyadelphous
polyembryonic
polygamodioecious
polygamous
posterior
prominulous
protandrous
proximal
pseudopuberulous
pubescence
pubescent
pulp-vesicle
pulpy
pulvinate
pulvinus
punctate
punctiform
punctulate
purgative
pustular
pustule
pyrene
pyriform
raceme
racemose
rachis
radicle
ramiflorous
raphe
receptacle
reniform
resinous
reticulate
retrorse
retuse
revolute
feathery; feather-like
an aerating root; e.g., in Rhizophoraceae
having stamens grouped into several bundles
having more than one embryo in a seed
(plant) having bisexual flowers and unisexual flowers on
different individuals of the same species
(plant) having bisexual flowers and unisexual flowers on the
same, or on different individuals of the same species
on the side next or close to the axis
slightly raised, standing out a little from the surface
(flowers) having anthers mature before the pistils in the same
flower
the part nearest to the axis
false or resembling
slightly hairy
hairiness
clothed with soft hair
air cavity or small bladder in the pulp of a fruit; e.g., Citrus
fruit
with juicy or fleshy tissue
cushion-shaped
the swollen base of a leaf-stalk
marked with dots, depressions or translucent glands
in the form of a point or a dot
with minute dots
strongly laxative
having slight elevation like blisters
a blister-like prominence; pimple
a stone or kernel of a fruit; nutlet
pear-shaped
an indeterminate or centripetal inflorescence with lengthened
axis and equally stalked flowers
raceme-like or having racemes
the central elongated axis of an inflorescence or a compound
leaf
the rudimentary root of a plant embryo
flowering on the branches
a ridge of fibrous vascular tissue connecting the base of
nucellus with the placenta
that part of flower stalk (usually fleshy) which bears other
floral organs
kidney-shaped
producing resin or covered with resin
netted like net-work, as venation of a leaf
(anther) turned or directed backwards
with a shallow notch at a rounded apex
rolled back from the margin or apex
483
rheumatism
rhomboid
rosette
rugose
rugulose
ruminate
saccate
sagittate
samara
sarcotesta
sarcotheca
scabies
scabrous
scalariform
scale
scandent
schizocarp
scurfy
semi-inferior
semi-superior
sepal
septicidal
seriate
sericeous
serrate
serrulate
sessile
setaceous
sinuate/sinuous
spathaceous
spathe
spathulate
spicate
spike
stamen
staminate
staminode
stellate
stigma
any disease marked by inflammation and pain in the joints ,

muscles or fibrous tissue
approaching a rhombic outline; quadrangular with the lateral
angles obtuse
arranged in a condensed circular fashion
covered with wrinkles
somewhat wrinkled
(endosperm) having a mottled appearance through infolding of
inner seed-coat
sac-shaped or pouch-shaped
shaped like the barbed head of an arrow
a winged, non-spliting, flat, one-seeded fruit
the fleshy outer portion of seed-coat
fleshy pollen case or theca
a contagious skin disease causing severe itching
rough to the touch due to the presence of stiff hairs, scales, or
points
having markings suggestive of a ladder
any thin scarious body, usually a degenerate leaf, sometimes of
epidermal origin
climbing
a dry fruit which splits into two or more one-seeded portions
covered with bran-like scales
partially inferior
partially superior
leaf-like divisions of a calyx
dehiscing through the dissepiments or lines of junction
disposed in series of rows, either transverse or longitudinal
clothed with close-pressed soft and straight pubescence
beset with sharp teeth directed upwards on the margin
serrate but the teeth minute
without stalk
bristle-like
with a deep wavy margin
resembling or bearing a spathe
a large bract sheathing or enclosing an inflorescence or a
flower cluster
oblong, with the basal end attenuate like a druggists spatula
disposed in or resembling a spike
an indeterminate inflorescence, with sessile flowers seated
along a common axis
the male organ of a flower, consisting of a stalk or a filament
and an anther containing pollen
having stamens
rudimentary sterile stamen
star-like or star-shaped with slender segments radiating out
from a common centre
that portion of a pistil or style which receives the pollen
484
stigmatic
stigmatose
stimulant
stipe
stipellae
stipitate
stipular
stipulary
stipule
stomatal crypts
strand
striate
strigose
style
subulate
sulcate
superior
suture
syncarpous
tanniferous
tendril
tenuinucellate
tepal
terete
tessellate
testa
tetrad
theca
thyrse
thyrsoid
tomentellous
tomentose
tomentum
torulose
torus
translucent
trapezoid
triad
trichome
relating to or having the function of a stigma

provided with stigmas or having conspicuous stigmas
a stimulus-producing agent
stalk
a minute stipule on a partial petiole of a compound leaf
having a stipe or a special stalk
having stipules or relating to stipules
occupying the place of stipules
an appendage of a leaf on each side of the leaf-insertion
simple glandular pits or cavities on stomata
a bundle of vascular tissue, resembling a cord
marked with fine longitudinal parallel lines, as grooves or
ridges
beset with sharp-pointed appressed straight and stiff hairs or
bristles
the slender upper part of an ovary supporting a stigma
awl-shaped; narrowing and tapering from the base to a fine
point
grooved or furrowed
(ovary) with all the floral envelopes inserted below it, on the
torus
a line or a junction of two parts immovably connected; line of
dehiscence
composed of two or more united carpels
producing tannin
a slender stem or leaf-outgrowth which functions as a climbing
apparatus
(ovule) with a thin nucellus
free segment of a perianth not differentiated into sepals and
petals
cylindrical and usually tapering, rounded in cross-section
checkered
the outer coat of a seed, usually hard and brittle
a group of four; a body formed of four cells, as in the
formation of pollen in the pollen-mother-cells
a pollen-case
a mixed inflorescence with the main axis indeterminate, and
the secondary and ultimate axes cymose
resembling a thyrse
sparingly or minutely tomentose
densely pubescent with matted wool, or short hairs
pubescence of matted interwoven hairs
cylindrical with swollen portions at intervals; beaded
same as receptacle
transmitting rays of light without being transparent
having a shape of a trapezium
group of three
hair-like outgrowth of the epidermis
485
trichotomous
trifid
trifoliolate
tristylous
truncate
tubercle
tuberculate
turbinate
umbel
undulate
ungulate
uniseriate
unisexual
unitegmic
urceolate
valvate
vein
velutinous
velvety
venation
vermifuge
verrucose
verruculose
versatile
vessel
vestigial
villous
viviparous
zygomorphic
three-forked, branching into three divisions

three-cleft
with three leaflets
with three styles
squared at the tip or base as if cut off with a straight blade
a small rounded protuberance
having tubercles
top-shaped; inversely conical
an indeterminate inflorescence in which a cluster of pedicels
spring from the same point, like the ribs of an umbrella
wavy
hoofed; clawed
in one horizontal row or series
(flower) with either stamens or pistils only, not both
(ovule) having one layer of integument
pitcher-shaped, hollow and contracted at the mouth like an urn
or a pitcher
opening by valves, as in most dehiscent fruits or anthers; when
parts of a flower-bud meet exactly without overlapping
a thread or strand of vascular tissue in a flat organ, as a leaf
velvety due to a coating of fine soft hairs
densely covered with fine short soft erect hairs
the disposition of veins; the mode of veining
that expels intestinal worms
warty
very warty; much covered with warts
swinging freely on its support, as many anthers on their
filaments
any tube or canal with properly defined walls in which fluids
circulate
small and imperfectly developed; the remaining trace of an
organ which fully developed in some ancestral form
covered with long fine hairs
germinating or sprouting from seed or bud while still attached
to the parent plant
not radially symmetrical
486
INDEX TO SCIENTIFIC NAMES

(compiled by Rusea Go )
Names in italics here refer to synonyms; family names given in upper case are those of families
revised in this volume. Pages with illustrations are listed in italics.
Acer
1
Acer caesiaefolium
3
Acer caesium
3
Acer curranii
3
Acer javanicum
3
Acer laurinum
2, 3
Acer niveum
3
Acer philippinum
3
ACERACEAE
V, 1, 453
Acioa heteropetala
163
Acronychia
352, 353, 356, 358, 381
Acronychia apiculata
358
Acronychia arborea
358
Acronychia laurifolia
358
Acronychia pedunculata
358, 359
Acronychia porteri
381, 382
Adenanthera triphysa
424
Adolphia
306
Aegiphila viburnifolia
113
Agathidanthes
253
Agathidanthes javanica
255
Agathis
XLVII, 27, 28
Agathis alba
29
Agathis beccarii
29
Agathis borneensis
XLV, 28, 29, 31
Agathis dammara
29
Agathis endertii
28, 29
Agathis kinabaluensis
29, 30
Agathis latifolia
29
Agathis lenticula
28, 30
Agathis loranthiifolia
29
Agathis macrostachys
29
Agathis orbicula
28, 32
Agathis rhomboidalis
29
Agathisanthes
253
Agathisanthes javanica
255
Agelaea
188
Agelaea borneensis
188
Agelaea insignis
188
Agelaea macrophylla
187, 188
Agelaea trinervis
188
Ailanthus
421, 422, 423
Ailanthus blancoi
423
Ailanthus integrifolia
423
Ailanthus peekelii
423
Ailanthus philippinensis
424
Ailanthus triphysa
423, 424, 425
ALANGIACEAE
V, 5, 199
Alangium
5, 6, 199
Alangium sect. Alangium
6
Alangium sect. Angolam
6
Alangium sect. Canostigma
6
Alangium sect. Marlea
6
Alangium sect. Rhytidandra
6
Alangium begoniaefolium
10, 13
Alangium bogoriense
10
Alangium borneense
10
Alangium chinensis var. tomentosum
10
Alangium circulare
XLIX, 7
Alangium ebenaceum
10
Alangium ebenaceum var. B
10
Alangium ebenaceum var. C
10
Alangium ebenaceum var. D
10
10
Alangium ebenaceum var. E
Alangium ebenaceum var. G
10
Alangium ebenaceum var. tutela
10
Alangium griffithii
7, 8
Alangium havilandii
7, 8
Alangium hirsutum
12
Alangium javanicum
7, 9
Alangium javanicum form B
10
Alangium javanicum form C
10
Alangium javanicum form D
10
Alangium javanicum var. ebenaceum 10, 11
Alangium javanicum var. javanicum 10, 11
Alangium javanicum var. meyeri
10, 11
Alangium kinabaluense
14
Alangium kurzii
7, 10
Alangium lamarckii
12
Alangium longiflorum
6, 12
Alangium meyeri
10
Alangium mezianum
10
487
Alangium nobile
7, 13
Alangium ridleyi
10
Alangium rotundatum
13
Alangium rotundifolium
7, 13
Alangium salvifolium subsp. hexapetalum 12
Alangium scandens`
6
Alangium sessiliflorum
10
Alangium tutela
10
Alangium sp. 1
6
Aleurocanthus woglumi
407
Allanthospermum
422, 421, 427
Allantospermum borneense
XLVII, 427
subsp. borneense
426, 429
subsp. rostratrum
429
Allantospermum multicaule
427
Alphitonia
306, 307
Alphitonia excelsa
308, 309
Alphitonia incana
308
Alphitonia philippinensis
308
Ampacus accedens
386
Ampacus alba
391
Ampacus aromatica
392
Ampacus glabra
390
Ampacus macrophylla
386
Ampacus robusta
390
Ampacus roxburghiana
392
Ampacus triphylla
394
Amyris graveolens
370
Anacardiaceae
XIII, XIV, XVIII,
46, 352, 354, 480
Anacolosa
271, 274
Anacolosa arborea
274
Anacolosa frutescens
272, 274
Anacolosa heptandra
274
Anacolosa luzoniensis
274
Anacolosa maingayi
284
Anacolosa sp.
274
Angelesia
165
Angelesia splendens
165
Anisophyllea
15, 16, 17, 25, 322
Anisophyllea beccariana
17, 18
Anisophyllea chartacea
17, 18
Anisophyllea corneri
17, 19
Anisophyllea disticha
16, 17, 19, 23
Anisophyllea ferruginea
XLVII, 17, 21
Anisophyllea glandulifolia
25
Anisophyllea globosa
17, 20, 22
Anisophyllea impressinervia
17, 22
Anisophyllea nitida
17, 23
Anisophyllea rhomboidea XLIX, 17, 19, 23
ANISOPHYLLEACEAE
V, 15, 16, 322
Anisophylleaceae tribe Anisophylleae
15
Anisophyllum
16
Anisophyllum trapezoidale
19
Anisoptera grossivenia XLV, XLVI, XLVII
Araliaceae
199
Aralidium
199
Araucaria
27
ARAUCARIACEAE
V, 27
Ardisia ochracea
127
Arrabidaea magnifica
33
Atalantia
411
Atalantia disticha
412
Atalantia disticha var. paniculata
412
Atalantia maritima
412
Atalantia nitida
411
Atalantia paniculata
412
Atuna
156, 157, 158
Atuna cordata
L, 158, 159
Atuna elata
161
Atuna excelsa
161
Atuna nannodes
158
Atuna racemosa
158, 160
Atuna racemosa subsp. excelsa
161
Atuna racemosa subsp. racemosa 156, 161
Atuna villamilii
161
Aubletia caseolaris
450
Aucuba
199
Aucuba japonica
199
Averrhoa
287, 289
Averrhoa bilimbi
287, 289
Averrhoa carambola
289
Averrhoaceae
288
Avicennia
324, 450
Barringtonia
421, 442
Begoniaceae
209
Berchemia
306
Bergera koenigii
406
Bhesa
107, 109, 111
Bhesa paniculata
110, 111.112
Bhesa robusta
111, 112
Bignonia indica
38
Bignonia pentandra
38
Bignonia spathacea
37
Bignonia tripinnata
38
BIGNONIACEAE
V, 33
488
Biophytum
287, 288
Biophytum sensitivum
288
Bischofia
454
Bischofiaceae
454
Blatti
447
Blatti acide
450
Blatti alba
449
Blatti caseolaris
450
Blatti pagatpat
450
Bombacaceae
XIII, XVIII
Borneodendron aenigmaticum
XLVIII
Boswellia
46
Boswellia sacra
46
Brackenridgea
257, 258, 259
Brackenridgea sect. Brackenridgea
259
Brackenridgea sect. Notochnella
259
Brackenridgea denticulata
259
Brackenridgea hookeri
259, 261
Brackenridgea palustris
259, 260
Brackenridgea serrulata
260
Brassicaceae
101, 102
Brongniartia
246
Brongniartia coriacea
247
Brucea
421, 422, 429
Brucea amarissima
431
Brucea javanica
428, 429
Brucea sumatrana
431
Brucea sumatrensis
431
Bruguiera
321, 322, 323
Bruguiera 10-angulata
326
Bruguiera angularis
326
Bruguiera australis
326
Bruguiera capensi
324
Bruguiera caryophylloides
324
Bruguiera conjugata
325
Bruguiera cylindrica
324, 325, 326
Bruguiera decandra
335
Bruguiera eriopetala
326
Bruguiera gymnorrhiza
323, 324
Bruguiera malabarica
324, 326
Bruguiera oxyphylla
326
Bruguiera parietosa
326
Bruguiera parviflora
324, 325, 326, 327
Bruguiera rheedii
324
Bruguiera ritchiei
325
Bruguiera rumphii
324
Bruguiera sexangula
323, 326
Bruguiera wighti
324
Bruinsmia
463, 465
Bruinsmia polysperma
465
Bruinsmia styracoides
463, 464, 465
Bulbophyllum
xl
Burkillanthus
261, 352, 353, 357
Burkillanthus malaccensis
261, 360
BURSERACEAE
V, XIV, XIX,
45, 46, 352, 421
Burseraceae tribe Bursereae
46
Burseraceae tribe Canarieae
46
Calophyllum
XIV, XIX
Calophyllum nodosum
XLVIII
Calosanthes indica
38
Campanulaceae
213
Campylocerum
265
Camunium
406
Canariopsis aspera
51
Canarium
45, 46, 47, 48, 65, 84, 174
Canarium sect. Canarium
48
Canarium sect. Pimela
48
Canarium sect. Tenuipyrena
65
Canarium subg. Africanarium
48
Canarium subg. Canariellum
48
Canarium subg. Canarium
48
Canarium acutum
58
Canarium album
46
Canarium angulatum
68
Canarium apertum
46, 50
Canarium asperum
L, 49, 51
Canarium asperum subsp. asperum
51
var. asperum
51
var. clementis
51
Canarium asperum subsp. papuanum
51
Canarium beccarii
59
Canarium brunneum
99
Canarium caudatum
49, 51
Canarium caudatum forma auriculiferum 52
Canarium caudatum forma caudatum
52
Canarium connarifolium
83
Canarium costatum
66
Canarium crassifolium
73
Canarium cuspidatum
73
Canarium decumanum
L, 49, 52, 62, 63
Canarium denticulatum
49, 53
Canarium denticulatum subsp. denticulatum
53
Canarium denticulatum subsp. kostermansii
53
489
Canarium dichotomum
49, 53
Canarium divergens
XLVII, 49, 54
Canarium endertii
53
Canarium expansum
67
Canarium fissistipulum
53
Canarium flavum
58
Canarium fragile
69
Canarium fusco-calycinum
50, 54
Canarium giganteum
58
Canarium glaucum
58
Canarium grandifolium XLVII, 50, 55, 62
Canarium hirsutum
50, 55
Canarium hirsutum subsp. hirsutum
var. hirsutum forma scabrum
56
Canarium hirtellum
62
Canarium hirtipetalum
92
Canarium hispidum
55
Canarium incurvatum
68
Canarium indicum
46, 48
Canarium kedondon
73
Canarium kinabaluensis
49, 56
Canarium korthalsii
97
Canarium kostermansii
49, 56
Canarium kunstleri
53
Canarium laciniatum
53
Canarium laevigatum
90
Canarium latistipulatum
50, 57, 64
Canarium laxum
69
Canarium littorale
49, 50, 57
Canarium littorale forma littorale
58
Canarium littorale forma pruinosum
58
Canarium littorale forma purpurascens 58
Canarium littorale forma rufum
58
Canarium littorale forma tomentosum
58
Canarium luzonicum
46, 48
Canarium megalanthum
48, 50, 58
Canarium merrillii
50, 59
Canarium micrantherum
97
Canarium minahassae
73
Canarium molle
51
Canarium montanum
73
Canarium motleyanum
62
Canarium multifidum
59
Canarium nitens
68
Canarium nitidum
61
Canarium odontophyllum
48, 50, 59, 60
Canarium ovatum
46, 48
Canarium palawanense
59
Canarium parvifolium
61
Canarium patentinervium
49, 61, 63
Canarium pauciflorum
52, 86
Canarium pilosum
49, 50, 61
Canarium pilosum subsp. borneensis
62
Canarium pilosum subsp. pilosum
62
Canarium pilosum var. hirtellum
62
Canarium pimela
48, 63, 64
Canarium planchonii
95
Canarium pruinosum
58
Canarium pseudodecumanum
48, 49, 62
Canarium pseudopatentinervium
49, 63
Canarium pseudopimela
49, 63
Canarium reticulatum
73
Canarium rubiginosum
73, 94
Canarium rufum
58
Canarium rugosum
74
Canarium rugosum var. sumatranum
74
Canarium sarawakanum
50, 64
Canarium serricuspe
58
Canarium serrulatum
50, 58
Canarium subcordatum
55
Canarium villosum
51
Canarium virgatum
74
Canarium vulgare
46, 48
Cantanola
188
CAPPARACEAE
V, 101, 102
Capparaceae subfam. Cleomoideae
102
Capparis
102
Capparis magna
103
Capparis spinosa
101
Capusia
154
Carallia
321, 322, 328
Carallia arguta
330
Carallia borneensis
332, 329, 331
Carallia brachiata
329, 330
Carallia calycina
330
Carallia celebica
330
Carallia cerisopsifolia
330
Carallia confinis
330
Carallia coriifolia
329, 332
Carallia cuprea
330
Carallia cuspidata
330
Carallia floribunda
330
Carallia lucida
330
Carallia mindanaensis
329
Carallia multiflora
330
Carallia scortechinii
330
Carallia spinulosa
330
Carallia timorensis
330
490
Carallia viridifolia
330
Carallia sp. 1
329, 332
Carallia sp. 2
329, 333
Carallia sp. 3
328, 333
Carallia sp. 4
329, 333
Carallia sp. 5
328, 334
Cardiocarpus amarus
442
Cardiophora hindsii
442
Carya
233
Caryospermum
150
Caryospermum philippinensis
150
Cassine
113
Cassine viburnifolia
108, 113, 115
Casuarina equisetifolia
XLIV
Ceanothus
305
Ceanothus asiaticus
311
CELASTRACEAE
V, XIV, XVIII,
107, 322, 453
Celastrales
322
Celastrus
108, 154
Celastrus alpestris
150
Celastrus robustus
112
Celtis angustifolia
318
Celtis grewioides
318
Cenarium
47
Ceratostachys
253
Ceriops
321, 322, 334, 335
Ceriops boviniana
335
Ceriops candolleana
335
Ceriops decandra
335
Ceriops fosteniana
335
Ceriops lucida
336
Ceriops pauciflora
335
Ceriops roxburghiana
335
Ceriops tagal
335, 337
Ceriops timoriensis
336
Ceriops zippeliana
335
Chalcas koenigii
406
Chalcas paniculata
406
Chamaepericlymenum suecicum
199
CHRYSOBALANACEAE
V, 155, 156
Cinnamomum
433
Citrofortunella microcarpa
364
Citrus
351, 352, 353, 355
356, 357, 362, 363, 483
Citrus acida
365
Citrus angulata
395
Citrus aurantifolia
365
Citrus aurantium subsp. saponacea
366
Citrus aurantium var. grandis

364
Citrus aurantium var. medica
363
Citrus aurantium var. sinensis
365
Citrus decumana
364
Citrus deliciosa
364
Citrus grandis
363, 364
Citrus halimii
362, 364
Citrus hystrix
365, 366
Citrus limon
364
Citrus limonum
364
Citrus macroptera
362, 366
Citrus macroptera var. macroptera 366, 367
Citrus madurensis
364
Citrus malaccensis
261
Citrus maxima
364
Citrus medica
363
Citrus medica var. limon
364
Citrus microcarpa
364
Citrus mitis
364
Citrus nobilis
364
Citrus papeda
365
Citrus papuana
366
Citrus reticulata
363, 364
Citrus reticulata var. austera
364
Citrus sinensis
365
Citrus torosa
365
Clausena
352, 353, 354, 356, 369, 371
Clausena calciphila
369, 370
Clausena excavata
368, 369, 370
Clausena excavata var. excavata
370
Clausena javanensis
370
Clausena lansium
369
Clausena wampi
369
Cleome
102
Clethra
181
Clethra canescens
181, 183
Clethra canescens var. canescens
183
Clethra canescens var. clementis
183
Clethra canescens var. ledermannii
183
Clethra canescens var. luzonica
183
Clethra canescens var. novoguinensis 183
Clethra clementis
183
Clethra elongata
184
Clethra longispicata
183, 184
Clethra pachyphylla
182, 183, 184
CLETHRACEAE
V, 181
Clusiaceae
XIV, XLVII
Cnestis
188
Colubrina
305, 306, 311
491
Colubrina anomala
312
Colubrina arborescens
311
Colubrina asiatica
311
Colubrina beccariana
310, 311, 312
Colubrina elliptica
311
Colubrina excelsa
308
Colubrina ferruginosa
311
Combretocarpus
15, 16, 25, 26, 322
Combretocarpus motleyi
25
Combretocarpus rotundatus XLVIII, 24, 25
Combretum
25
Commiphora myrra
46
Commiphora opobalsamum
46
Compositae
480
Coniferae
28
CONNARACEAE
V, 187, 288
Connaropsis
289
Connaropsis acuminata
290
Connaropsis diversifolia
290
Connaropsis glauca
291
Connaropsis grandiflora
290
Connaropsis griffithii
290
Connarus
83, 187, 188, 189, 190
Connarus agamae
L, 189, 190, 193, 195
Connarus borneensis
192
Connarus cullionensis
192
Connarus densiflorus
191
Connarus ellipticus
190
Connarus euphlebius
191
Connarus falcatus
191
Connarus grandis
190
Connarus hebephyllus
192
Connarus impressinervis
190
Connarus jackiana
192
Connarus lucens
191
Connarus monocarpus
191
Connarus monocarpus subsp. malayensis 191
Connarus mutabillis
192
Connarus odoratus
192
Connarus pachyphyllus
191
Connarus plumoso-stellatus
192
Connarus semidecandrus
192
Connarus stellatus
192
Connarus urdanetensis
197
Connarus villosus
192
Connarus winkleri
191
Connarus sp. A.
189, 190
Connarus sp. B.
189, 190
Cookia chlorosperma
372
Cookia cyanocarpa
374
Cookia wampi
369
CORNACEAE
V, 5, 199, 200, 253
Cornales 5, 322
Cornus mas
199
Cotylelobium lanceolatum
XLV
Craterianthus
341
Crateva
101, 102, 103
Crateva brownii
104
Crateva lophosperma
103
Crateva macrocarpa
104
Crateva magna
103, 105
Crateva membranifolia
104
Crateva nurvala
103
Crateva religiosa
103, 104, 105, 106
Cratoxylum
219, 220, 221
Cratoxylum sect. Cratoxylum
221
Cratoxylum sect. Isopterygium
221
Cratoxylum sect. Tridesmos
221
Cratoxylum arborescens
219, 221, 223
Cratoxylum celebicum
226
Cratoxylum cochinchinense
219, 221, 222, 223
Cratoxylum cochinchinense var. calcareum
225
Cratoxylum formosum 219, 221, 223, 224
Cratoxylum glaucum
219, 221, 225
Cratoxylum hypericum
226
Cratoxylum ligustrinum
223
Cratoxylum maingayi
219, 221, 223, 225
Cratoxylum myrtifolium
223
Cratoxylum polyanthum
223
Cratoxylum procerum
225
Cratoxylum sumatranum
221, 226
Cratoxylum sumatranum subsp. blancoi 226
Cratoxylum sumatranum
subsp. sumatranum
226
Crossostylis
322
Cucurbitaceae
209
Cunoniaceae
322, 453
Curtisia
199
Curtisia faginea
199
Curtisiana
65
Curtisiana penangensis
69
Cyclandrophora
157
Cyclandrophora asperula
161
Cyclandrophora elata
161
Cyclandrophora excelsa
161
Cyclandrophora glaberrima
161
492
Cyclandrophora nannodes
159
Cyclandrophora villamilii
161
Dacrydium comosum
XLIX
Dacryodes
46, 47, 65, 76, 84, 87
Dacryodes angulata
68
Dacryodes costata
65, 66, 67
Dacryodes elmeri
L, 65, 67
Dacryodes expansa
66, 67
Dacryodes incurvata
66, 68
Dacryodes kostermansii
71
Dacryodes laxa
65, 68
Dacryodes longifolia
66, 69
Dacryodes longifolia var. penangensis 69
Dacryodes macrocarpa
66, 69
Dacryodes macrocarpa var. genuina
71
Dacryodes macrocarpa var. kostermansii 71
Dacryodes macrocarpa var. macrocarpa 71
Dacryodes macrocarpa var. patentinervia 71
Dacryodes nervosa
66, 72
Dacryodes rostrata
65, 66, 70, 72
Dacryodes rostrata f. cuspidata
65, 73
Dacryodes rostrata f. genuina
73
Dacryodes rostrata f. rostrata
73
Dacryodes rubiginosa
XLIX, 66, 73
Dacryodes rugosa
65, 66, 67, 74
Dacryodes rugosa var. genuina
74
Dacryodes rugosa var. rugosa
74
Dacryodes rugosa var. virgata
74
Dacryodes scandens
62
Dactylocladus stenostachys
XLVIII
Dalrymplea
454
Dalrymplea javanica
460, 461
Dammara
28
Dammara loranthiifolia
29
Dapania
288
Dapania grandifolia
288
Daphniphyllopsis
253
Daphniphyllopsis capitata
255
Datisca
209
DATISCACEAE
V, 209
Deplanchea
35, 36
Deplanchea bancana
33, 34, 36
Deplanchea coriacea
36
Derris
187
Diacaecarpium rotundifolium
13
Diatoma brachiata
330
Dichapetalum tetramerum
196
Diemenia
169
Dilleniales
187
Diplanthera bancana
36
Dipterocarpaceae
XIII, XVIII, 201
Dipterocarpus beccarii
XLV
Dipterocarpus caudiferus
XLVI
Dipterocarpus crinitus XLV, XLVI, XLVII
Dipterocarpus globosus
XLVI, XLVII
Dipterocarpus lowii
XLVII
Dipterocarpus mandus
L
Dipterocarpus pachyphyllus
L
Dipterocarpus rigidus
XLVII
Dipterocarpus sarawakensis
XLVII
Dipterocarpus validus
L
Dipteronia
1
Discaria
306
Dolichandrone
35, 37
Dolichandrone longissima
37
Dolichandrone rheedii
37
Dolichandrone spathacea
33, 35, 37, 39
Dracontomelon cuspidatum
73
Driessenia microthrix
25
Drimys piperita
XLIX
Dryobalanops aromatica
XLVI, XLVII, XLIX
Dryobalanops beccarii
XLVII, XLVIII
Dryobalanops fusca
XLIX
Dryobalanops keithii
L
Dryobalanops lanceolata
XLV, XLVI, XLVIII
Dryobalanops rappa
225
Dryptopetalum coriaceum
339
Duabanga
443, 445, 446
Duabanga borneensis
446
Duabanga grandiflora
446
Duabanga moluccana
444, 445, 446
Duabanga taylorii
446
Durio
XIII, XVIII
Durio bukitrayaensis
XVIII
Dysoxylum dasyphyllum
423
Ebenaceae
XLVII
Elaeocarpaceae
322
Elaeodendron
113
Elaeodendron subrotundum
113
Elepanthus longifolius
197
Ellipanthus
187, 188, 195, 196
Ellipanthus beccarii
196
Ellipanthus beccarii var. beccarii XLIX, 196
Ellipanthus beccarii var. peltatus
196
Ellipanthus burebidensis
197
Ellipanthus luzoniensis
197
493
Ellipanthus mindanaensis
196
Ellipanthus sarawakensis
197
Ellipanthus tomentosus
194, 196
Ellipanthus tomentosus subsp. tomentosus
var. luzoniensis
197
Ellipanthus urdanetensis
197
Ellipanthus vidalii
197
Elodea formosa
224
Elodea sumatrana
226
Embelia urophylla
171
Engelhardia
XLVIII, 233, 235
Engelhardia aceriflora
244
Engelhardia apoensis
37, 235
Engelhardia chrysolepis
242
Engelhardia danumensis 235, 236, 237, 238
Engelhardia fenzelii
242
Engelhardia formosana
242
Engelhardia kinabaluensis
235, 237, 238
Engelhardia lepidota
241
Engelhardia mendalomensis 235, 237, 239,
Engelhardia mersingensis 235, 237, 240, 242
Engelhardia nudiflora
243
Engelhardia palembanica
243
Engelhardia parvifolia
243
Engelhardia permicrophylla
243
Engelhardia philippinensis
244
Engelhardia pterococca
242
Engelhardia pterococca var. aceriflora 244
Engelhardia rigida
235, 237, 238,
239, 240, 241
Engelhardia rigida var. rigida
241
Engelhardia rigida var. subsimplicifolia 240
Engelhardia roxburghiana
234, 235,
237, 240, 241
Engelhardia serrata
235, 237, 242
Engelhardia serrata var. cambodica
243
Engelhardia serrata var. nudiflora
243
Engelhardia serrata var. parvifolia
243
Engelhardia serrata var. serrata
243
Engelhardia spicata
235, 237, 244
Engelhardia spicata var. aceriflora
244
Engelhardia spicata var. colebrookeana 244
Engelhardia spicata var. formosana
242
Engelhardia spicata var. spicata
244
Engelhardia subsimplicifolia
240
Engelhardia wallichiana
242
Engelhardia zambalensis
241
Engelhardtia
233, 234
Ericaceae
XLVII
Ericales
181
Erythropalum
273
Erythropalum scandens
273
Eugenia
XIV, 202
Eugenia bankense
XLVIII
Euodia
384
Euodia accedens
386
Euodia alba
391
Euodia arborea
393
Euodia aromatica
392
Euodia bonwickii
388
Euodia concinna
393
Euodia confusa
389
Euodia glabra
389, 390
Euodia kingii
390
Euodia krukovii
390
Euodia latifolia
392
Euodia lunu-ankenda
392
Euodia macrophylla
386
Euodia malayana
393
Euodia obtusifolia
393
Euodia punctata
393
Euodia ridleyi
389, 390
Euodia robusta
390
Euodia roxburghiana
392
Euodia schullei var. ridleyi
389
Euodia speciosa
388
Euodia suaveolens var. ridleyi
389
Euodia subunifoliolata
394
Euodia tenuistyla
389
Euodia triphylla
394
Euodia triphylla var. pubescens
393
Euodia vilamilii
388
Euonymus
107, 111, 114, 117, 121
Euonymus acuminifolius
117
Euonymus acuminifolius var. borneensis 117
Euonymus alatus
119
Euonymus castaneifolius
116, 117, 118
Euonymus cochinchinensis
117, 118
Euonymus coriaceus
119
Euonymus elmeri
119
Euonymus glandulosus
117, 119
Euonymus javanicus
114, 117, 119
Euonymus micropetalus
119
Euonymus moultonii
118
Euonymus pahangensis
119
Euonymus philippinensis
118
Euonymus viburnifolius
113
Euphorbiaceae
XIV, XVIII, XXXVI,
494
XLVIII, 212, 454

Eurycoma
422, 431
Eurycoma longifolia
421, 430, 433
Eurycoma merguensis
433
Euscaphis
453
Euscaphis japonica
453
Euthemis
258, 260, 263
Euthemis ciliata
263
Euthemis hackenbergii
263
Euthemis leucocarpa
262, 263
Euthemis minor
262, 263
Euthemis obtusifolia
263
Euthemis robusta
263
Evodia accedens
386
Evodia nervosa
386
Exitelia corymbosa
168
Fagaceae
XIV, XIX
Fagara
416
Fagara avicennae
417
Fagara glabra
390
Fagara lunu-ankenda
392
Fagara myriacantha
419
Fagara triphylla
394
Ficus
300
Fortunella
351, 352, 353, 364
Frangula 306
Garcinia cymulosa
330
Garuga
45, 46, 74
Garuga abilo
75
Garuga brunnea
99
Garuga clarkei
75
Garuga floribunda var. floribunda
75, 77
Garuga floribunda var. gamblei
75
Garuga littoralis
75
Geraniales
287
Glyaspermum ramiflorum
302
Glycosmis
352, 353, 354, 356, 371, 372
Glycosmis chlorosperma
371, 372
Glycosmis chlorosperma var. bidiensis 373
Glycosmis chlorosperma var. chlorosperma
373
Glycosmis chlorosperma var. elmeri
373
Glycosmis citrifolia
372
Glycosmis clemensii
374
Glycosmis cyanocarpa
372, 374
Glycosmis cyanocarpa var. platyphylla 374
Glycosmis cyanocarpa
var. sapindoides
376
Glycosmis cymosa
374
Glycosmis elata
376
Glycosmis elmeri
373
Glycosmis greenei var. simplex
375
Glycosmis lanceolata
372, 374
Glycosmis longifolia
374
Glycosmis longisepala
XLIX, 372, 375
Glycosmis macarantha 371, 372, 375, 377
Glycosmis macrophylla
376
Glycosmis malayana
372
Glycosmis montana
375
Glycosmis monticola
372
Glycosmis oliveri
375
Glycosmis parviflora
371, 372
Glycosmis platyphylla
374
Glycosmis sapindoides
372, 376
Glycosmis sapindoides var. sapindoides 376
Glycosmis simplicifolia
375
Glycosmis superba
372, 376
Glyptopetalum
111, 121
Glyptopetalum glandulosum
119
Glyptopetalum palawanense
121
Glyptopetalum quadrangulare XLIX, 120,
121, 122, 124
118
Glyptopetalum scortechinii
Gomphia
257, 258, 265
Gomphia corymbosa
259, 260
Gomphia hookeri
259, 260
Gomphia hookeri var. corymbosa
260
Gomphia microphylla
265
Gomphia oblongifolia
265
Gomphia serrata
264, 265
Gomphia sumatrana
265
GOODENIACEAE
V, 213
Gouania
306, 307
Griselinia
199
Griselinia littoralis
199
Guaiacum abilo
75
Guttiferae
XIV, XIX, XLVII, 219, 220
Gynotroches
321, 322, 323, 336
Gynotroches axillaris
338, 339
Gynotroches dryptopetalum
339
Gynotroches lanceolata
339
Gynotroches micrantha
339
Gynotroches parvifolia
339
Gynotroches puberula
339
Gynotroches reticulata
339
Haloragis disticha
19
Haplolobus
47, 75, 76
Haplolobus beccarii
XLIX, 76
495
Haplolobus bintuluensis
76, 79
Haplolobus borneensis
86
Haplolobus inaequifolius
XLIX, 76, 79
Haplolobus kapitensis
76, 78, 80
Haplolobus leenhoutsii
76, 78, 80
Haplolobus sarawakanus
76, 81
Harmandia
273, 275
Harmandia kunstleri
277
Harmandia mekongensis
276, 277
Harrisonia
421, 422
Harrisonia perforata
422
Hasskarlia
454
Hebonga obliqua
424
Hemisantiria
65
Hemisantiria nitida
68
Hemisantiria rostrata
72
Hemisantiria rugosa
74
Hemisantria ? n. sp.
67
Hippocratea maingayi
128
Hippoxylum indicum
38
Hopea badiifolia
L
Hopea beccariana
XLVII
Hopea bullatifolia
L
Hopea centipeda
L
Hopea dasyrrachis
L
Hopea fluvialis
L
Hopea megacarpa
L
Hopea tenuinervula
XLVII
Hopea treubii
XLVII
Huertea
453
HYPERICACEAE
V, 219, 220
Hypericaceae tribe Cratoxyleae
220
Hypericaceae tribe Hypericeae
220
Hypericaceae tribe Vismieae
220
Hypericum
219, 220
Hypericum arborescens
223
Hypericum cochinchinense
223
Hypericum japonicum
219, 220
Hypericum petiolulatum
219, 220
Icacinaceae
271
Icicaster
83
Icicaster planchonii
95
Ilex daphniphylloides
255
ILLICIACEAE
V, 227
Illiciales
227
Illicium
227, 229
Illicium sect. Badiana
229
Illicium sect. Cymbostemon
229
Illicium sect. Illicium
229
Illicium anisatum
229
Illicium cauliflorum
229, 230, 231
Illicium kinabaluense
229, 230
Illicium religiosum
229
Illicium stapfii
228, 229, 230, 231
Illicium verum
229
Illicium sp.
230
Indovethia
258
Indovethia calophylla
258
Irvingella harmandiana
434
Irvingella malayana
434
Irvingella oliveri
434
Irvingia
421, 422, 433
Irvingia harmandiana
434
Irvingia longipedicellata
434
Irvingia malayana
432, 434
Irvingia oliveri
434
Irvingiaceae
421
Isopterys penangiana
469
Itea javanica
299
Ixonanthaceae
422
Jacaranda filicifolia
33
Jacaranda rhombifolia
33
Jambolifera pedunculata
358
Jambolifera porteri
382
JUGLANDACEAE
V, 233
Juglans
233
Juglans pterococca
242
Kaernbachia
454
Kandelia
321, 322, 339
Kandelia candel
340, 341
Kanilia caryophylloides
324
Kanilia parviflora
325
Kibara
245, 246
Kibara angustifolia
247
Kibara blumei
247
Kibara chartacea
247
Kibara coriacea
246
Kibara cuspidata
247
Kibara depauperata
247
Kibara ellipsoidea
247
Kibara grandifolia
247
Kibara macrophylla
247
Kibara mollis
247
Kibara motleyi
247
Kibara obtusa
L, 246, 247, 249
Kibara serrulata
247
Kibara streimannii
246
Kibara tomentosa
247
496
Kibara trichantha
247
Kokoona 108, 111, 122, 123, 124, 130, 153
Kokoona coriacea
123, 124
Kokoona lanceolata
127
Kokoona leucoclada
123, 124, 125
Kokoona littoralis
123, 124, 125
Kokoona littoralis var. bakoensis
127
Kokoona littoralis var. littoralis
127
Kokoona littoralis var. longifolia
127
Kokoona ochracea
123, 124, 127
Kokoona ovatolanceolata
123, 124, 126, 128, 129
Kokoona reflexa
123, 124, 128
Kokoona sabahana
L, 124, 128
Kokoona scortechinii
127, 128
Koompassia
XLVI
Koompassia excelsa
212
Koompassia malaccensis
187
Kostermanthus
156, 157, 161
Kostermanthus heteropetalus
162, 163
Kostermanthus malayanus
161
Krugiodendron
305
Kurrimia
111
Kurrimia luzonica
112
Kurrimia maingayi
112
Kurrimia minor
112
Kurrimia paniculata
112
Kurrimia pulcherrima
112
Labiatae
35
Lauraceae
201
Laurus caesia
3
Lavellea
283
Leguminosae
187, 212
Lepidobotryaceae
287
Lepidobotrys
287
Lepidocarpa costata
175
Leptospermum flavescens
XLIX
Licania
156, 157, 165
Licania splendens
164, 165
Limonellus angulosus
395
Limonia
351, 352, 353
Limonia aurantifolia
365
Limonia corymbosa
411
Limonia disticha
411
Limonia lucida
406
Limonia minuta
400
Limonia parviflora
372
Litsea palustris
XLVIII
Lobelia
213
Lobelia frutescens
217
Lobelia plumieri
217
Loeseneriella
109
Lophopetalum
108, 111, 123, 129, 130, 131, 153
Lophopetalum beccarianum
130, 131, 132, 133
Lophopetalum coriacea
124
Lophopetalum curtisii
135
Lophopetalum dubium
127
Lophopetalum fimbriatum
138
Lophopetalum fuscescens
134
Lophopetalum glabrum 130, 131, 132, 133
Lophopetalum havilandii
132
Lophopetalum intermedium
134
Lophopetalum javanicum
131, 132, 134
Lophopetalum littoralis
127
Lophopetalum maingayi
127
Lophopetalum multinervium 131, 132, 134
Lophopetalum oblongifolium
134
Lophopetalum oblongum
134
Lophopetalum pachyphyllum
XLIX, 131, 135
Lophopetalum pallidum
108, 131, 135,
Lophopetalum paucinervium
134
Lophopetalum reflexum
128
Lophopetalum rigidum
131, 136
Lophopetalum scortechinii
132
Lophopetalum sessilifolium 130, 131, 137
Lophopetalum subovatum
130, 131, 137
Lophopetalum subsessile
136
Lophopetalum wightianum
131, 138
Lophopetalum winkleri
138
Lunasia
352, 353, 354, 355, 379
Lunasia amara
378, 379
Lunasia amara var. amara
380
Lunasia amara var. babuyanica
380
Lunasia gigantifolia
379
Lunasia reticulata
379
Lusa radja
431
Luvunga
352, 353, 357
Lythraceae
445, 447
Macaranga
212
Maclurodendron
352, 353, 354, 356, 380, 381
Maclurodendron parviflorum
XLIX, 381
Maclurodendron porteri
381, 382, 383
Maclurodendron pubescens
L, 381, 382
Macrosolen rotundatus
25
497
Magnoliaceae
227
Magnoliales
227
Malpighiales
288
Mangifera
XIII, XVIII
Mangium
345
Manotes asiatica
433
Mansoa hymensala
33
Manungala pendula
439
Maranthes
156, 157, 167
Maranthes corymbosa
166, 168
Marlea begonifolia
13
Marlea densiflora
8
Marlea ebenacea
10
Marlea griffithii
8
Marlea nobilis
13
Marlea rotundifolia
13
Mastixia
199, 200, 201
Mastixia subg. Manglesia
200
Mastixia subg. Mastixia
200
Mastixia subg. Mastixia series Alternae 200
Mastixia subg. Mastixia series Oppositae
200
Mastixia subg. Pentamastixia
200
Mastixia subg.Tetramastixia
200
Mastixia acuminatissima
205
Mastixia bracteata
201
Mastixia caesia
205
Mastixia caudatifolia
205
Mastixia clarkeana
208
Mastixia clarkeana var. macrophylla 208
Mastixia cuspidata
201, 202
Mastixia cuspidata var. margarethae
205, 201
Mastixia eugenioides
201, 202
Mastixia glauca
XLIX, 201, 203
Mastixia kimanila
205
Mastixia korthalsiana
206
Mastixia korthalsiana var. macrophylla 208
Mastixia laxa
205
Mastixia laxa var. angustifolia
205
Mastixia macrocarpa
201, 203
Mastixia maingayi
206
Mastixia maingayi var. subtomentosa 206
Mastixia margarathae
205
Mastixia megacarpa
204
Mastixia parvifolia
204
Mastixia pentandra
201, 204
Mastixia pentandra subsp. cambodiana 204
Mastixia pentandra subsp. chinensis
204
Mastixia pentandra subsp. moluccana 204

Mastixia pentandra subsp. pentandra
204
Mastixia pentandra subsp. philippinensis 204
Mastixia pentandra subsp. scortechinii 204
Mastixia pentandra var. cuspidata
201
Mastixia premnoides
208
Mastixia propinqua
206
Mastixia rostrata
201, 204, 205
Mastixia rostrata subsp. caudatifolia
202, 205, 207
Mastixia rostrata subsp. rostrata
205
Mastixia scortechinii
204
Mastixia trichotoma
201, 205
Mastixia trichotoma var. benculuana 208
Mastixia trichotoma var. clarkeana
208
Mastixia trichotoma var. korthalsiana 206
Mastixia trichotoma var. maingayi
206
Mastixia trichotoma var. rhynchocarpa 208
Mastixia trichotoma var. simalurana
208
Mastixia trichotoma var. tenuis
208
Matthaea
245, 246, 248
Matthaea calophylla
248
Matthaea ellipsoidea
248
Matthaea latifolia
248
Matthaea sancta
248, 250
Maytenus
108
Maytenus emarginatus
108
Meesia
265
Meesia serrata
265
Melastomataceae
25
Meliaceae
XXXII, 46, 352, 421
Melicope
104, 352, 353, 355, 384, 385
Melicope accendens
387, 386
Melicope bonwickii
L, 385, 388
Melicope clemensiae
385, 388
Melicope conferta
358
Melicope confusa
385, 389
Melicope denhamii
L, 385, 389
Melicope glabra
385, 390
Melicope hookeri
386, 390
Melicope incana
386, 391
Melicope jugosa
L, 385, 391, 393
Melicope latifolia
385, 392
Melicope lunu-ankenda
386, 392
Melicope sororia
L, 393
Melicope subunifoliolata
L, 385, 394
Melicope tetrandra
414
Melicope triphylla
385, 394
Melicope unifoliolata
382
498
Menispermaceae
XXVIII
Merope
352, 353, 355, 357, 395
Merope angulata
395, 397, 405
Merrillia
352, 353, 354, 356, 396
Merrillia caloxylon
398, 399
Micromelum
352, 353, 354, 356, 389
Micromelum glabrescens
400
Micromelum minutum
400, 402
Micromelum minutum var. minutum
400, 401
Micromelum pubescens
400
Microtropia
138
Microtropis
107, 109, 138, 139, 140
Microtropis argentea
139, 140, 141
Microtropis bicolor
149
Microtropis borneensis 139, 140, 141, 142
Microtropis fascicularis 139, 140, 143, 144
Microtropis grandifolia
139, 140, 144
Microtropis grandifolia var. grandifolia 144
Microtropis grandifolia var. longipetiolatus
144
Microtropis keningauensis
139, 140, 144
Microtropis kinabaluensis 139, 141, 144, 146
Microtropis kinabaluensis var. acuminata
144
Microtropis ovata
139, 140, 145
Microtropis pauciflora
149
Microtropis platyphylla 139, 141, 142, 145
Microtropis rigida
139, 140, 146
Microtropis sabahensis L, 139, 140, 146,147
Microtropis sarawakensis 139, 140, 141, 147
Microtropis sterrophylla
144
Microtropis suborbiculata
149
Microtropis sumatrana
139, 141, 147
Microtropis valida
139, 140, 149
Microtropis wallichiana
139, 140, 149
Millingtonia hortensis
33
Millingtonia pinnata
42
Monanthocitrus
352, 353, 355, 357, 402
Monanthocitrus bispinosa
402
Monanthocitrus oblanceolata
L, 402, 403
MONIMIACEAE
V, 245
Monimiaceae subfam. Mollinedieae
245
Moraceae
XL
Morinda
434
Murraya
352, 353, 354, 356,
371, 396, 398, 405, 406
Murraya sect. Bergera
406
Murraya sect. Murraya
396, 398, 406
Murraya burmanni
370
Murraya caloxylon
398
Murraya exotica
405, 406.407
Murraya foetidissima
406
Murraya koenigii
405, 406
Murraya odorata
406
Murraya paniculata
404, 405, 406, 407
Murraya paniculata var. exotica
406
Murraya paniculata var. paniculata 406, 407
Murraya sumatrana
406
Myricaceae
233
Myrtaceae
XIV, XLVII
Myrtales
156, 322
Neckia
258
Neckia serrata
258
Neolamarckia cadamba
212
Nepenthes
XXXVIII
Nyctocalos
35
Nypa
450
Nypa fruticans
396
Nyssa
199, 253
Nyssa arborea
255
Nyssa bifida
255
Nyssa javanica
253, 254, 255
Nyssa sessiliflora
255
NYSSACEAE
V, 199, 253
Ochanostachys
271, 274, 277
Ochanostachys amentacea
278, 279
Ochanostachys bancana
278
OCHNACEAE
V, 257
Ochnaceae subfam. Ochnoideae
257
Ochnaceae subfam. Sauvagesioideae
257
Ochranthe
454
Octomeles
209
Octomeles sumatrana XLVI, 210, 211, 212
OLACACEAE
V, 271
Olacales
271
Olacineae
271
Olax
273
Olax imbricata
273
Opiliaceae
271
Orchidaceae
XIII, XXXVI
Oreomunnea
234
Oreorhamnus
312
Oroxylum
35, 38
Oroxylum indicum
33, 35, 38, 40
Ouratia angustifolia
265
Ouratia beccariana
265
Ouratia borneensis
265
499
Ouratia megacarpa
265
Ouratia neriifolia
265
OXALIDACEAE
V, 287, 288
Oxalis
287, 288
Oxalis corniculata
288
Oxalis corymbosa
288
Pachylobus
65
Pagapate
447
Paliurus
305, 306
Paliurus spina-christin
305
Paracelastrus
138
Paramignya
352, 353, 357
Paramignya angulata
395
Paramignya longispina
395
Parashorea malaanonan
L
Parashorea tomentella
L
Parastemon
155, 156, 157, 169
Parastemon grandifructus
157, 169, 171
Parastemon spicatum
171
Parastemon urophyllus
169, 170, 171
Parinari
156, 157, 161, 167, 172, 173
Parinari sect. Sarcostegia
167
Parinari subg. Cyclandrophora
157
Parinari subg. Exitelia
167
Parinari subg. Sarcostegia
167
Parinari argenteo-sericea
L, 173
Parinari ashtonii
179
Parinari bicolor
175
Parinari canarioides
173, 174
Parinari costata
173, 175
Parinari costata subsp. costata
175
Parinari costata subsp. rubiginosa
175
Parinari elmeri
173, 177
Parinari gigantea
178
Parinari heteropetala
161
Parinari heteropetalum
163
Parinari metallica
XLVII, 173, 178
Parinari myriandra
161
Parinari nannodes
158
Parinari oblongifolia
173, 176, 178
Parinari rigida
173, 179
Parinari rubiginosum
175
Parinarium
172
Parinarium subg. Petrocarya
172
Parinarium asperulum
161
Parinarium borneensis
178
Parinarium corymbosum
168
Parinarium costatum var. rubiginosum 175
Parinarium elatum
161
Parinarium glaberrimum
161
Parinarium griffithianum
168
Parinarium myriandrum
163
Parinarium scabrum
161
Parinarium villamilii
161
Pellacalyx
321, 322, 323, 341, 343
Pellacalyx axillaris
342, 343
Pellacalyx cristatus
343, 344
Pellacalyx lobbii
343, 344
Pellacalyx symphiodiscus
343, 345
Perrottetia
107, 109, 150
Perrottetia alpestris
150
Perrottetia alpestris subsp philippinensis
148, 150
Perrottetia philippinensis subsp. alpestris150
Perrottetia philippinensis subsp. moluccana
150
Petalinia
277
Petalinia bancana
278
Petrocarya
172
Petrocarya excelsa
161
Phyllocladaceae
28
Phyllocladus hypophyllus
XLIX
Picrasma
421, 422, 435
Picrasma andamanica
435
Picrasma denhamii
389
Picrasma javanica
435, 437
Picrasma nepalensis
435
Picrasma philippinensis
435
Picroxylon siamense
433
Pilocarpus amara
379
Pimela dichotoma
53
Pinus dammara
29
PITTOSPORACEAE
V, 297
Pittosporum
297, 298, 299
Pittosporum accuminatissimum
303
Pittosporum clementis
302
Pittosporum comptum
302
Pittosporum epiphyticum
303
Pittosporum ferrugineum
298, 299
Pittosporum javanicum
299
Pittosporum linearifolium
L, 298, 299, 300, 301
Pittosporum longisepalum
298, 299, 300
Pittosporum nativitatis
299
Pittosporum ramiflorum
298, 299, 302
Pittosporum ramiflorum var. parviflorum
302
Pittosporum resiniferum L, 298, 299, 303
500
Pittosporum rufescens
299
Pittosporum silamense
298, 299, 303
Pittosporum versteeghii
299
Plaesiantha
341
Plaesiantha lobbii
344
Pleiospermium
352, 353, 354, 355, 356, 357, 407, 409
Pleiospermium latialatum
409
Pleiospermium longisepalum 408, 409, 410
Podocarpaceae
XLVII, 28
Polygalaceae
154, 467, 469
Polyosma hookeri
14
Protium
47, 81
Protium connarifolium
L, 82, 83
Protium philippinensis
83
Pseudellipanthus
195
Pseudellipanthus beccarii
196
Pseudellipanthus peltatus
196
Pseuditea javanica
299
Pterocymbium tubulatum
XLVI
Pterospermum javanicum
XLVI
Pterilema
234
Pterilema aceriflorum
244
Quassia
422, 436
Quassia borneensis
XLVII, 436, 439
Quassia indica
421, 438, 439
Quinaria lansium
369
Radermachera
33, 35, 41
Radermachera corymbosa
42
Radermachera lobbii
42
Radermachera lobbii subsp. acuminata 42
Radermachera pinnata
41, 42
Radermachera pinnata subsp. acuminata
42, 43
Radermachera pinnata subsp. pinnata
42
Radermachera ramiflora
41, 42, 44
Radermachera whitfordii
42
Radermachera sp. A.
41, 44
Rafflesia
XIV, XIX
Rafflesiaceae
XIV, XIX
Reissantia
108
Rex amaroris
442
Reynosia
305
RHAMNACEAE
V, 305, 453
Rhamnaceae tribe Colletieae
306
Rhamnaceae tribe Gouanieae
306
Rhamnaceae tribe Rhamneae
306
Rhamnaceae tribe Ventilagineae
306
Rhamnaceae tribe Zizipheae
306
Rhamnus
306, 312
Rhamnus borneensis
313, 315
Rhamnus borneensis var. borneensis
313
Rhamnus cathartica
313
Rhamnus incanus
308
Rhamnus lancifolia
313
Rhizophora
321, 322, 323, 325, 326, 345, 347
Rhizophora apiculata
346, 347
Rhizophora australis
326
Rhizophora candel
341
Rhizophora candelaria
347
Rhizophora caryophylloides
324
Rhizophora caseolaris
449, 450
Rhizophora ceratophylloides
324
Rhizophora conjugata
325, 347
Rhizophora cylindrica
324, 325, 326
Rhizophora decandra
335
Rhizophora eriopetala
326
Rhizophora glomerulata
335
Rhizophora gymnorrhiza
324
Rhizophora lingissima
348
Rhizophora macrorrhiza
348
Rhizophora mangle
348
Rhizophora mucronata
347, 348
Rhizophora mucronata var. stylosa
348
Rhizophora palun
324
Rhizophora parviflora
325
Rhizophora plicata
326
Rhizophora polyandra
326
Rhizophora sexangula
326
Rhizophora stylosa
347, 348
Rhizophora tagal
335
Rhizophora timorensis
335
Rhizophora tinctoria
324
RHIZOPHORACEAE
V, XIV, XVIII, 6, 15, 321, 322, 483
Rhizophoraceae tribe Gynotrocheae
322
Rhizophoraceae tribe Macarisieae
322
Rhizophoraceae tribe. Rhizophoreae
322
Rhizophorales
322
Rhododendron quadrasianum
XLIX
Rhus javanica
429
Rosaceae
155, 156
Rosales
156, 187
Roucheria
289
Roucheria macrophylla
292
Rourea
189, 288
Rourea diversifolia
290
501
Rourea mimosoides
187
Roureopsis
188
Roureopsis acutipetala
188
Roureopsis emarginata
188
Rubiaceae
5, 212, 322, 434
RUTACEAE
V, 351, 352, 421
Rutaceae subfam. Aurantioideae 352, 353
Rutaceae subfam. Flindersioideae
353
Rutaceae subfam. Rhabdodendroideae 353
Rutaceae subfam. Rutoideae
352, 353
Rutaceae subfam. Toddalioideae 352, 353
Rutales
288
Sageretia
306
Sagittipetalum
328
Sagittipetalum mindanaensis
329
Sagittipetalum palawanense
329
Salacia
109, 153
Salacia bartletti
274
Samadera
436
Samadera brevipetala
439
Samadera indica
439
Santalales
271
Santalodes
189
Santalodes diversifolium
290
Santaloides
189
Santiria
45, 46, 47, 76, 83, 84, 93
Santiria sect. Icicopsis
84
Santiria sect. Santiria
84
Santiria apiculata
84, 85, 87, 89
Santiria apiculata var. apiculata
86
Santiria apiculata var. pilosa
86
Santiria beccarii
86
Santiria borneensis
88
Santiria brachystachys
86
Santiria caesia
93
Santiria conferta
85, 86
Santiria conferta var. wrayi
86
Santiria costata
66
Santiria dacryodifolia
85, 87
Santiria fasciculata
74
Santiria glabra
86
Santiria glabrifolia
90
Santiria grandiflora
84, 87
Santiria griffithii
83, 84, 85, 88, 94
Santiria havilandii
95
Santiria impressinervis
85, 89
Santiria kalkmaniana
85, 89, 91
Santiria laevigata
84, 85, 90
Santiria laevigata forma glabrifolia
90
Santiria laevigata forma laevigata

90
Santiria laevigata forma typica
90
Santiria latifolia
93
Santiria laxa
69
Santiria longifolia
69
Santiria macrocarpa
71
Santiria megaphylla
XLVII, 85, 92
Santiria minimiflora
94
Santiria minutiflora
86
Santiria mollis
84, 92
Santiria mollissima
97
Santiria montana
73
Santiria multiflora
97
Santiria nervosa
72
Santiria nigricans
85, 93
Santiria nitida
68
Santiria oblongifolia
84, 85, 93
Santiria pedicellata
94
Santiria pilosa
86
Santiria planchonii
95
Santiria rostrata
72
Santiria rubiginosa
73, 83, 84, 85, 94
Santiria rubiginosa var. latipetiolata
94
Santiria rubiginosa var. pedicellata
94
Santiria rubiginosa var. rubiginosa
95
Santiria rugosa
74
Santiria samarensis
73
Santiria sarawakana
84, 95
Santiria serrulata
50
Santiria tomentosa
84, 88, 95
Santiria violacea
90
Santiria virgata
47
Santiria wrayi
86
Sapindaceae
1, 187, 352, 453, 454
Sapindales
352
Sapindus
376
Sarawakodendron
107, 109, 153
Sarawakodendron filamentosum
XLVII, 151, 153
Sarcodiscus
246
Sarcodiscus chloranthiformis
247
Sarcotheca
287, 288, 289, 290
Sarcotheca acuminata
290
Sarcotheca diversifolia
289, 290
Sarcotheca glauca
290, 291, 293
Sarcotheca macrophylla
XLIX, 290, 292
Sarcotheca oblongifolia
294
Sarcotheca ochracea
290, 294
Sarcotheca rubrinervis
L, 290, 294
502
Sarcotheca subtriplinervis
290
Saxifragaceae
454
Scaevola
213, 214
Scaevola chanii
214, 215
Scaevola frutescens
217
Scaevola koenigii
217
Scaevola leschenaultii
217
Scaevola macrocalyx
217
Scaevola merrillii
216
Scaevola merrillii var. mollis
216
Scaevola micrantha
L, 214, 216
Scaevola muluensis
214, 217
Scaevola pedunculata
216
Scaevola piliplena
217
Scaevola plumieri
214
Scaevola sericea
214, 217
Scaevola taccada
217
Scaevola verticillata
214, 218
Schisandraceae
227
Schismatoglottidinae
XXXVIII
Schuurmansia
257, 258, 276,
Schuurmansia angustifolia
268
Schuurmansia borneensis
268
Schuurmansia elegans
266, 268
Schuurmansia parviflora
268
Schuurmansiella
258, 268
Schuurmansiella angustifolia
XLIX, 268, 269
Sciadicarpus
246
Sciadicarpus brongniartii
247
Sclerostylis lanceolata
374
Sclerostylis spinosa
395
Scorodocarpus
274, 278
Scorodocarpus borneensis
271, 280, 281
Scrophulariaceae
35
Scutinanthe
45, 46, 47, 97
Scutinanthe brunnea
96, 97, 99
Severinia
352, 353, 354, 356, 357, 410, 411, 412
Severinia disticha
410, 411, 412, 413
Severinia paniculata
410, 411, 412
Shorea acuta
XLVII
Shorea agamii subsp. diminuta
L
Shorea albida
XLVI, XLVIII, 26, 225
Shorea almon
L
Shorea alutacea
XLIX
Shorea asahi
L
Shorea bakoensis
XLIX
Shorea chanii
L
Shorea collaris
L
Shorea crassa
XLVII
Shorea curtisii
XLVI, XLVII, XLIX
Shorea cuspidata
XLVII, XLIX
Shorea dasyphylla
XLIX
Shorea dealbata
XLIX
Shorea elliptica
XLVII, XLIX
Shorea falcifera
XLVI, XLVII, XLIX
Shorea falciferoides
L
Shorea flaviflora
XLVIII
Shorea flemmichii
XLVII
Shorea geniculata
XLVII
Shorea iliasii
L
Shorea induplicata
XLIX
Shorea kunstleri
XLVII
Shorea ladiana
XLVII
Shorea laevis
XLV, XLVIII
Shorea laxa
XLVII
Shorea lunduensis
XLIX
Shorea macroptera
XLVI, XLVII
Shorea multiflora
XLVIII
Shorea ovata
XLVII
Shorea pallidifolia
XLIX
Shorea parvifolia
XLV, XLVI, XLVII, XLVIII, L
Shorea resinosa
XLIX
Shorea richetia
XLIX
Shorea rubella
XLVII
Shorea slootenii
XLVII
Shorea splendida
XLIX
Shorea stenoptera
XLIX
Shorea subcylindrica
XLIX
Shorea superba
XLVI
Shorea symingtonii
L
Shorea waltonii
L
SIMAROUBACEAE V, 46, 352, 421, 422
Siphonodon
107, 108, 109, 154
Siphonodon celastrineus
152, 154
Siphonodon pyriformis
154
Smythea
306, 307
Solenospermum
130
Solenospermum apiculatum
137
Solenospermum aquatile
134
Solenospermum javanicum
134
Solenospermum littorale
127
Solenospermum oblongifolium
134
Solenospermum pallidum
135
Solenospermum paucinervium
134
Solenostigma angustifolium
318
503
Sonneratia
443, 445, 447
Sonneratia acida
450
Sonneratia alba
449
Sonneratia alba x Sonneratia caseolaris 447
Sonneratia alba x Sonneratia ovata
447
Sonneratia caseolaris
443, 447, 449, 450,
Sonneratia evenia
450
Sonneratia lanceolata
450
Sonneratia neglecta
450
Sonneratia obovata
450
Sonneratia ovalis
450
Sonneratia ovata
443, 447, 448, 449, 450
Sonneratia pagatpat
450
SONNERATIACEAE
V, 443, 445, 446,
Soulamea
421, 422, 441
Soulamea amara
421, 440, 442
Spathodea campanulata
33
Stalagmites lamponga
330
Staphylea
453
STAPHYLEACEAE
V, 453, 454
Stemonurus frutescens
274
Sterculiaceae
212
Stixis
101, 102
Stixis ovata
102
Strombosia
271, 273, 274, 283
Strombosia ceylanica
283, 284
Strombosia javanica
282, 283
Strombosia latifolia
284
Strombosia lucida
284
Strombosia maingayi
284
Strombosia multiflora
284
Strombosia rotundifolia
284
STYRACACEAE
V, 463
Styrax
463, 465
Styrax agreste
463
Styrax benzoin
463
Styrax javanicum
9
Styrax officinalis
465
Symmetria
328
Symmetria obovata
330
Symplocos
463
Syzygium
XIV, XIX
Syzygium bankense
XLVIII
Tabebuia rosea
33
Taeniochlaena
188
Tapiscia
453
Tecoma stans
33
Temminckia
213
Temminckia micrantha
216
Terminthodia
412
Terminthodia viridiflora
414
Tetracrypta
16
Tetractomia
352, 353, 355, 412
Tetractomia beccarii
414
Tetractomia holttumi
414
Tetractomia latifolia
414
Tetractomia montana
414
Tetractomia obovata
414
Tetractomia parviflora
414
Tetractomia tetrandrum
414, 415
Tetradium
384
Tetramelaceae
209
Tetrameles
209
Tetramerista
257
Tetrameristaceae
257
Theaceae
XLVII
Toddalia 352, 353, 355
Tricaphis
454
Triceros
454
Trichocarya
169
TRIGONIACEAE
V, 467
Trigoniastrum
467
Trigoniastrum hypoleucum
467, 468
Trigonochlamys
83
Trigonochlamys grandifolia
55
Trigonochlamys griffithii
88
Triomma
45, 46, 99
Triomma malaccensis
98, 100
Triphasia
352, 351, 353
Triplochiton scleroxylon
212
Turpinia
453, 454, 455
Turpinia borneensis
L, 455, 456
Turpinia calciphila
455, 456, 457
Turpinia grandis
455, 456, 457
Turpinia latifolia
460, 461
Turpinia laxiflora
460, 461
Turpinia montana var. borneensis
456
Turpinia nepalensis
455
Turpinia nitida
L, 454, 455, 456, 459
Turpinia sphaerocarpa 455, 456, 460, 462
Turpinia sphaerocarpa var. microcerotis
458, 461
Turpinia sphaerocarpa var. sphaerocarpa 461
Turpinia stipulacea
454, 455, 456, 461
Umbelliferae
476
Upuna borneensis
XLIX
Vatica
201
Vatica endertii
L
504
Vatica compressa
XLIX
Vatica dulitensis
XLVIII
Vatica granulata
L
Vatica pedicellata
XLIX
Vatica umbonata
XLVIII
Ventilago
306, 307
Verbenaceae
35
Viburnum
113
Vitex premnoides
208
Weinmannia blumei
XLIX
Wenzelia
402
Winteraceae
227
Xanthomyrtus spp.
XLIX
Xanthophyllum
154, 171, 469
Xanthophyllum subglobosum
154
Ximenia
273
Ximenia americana
271, 273
Ximenia borneensis
281
Zanthoxylon aromaticum
392
Zanthoxylum
352, 353, 354, 355, 416
Zanthoxylum avicennae
416, 417
Zanthoxylum diversifolium
417
Zanthoxylum iwahigense
417
Zanthoxylum macrophyllum
386
Zanthoxylum myriacanthum 416, 418, 419
Zanthoxylum nitidum
417
Zanthoxylum roxburghianum
392
Zanthoxylum scandens
417
Zanthoxylum triphyllum
394
Ziziphus
306, 314,
Ziziphus angustifolius
316, 317, 318
Ziziphus borneensis
317
Ziziphus calophylla
318
Ziziphus crebrivenosa
318
Ziziphus cumingiana
317
Ziziphus forbesii
318
Ziziphus grewioides
318
Ziziphus havilandii
317
Ziziphus horsfieldii
317
Ziziphus inermis
318
Ziziphus jujuba
314
Ziziphus mauritiana
314
Ziziphus suluensis
318
505
INDEX TO VERNACULAR NAMES

(compiled by Rusea Go)
Scientific names in brackets are the correct names corresponding to vernacular names listed here.
aam (Gomphia serrata (Gaertn.) Kanis)

adas china (Illicium verum Hook. f.)
akar malam (Agelaea macrophylla (Zoll.) Leenh.)
ala (Illicium stapfii Merr.)
alan (Shorea albida Symington)
ambibiliw (Kibara coriacea (Blume) Tulasne)
ambun ambun (Anisophyllea disticha (Jack) Baill.)
antimagas gimbaan (Gomphia serrata (Gaertn.) Kanis)
ara bukit (Pittosporum ferrugineum Aiton)
aremajuh (Sarcotheca glauca (Hook. f.) Hallier f.)
asam-asam (Dacryodes macrocarpa (King) H.J. Lam)
asam daham (Sarcotheca glauca (Hook. f.) Hallier f.)
asam piai (Sarcotheca glauca (Hook. f.) Hallier f.)
asem-asem (Sarcotheca rubrinervis Hallier f.)
atap (Trigoniastrum hypoleucum Miq.)
bajan (Kokoona spp.)
bajan paya (Kokoona ovatolanceolata Ridl.)
bakau (Rhizophora spp.)
bakau aleh aleh (Kandelia candel (L.) Druce)
bakau kurap (Rhizophora mucronata Lam.)
bakau lali (Ceriops decandra (Griff.) Ding Hou)
bakau minyak (Rhizophora apiculata Blume)
balak bekatan (Combretocarpus rotundatus (Miq.) Danser)
balik angin (Alphitonia exelsa (Fenzl) Reiss ex Endl.)
balimbing (Picrasma javanica Blume)
bangkawang (Maranthes corymbosa Blume)
bangkita (Rhizophora apiculata Blume)
bantis (Mastixia trichotoma Blume)
barat-barat (Cassine viburnifolia (Juss.) Ding Hou)
bawang hutan (Scorodocarpus borneensis (Baill.) Becc.)
bedara (Eurycoma longifolia Jack)
bedung (Pittosporum resiniferum Hemsl.)
belian landak (Anisophyllea ferruginea Ding Hou)
belimbing (Averrhoa spp.)
belimbing bulat (Sarcotheca diversifolia (Miq.) Hallier f.)
belimbing manik (Sarcotheca macrophylla Blume)
benuang (Octomeles sumatrana Miq.)
benung kasung (Duabanga moluccana Blume)
berus berus (Kandelia candel (L.) Druce)
berus kurong (Bruguiera gymnorrhiza (L.) Lam.)
berus lenggadai (Bruguiera parviflora (Roxb.) Wight & Arn. ex Griff.)
berus merah (Bruguiera gymnorrhiza (L.) Lam.)
berus ngayong (Bruguiera cylindrica (L.) Blume)
506
267
229
187
230
XLVI, 26, 265
247
21
267
300
292
71
292
292
295
469
122
128
345
341
348
335
348
26
308
436
168
348
208
114
281
433
303
21, 275
289
291
292
210
446
341
325
326
325
324
berus puteh (Bruguiera cylindrica (L.) Blume)

324
berus putut (Bruguiera cylindrica (L.) Blume)
326
beus (Bruguiera spp; Kandelia candel (L.) Druce)
323, 324, 326, 341
biansu gunong (Mastixia cuspidata Blume)
202
biku-biku (Bhesa paniculata Arn.)
112
bindang (Agathis borneensis Warb.)
29
binkuli (Oroxylum indicum (L.) Vent.)
41
binong (Octomeles sumatrana Miq.)
210
binuang (Octomeles sumatrana Miq.)
210, 212
binutan (Radermachera pinnata (Blanco) Merr.)
42
bogua (Harrisonia perforata (Blanco) Seemen)
422
bowlong (Illicium stapfii Merr.)
230
buah piang (Sarcotheca diversifolia (Miq.) Hallier f.)
291
bulok (Agathis endertii Meijer Drees & Agathis orbicula de Laub.)
30, 32
bulu (Agathis borneensis Warb.)
29
bulu bulu (Gynotroches axillaris Blume)
339
bunga lawang (Illicium verum Hook. f.)
229
butun laut (Scaevola sericea Vahl)
218
chinaga-lampong (Gomphia serrata (Gaertn.) Kanis)
267
dadam (Alangium havilandii Bloemb.)
9
dajak (Anisophyllea ferruginea Ding Hou)
21
danguh (Pellacalyx axillaris Korth.)
344
dat (Cratoxylum arborescens (Vahl) Blume)
223
dat tetong (Cratoxylum formosanum (Jack) Dyer)
224
daun kari (Murraya koenigii (L.) Spreng.)
406
derum (Cratoxylum spp.)
219, 220
diit (Cratoxylum arborescens (Vahl) Blume)
223
empudu (Bruinsmia styracoides Boerl. & Koord.)
465
engkop engkop (Anisophyllea nitida Madani)
23
entalun (Engelhardia serrata Blume)
243
219, 224
entemu (Cratoxylum formosanum (Jack) Dyer)
entrang (Turpinia sphaerocarpa Hassk.)
461
gadong hutan (Alangium griffithii (Clarke) Harms)
8
geroking (Cratoxylum arborescens (Vahl) Blume)
223
geronggan lompong (Cratoxylum glaucum Korth.)
225
geronggang (Cratoxylum spp.)
219, 220, 223, 224, 225
geronggang gajah (Cratoxylum arborescens (Vahl) Blume)
223
geronggang puteh (Cratoxylum glaucum Korth.)
225
geronggang timau (Cratoxylum glaucum Korth.)
225
gerunggang (Cratoxylum maingayi Dyer)
226
gilas (Parastemon urophyllus (Wall. ex A.DC.) A.DC.)
172
gimurai (Oroxylum indicum (L.) Vent.)
41
gitan gizu (Sarcotheca glauca (Hook. f.) Hallier f.)
292
iba jantan (Sarcotheca diversifolia (Miq.) Hallier f.)
291
iba talon (Sarcotheca rubrinervis Hallier f.)
295
icerawas burung (Dacryodes macrocarpa (King) H.J. Lam var. macrocarpa)
71
idat (Cratoxylum arborescens (Vahl) Blume)
223
ikor mata (Sarcotheca ochracea Hallier f.)
294
imah (Ochanostachys amantacea Mast.)
278
507
ionadiandau (Eurycoma longifolia Jack)

433
ira prumpuan (Sarcotheca rubrinervis Hallier f.)
295
itan beruang (Mastixia trichotoma Blume)
208
jadam (Alangium javanicum (Blume) Wangerin)
9
jadam paya (Alangium havilandii Bloemb.)
9
jaloot (Brucea javanica (L.) Merr.)
431
jambul merah (Jacaranda rhombifolia G.F.W. Meijer)
33
jampang (Tetractomia tetrandrum (Roxb.) Merr.)
414
jampang rusa (Tetractomia tetrandrum (Roxb.) Merr.)
414
jenangan (Alangium havilandii Bloemb.)
9
jerit (Anacolosa frutescens (Blume) Blume)
275
jiwang (Sarcotheca diversifolia (Miq.) Hallier f.)
291
kacang-kacang (Quassia indica (Gaertn.) Noot.)
441
kait-kait (Harrisonia perforata (Blanco) Merr.)
422
kajo jelan (Cratoxylum formosanum (Jack) Dyer)
224
kambayau (Dacryodes rostrata (Blume) H.J. Lam f. rostrata)
73
kandis batu (Gynotroches axillaris Blume)
339
kandis daham (Sarcotheca glauca (Hook. f.) Hallier f.)
292
kata mudung (Cratoxylum arborescens (Vahl) Blume)
223
kayu ajung (Parastemon grandifructus Prance)
171
kayu jadi (Agathis borneensis Warb.)
29
kayu labu (Ziziphus angustifolius (Miq.) Hatusima ex Steenis)
319
kayu pahit (Picrasma javanica Blume)
436
kayu runap (Anisophyllea disticha (Jack) Baill.)
21
kayu tom (Combretocarpus rotundatus (Miq.) Danser)
26
kayu tulang (Allanthospermum borneensis Forman)
429
kayu wulu (Mastixia pentandra Blume)
204
kebuan (Crateva magna (Lour.) DC.)
104
46, 47, 65, 71, 73, 84, 97, 100
kedondong (Canarium spp.; Dacryodes spp.)
kedondong asam (Triomma malaccensis Hook. f.)
100
keladang (Gomphia serrata (Gaertn.) Kanis)
267
kelaju (Dolichandrone spathacea (L. f.) K. Schum.)
37
kelalud (Gynotroches axillaris Blume)
339
kelana (Ellipanthus beccarii Pierre var. peltatus (Schellenb.) Leenh.)
196
kelapahit (Picrasma javanica Blume; Quassia indica (Gaertn.) Noot.)
441
kelin (Ellipanthus tomentosus Kurz)
197
kelutak (Gomphia serrata (Gaertn.) Kanis)
267
kembayau (Dacryodes spp.; Dacryodes rostrata (Blume) H.J. Lam)
65, 73
kemenyan (Stryrax benzoin Dryand.)
463
kemuning (Murraya paniculata (L.) Jack)
407
kemuning hutan (Carallia borneensis Oliver)
329
kenari (Canarium vulgare Leenh.)
46, 47
kepajang (Anisophyllea ferruginea Ding Hou)
21
kepajang landak (Anisophyllea ferruginea Ding Hou)
21
kerakas payau (Gynotroches axillaris Blume)
339
kerakas payoh (Gynotroches axillaris Blume)
339
keramoh (Daryodes rostrata f. cuspidata (Blume) H.J. Lam)
65, 73
kerantai (Santiria spp.)
83
kerapa-kerapa (Sarcotheca diversifolia (Miq.) Hallier f.)
291
508
keruas (Dacryodes macrocarpa (King) H.J. Lam var. macrocarpa)

keruntum (Combretocarpus rotundifolius (Miq.) Danser)
kimayau (Dacryodes spp.)
kirap (Cratoxylum glaucum Korth.)
kolambang (Gomphia serrata (Gaertn.) Kanis)
kolintuhan (Clethra spp.)
kondolon (Alangium spp.)
kuinin (Brucea javanica (L.) Merr.)
kujuk langit (Radermachera pinnata (Blanco) Seemen)
kukualang (Harrisonia perforata (Blanco) Merr.)
kupi kupi (Gynotroches axillaris Blume)
kurong (Bruguiera gymnorrhiza (L.) Lam.)
laba (Turpinia calciphila J.T. Pereira)
labak (Kibara coriacea (Blume) Tulasne)
labakan (Cratoxylum arborescens (Vahl) Blume)
ladin (Gomphia serrata (Gaertn.) Kanis)
ladit (Anacolosa frutescens (Blume) Blume)
lampyos (Sarcotheca rubrinervis Hallier f.)
langarai (Bruguiera parviflora (Roxb.) Wight & Arn. ex Griff.)
lanjing lanjing (Bruguiera parviflora (Roxb.) Wight & Arn. ex Griff.)
lemeng (Octomeles sumatrana Miq.)
lenggadai (Bruguiera spp.)
lia (Trigoniastrum hypoleucum Miq.)
limau besar (Citrus grandis (L.) Osbeck)
limau buaya (Merope angulata (Willd.) Swingle)
limau hantu (Burkillanthus malaccensis (Ridl.) Swingle;
Citrus macroptera Montr.)
limau hutan (Citrus halimii Stone; Citrus macroptera Montr.)
limau kesturi (Citrus microcarpa Bunge)
limau laut (Merope angulata (Willd.) Swingle)
limau limau (Pleiospermium longisepalum Swingle)
limau manis (Citrus reticulata Blanco; Citrus sinensis (L.) Osbeck)
limau mata kerbau (Citrus limon (L.) Burm. f.)
limau nipis (Citrus aurantifolia (Christm. & Panz.) Swingle)
limau purut (Citrus hystrix DC.)
limau susu (Citrus medica L.)
limo antu (Pleiospermium latialatum Swingle)
limo bali (Pleiospermium latialatum Swingle)
limo to (Pleiospermium latialatum Swingle)
linggayong (Kandelia candel (L) Druce)
linggayong laut (Kandelia candel (L) Druce)
longugan (Illicium kinabaluensis A.C.Sm.)
maba (Turpinia sphaerocarpa Hassk.)
machit laling (Cratoxylum cochinchinense (Lour.) Blume)
madak (Cratoxylum arborescens (Vahl) Blume)
magas (Duabanga moluccana Blume)
majang-majang (Gomphia serrata (Gaertn.) Kanis)
mamungal (Quassia borneensis Noot.)
manat (Cratoxylum arborescens (Vahl) Blume)
509
71
26
65
225
267
181
5
431
42
422
339
325
457
247
223
267
275
295
326
326
210
323, 326
469
364
396
362, 366
366, 410
364
396
410
364, 365
364
365
365
363
409
409
409
341
341
230
461
224
223
446
246
439
223
mandailas (Parastemon grandifructus Prance;

Parastemon urophyllus (Wall. ex A.DC.) A. DC.)
171
manggilan (Agathis borneensis Warb.)
29
manuggal (Quassia borneensis Noot.; Quassia indica (Gaertn.) Noot.)
441
mara (Brucea javanica (L.) Merr.)
431
marapangi (Alangium kurzii Craib; Alangium rotundifolium (Hassk.) Bloemb.)
12, 14
mata buaya (Bruguiera sexangula (Lour.) Poir)
326
mata ulat (Kokoona spp.)
122
mayam kampong (Alangium griffithii (Clarke) Harms)
8
medang (Lauraceae species; Sarcotheca glauca (Hook. f.) Hallier f.)
201, 292
medang kanigara (Mastixia trichotoma Blume)
208
medang pahit (Quassia borneensis Noot.)
439
medang rawang (Tetratomia tetrandrum (Roxb.) Merr.)
414
medang surungan (Mastixia pentandra Blume)
204
melan (Cratoxylum formosanum (Jack) Dyer)
224
mempulak (Crateva magna (Lour.) DC.)
104
mendailas (Parastemon grandifructus Prance)
172
menengang (Anisophyllea spp.)
16
mengilas (Parastemon urophyllus (Wall. ex A.DC.) A.DC.)
172
mengilas babi (Trigoniastrum hypoleucum Miq.)
469
mengkudu (Irvingia malayana Oliver ex A.W. Benn.; Morinda spp.)
434
meransi (Carallia sp. 1)
332
merawai (Carallia sp. 1)
332
merbatu (Atuna spp.; Kostermanthus heteropetalus (Scort. ex King) Prance;
Maranthes corymbosa Blume; Parinari spp.)
157, 165, 168, 172
341
merbuloh (Pellacalyx spp.)
merinang (Ellipanthus beccarii Pierre var. peltatus (Scheenb.) Leenh.)
196
mertama (Anisophyllea corneri Ding Hou; Anisophyllea disticha (Jack) Baill.)
16, 19, 21
mertilan (Cratoxylum arborescens (Vahl) Blume;
Cratoxylum cochinchinense (Lour.) Blume)
223, 224
miapa (Trigoniastrum hypoleucum Miq.)
469
midong (Alangium spp.)
5
mirinos (Cratoxylum formosanum (Jack) Dyer)
224
momon (Engelhardia serrata Blume)
243
mopu (Anisophyllea spp.)
16
murai (Oroxylum indicum (L.) Vent.)
41
mutun (Combretocarpus rotundatus (Miq.) Danser)
26
ngali-nut (Canarium indicum L.)
46
ngayong (Bruguiera cylindrica (L.) Blume)
324
ngilas (Parastemon grandifructus Prance)
171
ngilis (Trigoniastrum hypoleucum Miq.)
469
nonok (Pittosporum ferrugineum Aiton)
300
nuad-mandau (Eurycoma longifolia Jack)
433
nyalin (Allantospermum borneense Forman; Trigoniastrum hypoleucum Miq.)
429, 469
nyalin bahe (Cratoxylum formosanum (Jack) Dyer)
224
nyalin bintek (Trigoniastrum hypoleucum Miq.)
469
obar-obar (Colubrina beccariana Warb.)
312
odok-odok (Colubrina beccariana Warb.)
312
owl (Engelhardia serrata Blume)
243
510
pahau pahau (Gynotroches axillaris Blume)

pait-pait (Brucea javanica (L.) Merr.; Quassia borneensis Noot.)
pamatudon (Canarium decumanum Gaertn.)
pangos (Crateva magna (Lour.) DC.)
panguban (Picrasma javanica Blume)
para bileh (Pleiospermium latialatum Swingle)
parang nyabor (Oroxylum indicum (L.) Vent.)
parang pamol (Oroxylum indicum (L.) Vent.)
paserujan gunugo (Pellacalyx lobbii (Hook. f.) Schimp.)
pasil-pasil (Ziziphus angustifolius (Miq.) Hatusima ex Steenis)
pasoh-pasoh (Dacryodes macrocarpa (King) H.J. Lam)
pati yata (Alphitonia exelsa (Fenzl) Reiss ex Endl.)
patok entilit (Mastixia rostrata Blume)
patok tilan (Cratoxylum formosanum (Jack) Dyer; Cratoxylum maingayi Dyer)
patoli entelit (Irvingia malayana Oliver ex. A.W. Benn.)
pauh kijang (Irvingia malayana Oliver ex. A.W. Benn.)
payas (Brucea javanica (L.) Blume)
pechi mata (Sarcotheca ochracea Hallier f.)
pedada (Sonneratia alba J. Smith; Sonneratia caseolaris (L.) Engl.)
pei (Anisophyllea beccariana Baill.)
perapan macas (Sarcotheca diversifolia (Miq.) Hallier f.)
perdu (Acer laurinum Hassk.)
perepat (Sonneratia alba J. Smith; Sonneratia caseolaris (L.) Engl.)
perepat hutan (Combretocarpus rotundatus (Miq.) Danser)
perepat paya (Combretocarpus rotundatus (Miq.) Danser)
perepat perepat (Combretocarpus rotundatus (Miq.) Danser)
perupok (Lophopetalum spp.)
perupok kuning (Kokoona ochracea (Elmer) Merr.)
petaling (Ochanostachys amentacea Mast.)
piang (Sarcotheca spp.; Sarcotheca glauca (Hook. f.) Hallier f.)
pidang (Cratoxylum glaucum Korth.)
pinggoh (Sarcotheca rubrinervis Hallier f.)
pinguh (Sarcotheca rubrinervis Hallier f.)
pokudata (Alphitonia excelsa (Fenzl) Reiss ex Endl.)
priabu (Mastixia trichotoma Blume)
pusing-pusing (Engelhardia serrata Blume)
putut (Bruguiera gymnorrhiza (L.) Lam.; Bruguiera sexangula (Lour.) Poir)
quintalai (Gomphia serrata (Gaertn.) Kanis)
rabong (Carallia brachiata (Lour.) Merr.)
radipah (Carallia brachiata (Lour.) Merr.)
raja tugag (Cratoxylum formosanum (Jack) Dyer)
rangkas-rangkas (Sarcotheca glauca (Hook. f.) Hallier f.)
rasak batu (Kostermanthus heteropetalus (Scot. ex King) Prance)
rasu (Pellacalyx lobbii (Hook. f.) Schimp.)
rawang (Maclurodendron porteri (Hook. f.) T.G. Hartley)
rawang mata (Tetractomia tetrandrum (Roxb.) Merr.)
rawang paya (Tetractomia tetrandrum (Roxb.) Merr.)
rogam (Sonneratia ovata Backer)
sabal (Dacryodes macrocarpa (King) H.J. Lam var. patentinervia Leenh.)
511
339
431
53
104
436
409
41
41
345
319
71
308
434
224, 226
205
433, 434
431
294
449, 450
18
291
3
449, 450
26
26
26
129
128
278
289, 292
225
295
295
308
208
243
325, 326
267
330, 333
330
224
292
165
345
382
414
414
451
71
sabutun (Combretocarpus rotundatus (Miq.) Danser)

26
sagad-berauh (Ochanostachys amentacea Mast.)
278
sagan-berauh (Scorodocarpus borneensis (Baill.) Becc.)
281
saipang (Pittosporum ferrugineum Aiton)
300
salang (Agathis borneensis Warb.)
29
salangugapit (Anacolosa frutescens (Blume) Blume)
275
salong banggi (Dacyodes rostrata var. cuspidata (Blume)H.J. Lam)
73
sambar bubu (Gynotroches axillaris Blume; Pellacalyx spp.)
341
sampaluan (Licania splendens (Korth.) Prance)
167
sangkuak (Anisophyllea ferruginea Ding Hou)
21
sansanglang (Engelhardia spp.)
233
santikal (Ochanostachys amentacea Mast.)
278
sasgah (Crateva magna (Lour.) DC.)
104
satu inchi (Alangium javanicum (Blume) Wangerin)
9
sawar bubu (Pellacalyx spp.)
339, 341
sawih (Duabanga moluccana Blume)
446
sebelu (Crateva magna (Lour.) DC.)
104
segera (Glycosmis chlorosperma (Blume) Spreng.)
373
seladah (Dacryodes spp.; Dacryodes macrocarpa (King) H.J. Lam
var. patentinervia Leenh.; Santiria spp.)
65, 71, 73, 83
selangan biabas (Cratoxylum cochinchinense (Lour.) Blume)
224
selangan tandok (Irvingia malayana Oliver ex A.W. Benn.)
434
sempetan (Mastixia eugenioides Matthew)
202
sengkanyat (Eurycoma longifolia Jack)
433
sengkayap (Eurycoma longifolia Jack)
433
serang (Crateva magna (Lour.) DC.; Melicope spp.)
104, 384
seraya (Shorea curtisii Dyer ex King)
XLVI
serunai (Bhesa paniculata Arn.)
112
serungan (Cratoxylum arborescens (Vahl) Blume;
Cratoxylum cochinchinense (Lour.) Blume;
Cratoxylum sumatranum (Jack) Blume)
219, 223, 224, 226
serungan batu (Cratoxylum spp.)
219
serungan labakan (Cratoxylum arborescens (Vahl) Blume)
223
serungan mampat (Cratoxylum sumatranum (Jack) Blume)
226
sibut (Dacryodes macrocarpa (King) H.J. Lam var. patentinervia Leenh.)
71
sidodot (Cratoxylum formosanum (Jack) Dyer)
224
simun (Bhesa paniculata Arn.)
112
sindok (Scorodocarpus borneensis (Baill.) Becc.)
281
sindu (Scorodocarpus borneensis (Baill.) Becc.)
281
sireh sireh (Anisophyllea corneri Ding Hou)
16, 19
sisit (Alangium havilandii Bloemb.)
9
soipang (Pittosporum ferrugineum Aiton)
300
t-izos-izos (Sarcotheca glauca (Gaertn.) Kanis)
292
tabarus (Sarcotheca spp.; Sarcotheca rubrinervis Hallier f.)
289, 291, 295
tabaus (Sarcotheca diversifolia (Miq.) Hallier f.)
291
tagal (Ceriops tagal (Pers.) C.B. Rob.)
336
taikakang (Cratoxylum cochinchinense (Lour.) Blume)
224
tait berak (Turpinia sphaerocarpa Hassk.)
461
tampaluan (Licania splendens (Korth.) Prance)
167
512
tangal (Ceriops tagal (Pers.) C.B. Rob.)

tanggal (Ochanostachys amentacea Mast.)
tangilan (Agathis lenticula de Laub.)
tansanglang (Engelhardia spp.)
tanug (Ceriops tagal (Pers.) C.B. Rob.)
tapang (Koompassia malaccensis Maingay ex Benth.)
tapong-tapong (Turpinia sphaerocarpa Hassk.)
tat (Cratoxylum arborescens (Vahl) Blume)
tebarus (Sarcotheca diversifolia (Miq.) Hallier f.)
tengar (Ceriops decandra (Griff.) Ding Hou; Ceriops tagal (Pers.) C.B. Rob.)
tengar samak (Ceriops tagal (Pers.) C.B. Rob.)
tengar tikus (Ceriops decandra (Griff.) Ding Hou)
tenghilan (Irvingia malayana Oliver ex A.W. Benn.)
tengoda (Anisophyllea ferruginea Ding Hou)
tenug (Ceriops tagal (Pers.) C.B. Rob.)
tepanga (Engelhardia serrata Blume)
terang bulu (Nyssa javanica (Blume) Wangerin)
tigarun (Crateva magna (Lour.) DC.)
tikolod (Carallia sp. 1)
timau (Cratoxylum glaucum Korth.)
tingo-tingo (Bruinsmia styracoides Boerl. & Koord.)
tombuid (Eurycoma longifolia Jack)
tongkat ali (Brucea javanica (L.) Merr.; Eurycoma longifolia Jack)
tongkat langit (Eurycoma longifolia Jack)
troduh (Scorodocarpus borneensis (Baill.) Becc.)
tubu (Agathis orbicula de Laub.)
tui (Dolichandrone spathacea (L. f.) K. Schum.)
tuih (Dolichandrone spathacea (L. f.) K. Schum.)
tuik-tuik hutan (Radermachera ramiflora Steenis)
tulang (Allantospermum borneense Forman)
tulang kwe (Trigoniastrum hypoleucum Miq.)
tulang payung piang (Sarcotheca glauca (Hook. f.) Hallier f.)
tulong (Agathis borneensis Warb.)
tumu (Agathis endertii Meijer Drees)
tumuh (Agathis orbicula de Laub.)
tungilan (Agathis lenticula de Laub.)
tungkat ali (Eurycoma longifolia Jack)
udok-udok (Colubrina beccariana Warb.)
wampee (Clausena lansium (Lous) Skeels)
wampi (Clausena lansium (Lous) Skeels)
yulu tambang (Carallia sp. 1)
513
336
278
32
233
336
187
461
223
291
335, 336
336
335
434
21
336
243
255
104
332
225
465
433
431, 433
433
281
32
37
37
44
429
469
292
29
30
32
32
433
312
369
369
332
TREE FLORA
of
SABAH AND SARAWAK
Tree Flora of Sabah and Sarawak Editorial Committee Chairman: E. Soepadmo (Forest Research
Institute Malaysia). Members: A. Latiff (Universiti Kebangsaan Malaysia), Ruth Kiew (Universiti
Pertanian Malaysia), H.S. Lee (Sarawak Forestry Department), L.G. Saw (Forest Research Institute
Malaysia), K.M. Wong (Sabah Forestry Department). Editorial assistant: R.C.K. Chung (Forest
Research Institute Malaysia). Advisors: P.S. Ashton (Harvard University, U.S.A.), J. Dransfield
(Royal Botanic Gardens, Kew, U.K.), C. Kalkman (Rijksherbarium, Leiden, the Netherlands), T.C.
TREE FLORA
of
SABAH AND SARAWAK
Volume Two
edited by
E. Soepadmo, K.M. Wong and L.G. Saw
Sabah Forestry
Forest Research Institute

Malaysia

L.G. Saw & B. Aslina (2007)
Sarawak Forestry
Copyright Forest Research Institute Malaysia (FRIM),

Sabah Forestry Department, and Sarawak Forestry Department 1996.
Forest Research Institute Malaysia (FRIM), Kepong, 52109 Kuala Lumpur, Malaysia
Perpustakaan Negara Malaysia Data Pengkatalogan-dalam-Penerbitan

Tree flora of Sabah and Sarawak / edited by E. Soepadmo,
K.M. Wong and L.G. Saw.
ISBN 983-9592-34-3 (v. 1)
ISBN 983-9592-56-4 (v. 2)
1. BotanySabah. 2. BotanySarawak. 3. BotanyClassification.
4. PlantsIdentification. I. Soepadmo, E. II. Wong, K.M. (Khoon Meng).
III. Saw, L.G.
581.095952
Electronic version by L. G. Saw & B. Aslina, Forest Research Institute Malaysia, 2007.
CONTENTS
Foreword
Acknowledgements
FAMILIES
1. Anacardiaceae (K.M Kochummen)
2. Boraginaceae (A.L. Dayang Awa)
3. Caprifoliaceae (Noorma Wati Haron)
4. Casuarinaceae (Runi S. Pungga)
5. Chloranthaceae (John B. Sugau)
6. Crypteroniaceae (J.T. Pereira)
7. Ctenolophonaceae (Runi S. Pungga)
8. Daphniphyllaceae (Balu Perumal)
9. Epacridaceae (S.P. Lim)
10. Erythroxylaceae (R.C.K. Chung)
11. Ixonanthaceae (Runi S. Pungga)
12. Leeaceae (A. Latiff)
13. Loganiaceae (K.M. Wong & John B. Sugau)
14. Lythraceae (P.C. Yii)
15. Malvaceae (Balu Perumal)
16. Myricaceae (A. Noorsiha)
17. Nyctaginaceae (A. Latiff)
18. Santalaceae (L.S.L. Chua)
19. Sapindaceae (F. Adema, P.W. Leenhouts & P.C. van Welzen)
20. Scyphostegiaceae (E.J.F. Campbell-Gasis)
21. Tetrameristaceae (John B. Sugau)
22. Ulmaceae (E. Soepadmo & Z. Edi Hamli)
23. Winteraceae (K.M. Wong)
Page
vii
ix
1
93
107
119
129
135
151
155
159
167
175
181
189
225
233
245
251
257
263
375
379
383
403
409
411
413
435
FOREWORD
t gives us great pleasure to provide this foreword. The production of the second volume, in just a
year after the first, is a noteworthy achievement. This volume, comprising 23 families, covers a
range of small and large families. We understand that, as in Volume 1, the revisions are not just
mere adaptations from previously revised treatments. This is attested by the discovery of 39 new
species during the course of the revisions of the families for this volume. Anacardiaceae (with 19
species) and Loganiaceae (also 19 species) had the most number of new species. Considering that
these families have been revised under the Flora Malesiana project, these new discoveries show that
continued exploration of the flora of Sabah and Sarawak will reward scientific research. The
importance and richness of the flora in Sabah and Sarawak makes it even more important to continue
documenting the flora in a systematic fashion.
As we have wished, this volume includes the work of botanists relatively new to taxonomic research.
Inasmuch as the project recognises the importance of documenting the tree flora, research and training
opportunities are also an integral part of the project. The Overseas Development Administration of the
United Kingdom has also continued to support both post-graduate and short-term training, a rather
fortunate aspect. Equally important but often not appreciated is the training resulting from the direct
interaction between the more senior botanists of the project and the newcomers in producing the
revisions.
For a project of such magnitude, funding has always been a major concern to those involved. We are
happy to have had the continued financial support of the Malaysian Government through the IRPA
(the Intensification of Research by Priority Areas) programme, the Overseas Development
Administration (ODA) of United Kingdom, and the International Tropical Timber Organization
(ITTO).
We wish to put on record our sincere thanks to these organizations.We also wish to congratulate and
thank all contributors of this volume. Finally, our congratulations also to the editors of this volume.
Dr. Abdul Razak Mohd. Ali

Director-General
Haji Awang Tengah Haji Awang Amin
Director
Malaysia
Mr. Cheong Ek Choon
Director
Malaysia
vii
ACKNOWLEDGEMENTS
nce more, we are able to perform the pleasant duty of recording our debt and
appreciation to many institutions and individuals in the successful completion of
Volume 2 of the Flora. The financial support of the Malaysian Government, the
Overseas Development Administration (ODA) of the United Kingdom and the International
Tropical Timber Organization (ITTO) has been fundamental to the smooth progress required for
this work. Dr. Abdul Razak Mohd Ali (Director-General, Forest Research Institute Malaysia),
Tuan Haji Awang Tengah bin Haji Amin (Director of the Sabah Forestry Department) and Mr.
Cheong Ek Choon (Director of the Sarawak Forestry Department) have ably directed resources
under their charge to ensure this continuation.
We would like to record special appreciation of the roles played by Dato Dr. B.C.Y. Freezailah
and Dr. Efransjah (ITTO), as well as Dr. Jeff Burley and Dr. Philip Bacon (Oxford Forestry
Institute) in the smooth management of the project. Our collaborating institutions in Malaysia
(the Universiti Kebangsaan Malaysia, Universiti Malaya, Universiti Malaysia Sarawak,
Universiti Pertanian Malaysia) and overseas (the Royal Botanic Gardens, Kew; the Oxford
Forestry Institute; the Rijksherbarium in Leiden; the Herbarium Bogoriense, Indonesia; the
Singapore Botanic Garden; the Arnold Arboretum of Harvard University) have continued to
give valuable assistance and support.
We have again benefitted from the use of facilities and loans of specimens for study approved
by the keepers and curators of the herbaria at the Arnold Arboretum (A), Bogor (BO), the
British Museum (BM), Cambridge (CGE), Kew (K), Kepong (KEP), Leiden (L), Sandakan
(SAN), Kuching (SAR), Singapore (SING), the Universiti Kebangsaan Malaysia at Bangi
(UKMB), the Universiti Malaya (KLU), the Universiti Pertanian Malaysia and the Universiti
Malaysia Sarawak. Prof. P.S. Ashton, Prof. C. Kalkman, Dr. J. Dransfield and Dr. T.C.
Whitmore kindly gave their time to advise on editorial matters. We are grateful to Prof.
Abdul Latiff Mohamad, Prof. Ruth Kiew and Mr. H.S. Lee for their help and cooperation as
members of the editorial committee. Many others have provided assistance during visits to
their institution by project botanists, including at Kew (Prof. G.T. Prance, its Director; Mr.
M.J.E. Coode, Dr. D. Kirkup, Mrs. Diane Bridson, Mr. N. Martland), Leiden (Prof. P. Baas,
its Director; Dr. M. Roos, Dr. J.F. Veldkamp, Dr. W.J.J.O. de Wilde, Dr. E.F. de Vogel; Dr.
Ding Hou, Dr. P. van Welzen, Dr. M.M.J. van Balgooy, Dr. P. Kessler, Dr. P.W. Leenhouts,
Dr. F. Adema, Mrs. Stans Kofman), Harvard (Prof. P.S. Stevens), Oxford (Mr. D. Filer, Prof.
D.J. Mabberley), and Singapore (Dr. W.K. Tan, Dr. L. Leong, Dr. S.C. Chin, Mr. E.P. Tay).
We are grateful also to the staff of the libraries and photographic units at the British Museum,
Kew, Leiden and Singapore for much assistance rendered.
We record our special appreciation for the help given by many on the home scene in Sarawak
(Mr. Lee Hua Seng, Abang Abdul Hamid Karim, Abang Mohtar Pawozan, Ms. Runi S.
Pungga, Mr. P.C. Yii, Haji Othman, Ms. Mohizah Mohamad, Mr. Julaihi Abdullah, Mr. S.
Teo), Sabah (Mr. Anuar Haji Mohammad, Mr. R.C. Ong, Dr. Y.F. Lee, Mr. A. Berhaman,
Ms. J.T. Pereira, Mr. J.B. Sugau, Mr. S.P. Lim, Mr. L. Madani, Mr. Dewol Sundaling) and
ix
Kepong (Dr. Wan Razali Wan Mohd., Mrs. Norhara Hussein, Mrs. Norli Raja Mohammad, Dr.
Abdul Rahim Haji Nik, Mrs. Wan Rahmah Wan Abdul Raof, Dr. N. Manokaran, Dr. Lillian
S.L. Chua, Mr. K.M. Kochummen, Mr. K.C. Khoo, Mrs. H.K. Kong, Mr. R.C.K. Chung, Ms.
Noorsiha Ayop, Mr. Edi Hamli, Mrs. Rusea Go, Mr. Mat Asri Ngah Sanah, Mr. Kamarudin
Saleh, Mrs. Lucy S. V. Rajoo, Ms. Shahani Saad).
Illustrators who have assisted in Volume 2 include Ms. Rosemary Wise who has kindly rallied
to our help, Ms. Yap Pak Hau, Mr. Mohd. Nizam Isa, Mr. Zainal Mustaffa, Mr. Joseph Pao,
Mr. J. Wessendorp and Mr. J.H. van Os. Their work ably accompanies the accounts presented
here by the many collaborators who have helped to make this volume possible.
E. Soepadmo
K.M. Wong
L.G. Saw
ANACARDIACEAE
K.M. Kochummen
Kepong, Malaysia
Hooker f., Fl. Br. Ind. 2 (1876) 7; King, J. As. Soc. Beng. 65, 3 (1896) 459; Merrill, EB (1921) 348; Ridley,
FMP 1 (1922) 517; Masamune, EPB (1942) 408; Browne, FTSB (1955) 45; Backer & Bakhuizen f., FJ 2 (1965)
146; Smythies, CST (1965) 1; Burgess, TBS (1966) 3; Ding Hou, Blumea 24 (1978) 1, FM 1, 8 (1978) 395;
Anderson, CLTS (1980) 134; Wong, DMT (1982) 5; Corner, WSTM 3rd ed. 1 (1988) 106; Kochummen, TFM 4
(1989) 9; Whitmore, Tantra & Sutisna, CLK 1 (1989) 12; Ng, Mal. For. Rec. 34, 1 (1991) 14; Kessler &
Sidiyasa, Trees of the Balikpapan-Samarinda Area (1994) 36.
Trees, erect or scandent shrubs, climbers or very rarely epiphytic shrubs, usually with acrid,
often turpentine smell from crushed living parts; bole often with buttresses or stilt-roots (in
swampy habitats), or without buttresses (Mangifera). Bark smooth, cracked, fissured, scaly
or dippled, sometimes with blotches of black exudate; inner bark reddish, pinkish or
yellowish, granular or laminated, with black, or white, creamy yellow, pink, brown or
colourless watery exudate that soon turns black or brown on drying. Sapwood whitish. Twigs
slender to stout, glabrous or hairy. Stipules absent. Leaves simple or imparipinnate, spirally
arranged, alternate or opposite (Bouea), rarely in pseudowhorls (Semecarpus in part), often
crowded at the ends of twigs; leaf or leaflet margins entire; petioles or petiolules often
swollen near the base, sometimes absent or very short. Inflorescences terminal, axillary, or
extra-axillary panicles; bracts and bracteoles caducous, rarely persistent; pedicels distinct,
obscure or absent, often articulate. Flowers bisexual or unisexual, radially symmetrical,
mostly 4- or 5-merous; floral axis between calyx and stamens (torus) often obscure,
sometimes distinct and elongated (Gluta, Swintonia); calyx 5- or 4-lobed, rarely 3-lobed,
sometimes calyptriform (hood-like) or splitting irregularly (Gluta spp.), caducous or
persistent and enlarged in fruit (Parishia spp.); petals 5 or 4, imbricate or valvate, rarely
contorted in bud, free, sometimes the basal part longitudinally adnate to the floral axis,
caducous or persistent and enlarged in fruit (Gluta in part, Swintonia); stamens equal or twice
the number of calyx lobes or petals, rarely more to numerous (Gluta spp.), inserted on the
margin of the disc, or just outside or inside of this margin, or on an enlarged torus (Gluta
spp.), filaments free or basally connate, glabrous, hairy, or papillose, anthers dorsifixed or
basifixed, sometimes dorso-basifixed, longitudinally dehiscent, 2-locular with 4 pollen sacs,
usually introrse, connective rarely prolonged, dilated and apically 2-lobed (Androtium); disc
usually present and distinct (obscure or absent in Gluta spp.), often fleshy, sometimes thin,
round, flat or concave above, pulvinate, rim-like, cup-like or with 5 gland-like lobes
(Swintonia), rarely cylindrical and stipe-like (some Mangifera spp.), often crenulate,
emarginate, rarely lobed; ovary free or basally adnate to the disc or receptacle, superior,
sometimes partly or wholly immersed in the disc or receptacle and appearing inferior (some
species of Melanochyla, Pegia, and Semecarpus), or inferior (Drimycarpus), usually sessile,
sometimes stipitate (Gluta), 1-carpellate and 1-locular, or syncarpous and 25(12)-carpellate
and 25(12)-locular, or apocarpous (46-carpellate in Androtium and Buchanania)
or carpels incompletely connate (5-carpellate in Dracontomelon and Koordersiodendron), usually

only one carpel fertile; styles 15(12), distinct or obscure, terminal or eccentric, stigmas 15(
12), distinct or obscure; pistillodes small, obscure or absent in male flower; ovule one in each
carpel, pendulous. Fruit a drupe, sometimes subtended by persistent enlarged calyx lobes
(Parishia), or enlarged petals (Gluta and Swintonia spp.), or enlarged fleshy pedicel and
receptacle (in Anacardium and Semecarpus), 15(12)-locular, 15(12)-seeded; epicarp thin,
mesocarp usually fleshy and resinous, sometimes waxy and oily, endocarp fibrous, crustaceous,
woody or bony. Seeds without or with little endosperm, rarely marked with sinuous intricate lines
(as in some Mangifera spp.); testa membranous or chartaceous, sometimes adnate to the
endocarp; embryo straight or curved, cotyledons free, rarely partly or incompletely united (Gluta
spp.), plano-convex, rarely unequal, radicle short.
Distribution. About 70 genera with c. 600 species, distributed mainly in the tropics and
subtropics with its highest diversity in the Malesian region. Eighteen genera and 95 species in
Sabah and Sarawak.
Ecology. Occurring in a variety of habitats, including peat swamp, limestone, riverine, kerangas,
hill and montane forest, to 2400 m. Most species are found scattered in natural forest. However,
certain species, e.g., Campnosperma coriaceum, are found growing gregariously. The deciduous
habit occurs in a number of genera; flowers and new leaves appear together in such species. The
flowers possess nectar-bearing organs or discs and are sometimes fragrant. They are evidently
pollinated by insects. The fruits are dispersed by animals, water or wind.
Uses. Several species belonging to a number of genera are widely cultivated for their edible fruits
and nuts. They include the introduced species Anacardium occidentale (cashew nut), Spondias
cytherea (hog-plum), the indigenous Bouea spp. (kundang) and Mangifera spp. (mango).
Important commercial timber trees include species of Campnosperma, Dracontomelon,
Koordersiodendron and Swintonia, the details of which are given under each genus. The
heartwood of some species is hard and durable and makes excellent furniture because of the
beautiful grain. Some species (although none from Sabah and Sarawak) produce valuable
economic products such as lacquers (from the resinous sap of Gluta laccifera, G. usitata and
Rhus verniciflua) and tannins (from species of Schinopsis and Rhus). A number of species are
used in traditional medicine.
Dermatitis. The sap is irritant in many species and can cause serious allergic contact dermatitis
which manifests as redness, swelling, vesiculation and weeping of the affected parts. In most
cases, the severe inflammation is controlled by systemic steroids given over a period of a week to
10 days. The intensity of reaction varies with the species and the sensitivity of the individual. The
worst irritant exudates are from species of Drimycarpus, Gluta, Melanochyla and Semecarpus. In
temperate countries poisonous Anacardiaceae are represented by species of Rhus, the so-called
poison ivies or poison oaks.
Taxonomy. Pinnate-leaved members of the Anacardiaceae, i.e. species of Dracontomelon,
Koordersiodendron, Parishia and Pentaspadon, can be confused with members of the
Burseraceae, Meliaceae and Sapindaceae. However, Anacardiaceae species have no stipules
ANACARDIACEAE (KOCHUMMEN)
whereas stipules are present in some Burseraceae (Canarium, Garuga). Technically, Anacardiaceae
have a single ovule in each locule of the ovary, whereas Burseraceae have two ovules per locule.
From the Meliaceae and Sapindaceae, Anacardiaceae may be distinguished by the resinous nature.
Many species of Aglaia in the Meliaceae also produce white sap, but in these species stellate hairs are
always present on the shoots, a character absent from the Anacardiaceae.
Key to genera
1. Leaves simple......................................................................................................................2
Leaves trifoliolate or pinnately compound........................................................................12
2. Leaves opposite......................................................................................................2. Bouea
Leaves alternate or spirally arranged...................................................................................3
3. Carpels free........................................................................................................................4
Carpels united......................................................................................................................5
4. Anther locules not separate, connective not prolonged.................................3. Buchanania
Anther locules separate, connective prolonged and apically bilobed..............1. Androtium
5. Ovary inferior. Leaf margin thickened........................................................6. Drimycarpus
Ovary superior. Leaf margin not thickened.........................................................................6
6. Ovary with 3 styles.....................................................................................15. Semecarpus
Ovary with 1 style...............................................................................................................7
7. Stigmas 3.............................................................................................................................8
Stigma 1...............................................................................................................................9
8. Trees. Leaves papillose beneath. Petals velvety hairy inside. Fruits hairy .........................
...................................................................................................................10. Melanochyla
Shrubs. Leaves not papillose beneath. Petals glabrous inside. Fruits glabrous ..................
..................................................................................................................14. Rhus (in part)
9. Calyx calyptriform (hood-like) or bursting irregularly at anthesis, without distinct lobes.
Disc absent. Stamens inserted on the torus (receptacle).........................................7. Gluta
Calyx not so, with distinct lobes. Disc present. Stamens not inserted on the torus.....10
10. Ovary 2-locular. Stamens 610, all fertile. Leaves with peltate scales..................................
................................................................................................................4. Campnosperma
Ovary 1-locular. Stamens 5 (very rarely 10), one to all of them fertile. Leaves without
peltate scales......................................................................................................................11
11. Petals with glandular ridges inside, caducous. Style often lateral....................9. Mangifera
Petals without glandular ridges inside, persistent in fruit. Style terminal.......17. Swintonia
12. Trees or shrubs.............................................................................................................13

Woody climbers...............................................................................................................19
13. Inner bark with abundant white sap. Flowers 4-merous. Fruit with long wings....................
...........................................................................................................................11. Parishia
Inner bark with scant white latex. Flowers 5-merous. Fruit without wings.......................14
14. Stamens or staminodes of the same number as petals...............................14. Rhus (in part)
Stamens or staminodes twice the number of petals...........................................................15
15. Petals valvate..............................................................................16. Solenocarpus (in part)
Petals imbricate at least at the upper parts.........................................................................16
16. Ovary 1-locular..........................................................................................12. Pentaspadon
Ovary 512-locular............................................................................................................17
17. Leaflets 1016 pairs, without domatia in the axils of lateral veins........................................
.........................................................................................................8. Koordersiodendron
Leaflets usually 39 pairs; domatia often present in the axils of lateral veins...................18
18. Leaflet base cuneate, often asymmetric. Ovary 512-locular; styles 512. Rare trees..........
....................................................................................................................13. Pleiogynium
Leaflet base broadly rounded, symmetric or asymmetric. Ovary 5-locular; styles 5. Very
common trees..........................................................................................5. Dracontomelon
19. Stamens of the same number as petals......................................................14. Rhus (in part)
Stamens or staminodes twice the number of petals...........................................................20
20. Petals valvate. Leaflets without domatia in the axils of lateral veins.....................................
....................................................................................................16. Solenocarpus (in part)
Petals imbricate. Leaflets with domatia in the axils of lateral veins......................................
Pegia Colebr.
Trans. Linn. Soc. 1, 15 (1827) 364; Ding Hou, FM 1, 8 (1978) 488.
Three species; India, Myanmar, Thailand, Laos, Vietnam, China, Borneo.
Woody climbers. Leaves alternate, imparipinnate; leaflets 34 pairs often with hairy
domatia in the axils of lateral veins. Flowers cream; ovary semi-inferior.
In Borneo, one species (P. sarmentosa (Lecomte) Hand.-Mazz.), recorded from Sabah
and Kalimantan only. Lowland to 1500 m.
1. ANDROTIUM Stapf
(Greek, andros = male, otion = ear-lobe; the stamens furnished with ear-like lobes)
rengas padang (Malay)
In Hooker, Icon. Pl. (1903) t. 2763; Merrill, EB (1921) 349; Masamune, EPB (1942) 408; Smythies, CST (1965)
1; Ding Hou, FM 1, 8 (1978) 420; Anderson, CLTS (1980) 134; Kochummen, TFM 4 (1989) 13.
Trees. Leaves spirally arranged, simple. Inflorescences axillary panicles, hairy. Flowers
bisexual; calyx 5(or 4)-lobed; petals 5 (or 4), imbricate, margin sparsely hairy; stamens twice
the number of petals, anthers basifixed, with 2 separate anther locules, connective prolonged,
dilated and apically two-lobed; disc intrastaminal, cup-shaped, crenulate along the margin;
ovary superior, subglobose, pilose; carpels 5, free, only one fertile, ovule one per carpel,
style obscure, stigma oblique. Fruit lentiform.
Distribution. Monotypic; Peninsular Malaysia, Borneo.
Ecology. Lowland forests including swamps.
Androtium astylum Stapf
Fig. 1.
(Latin, astylus = without style; the ovary)

In Hooker l.c. t. 2763; Merrill l.c. 349; Masamune l.c. 408; Ding Hou, FM 1, 8 (1978) 420; Anderson
l.c. 134; Kochummen l.c. 13. Type: Haviland 2860, Borneo, Sarawak, Kuching (holotype K; isotypes
BO, L, SING).
Small tree to 20 m tall, 40 cm diameter. Bark chocolate-brown with grey patches, finely
cracking; inner bark dark red, laminated. Sapwood pinkish. Terminal bud ovoid-ellipsoid,
hairy. Leaves coriaceous; elliptic, oblong or obovate, 3.514 x 25.5 cm; base cuneate,
margin recurved, apex acute, rarely obtuse or emarginate; midrib raised above, lateral veins
513 pairs, visible on both surfaces, intercostal veins reticulate, equally prominent as lateral
veins; petioles 0.51 cm long. Flowers white, in 18.5 cm long-hairy panicles. Fruits
resembling those of Buchanania, c. 12 x 15 mm, with slightly depressed apex.
Distribution. Peninsular Malaysia, Borneo. Uncommon in Sabah, recorded from Lamag,
Pinangah and Sandakan areas; common in Sarawak.
2. BOUEA Meisn.
(A. Boue, 17941881, German naturalist)
kundang, kundang rumenia (Malay)
Pl. Vas. Gen. (1837) Tab. Diagn. 75 & Comment. 55; King, J. As. Soc. Beng. 65, 3 (1896) 465;
Merrill, EB (1921) 350; Ridley, FMP 1 (1922) 519; Masamune, EPB (1942) 408; Backer &
Backhuizen f., FJ 2 (1965) 150; Ding Hou, Blumea 24 (1978) 4, FM 1, 8 (1978) 466; Anderson,
CLTS (1980) 135; Wong, DMT (1982) 112; Corner, WSTM 3rd ed. 1 (1988) 109; Kochummen, TFM
4 (1989) 13; Whitmore, Tantra & Sutisna, CLK 1 (1989) 12; Ng, Mal. For. Rec. 34, 1 (1991) 14;
Kessler & Sidiyasa, Trees of the Balikpapan-Samarinda Area (1994) 37.
Small to medium-sized trees. Terminal and axillary buds prominent. Leaves oppositedecussate, simple. Inflorescences usually axillary panicles. Flowers male and bisexual;
calyx 35-lobed; petals 35, imbricate, glabrous, keeled lengthwise; stamens 35, filaments
5
Fig. 1. Androtium astylum. A, flowering leafy twig; B, detail of lower leaf surface; C, flower in
longitudinal section. (All from S. 57105.)
subulate, glabrous, anthers basifixed; disc round, flat or slightly concave, glabrous; ovary
superior, ovoid or subglobose, 1-locular and 1-ovulate, puberulous or glabrous; style short,
stigma rounded and flat; pistillodes minute in male flowers. Fruits 1-locular; mesocarp
pulpy, edible. Seeds with testa adnate to the endocarp; embryo straight, cotyledons free,
plano-convex; germination semi-hypogeal.
Distribution. Three species; tropical SE Asia and Malesia; one species in Sabah and
Sarawak.
Uses. The trees have dense bushy crowns and can be recommended for planting as
ornamental shade trees. The ripe fruits of B. oppositifolia and the cultivated B. macrophylla
are eaten raw or steamed. The young fruits are sometimes pickled.
Timber. The common Malaysian name for the timber is kundang. The wood is moderately
hard to hard, the sapwood is not differentiated from the heartwood which is light brown with
a red tinge or grey-brown, and sometimes a core of dark coloured wood is formed in some
trees; the grain is interlocking and texture slightly coarse and even.
Notes. In the sterile stage, Bouea can be confused with some other genera with opposite
leaves such as Garcinia, Mesua (Clusiaceae) and Eugenia (Myrtaceae) but the resinous smell
from the crushed fresh leaves distinguishes it.
Bouea oppositifolia (Roxb.) Meisn.
Fig. 2.
(Latin, oppositus = opposite, folius = leaf; the opposite leaves)

l.c. 55; King l.c. 465; Ridley l.c. 520; Ding Hou, FM 1, 8 (1978) 466; Anderson l.c. 135; Corner l.c. 110;
Kochummen l.c. 14; Whitmore, Tantra & Sutisna l.c. 12; Kessler & Sidiyasa l.c. 37. Basionym: Mangifera
oppositifolia Roxb., Fl. Ind. ed. Wall. 2 (1824) 434. Type: Wallich 8490, Burma, Pegu (holotype K).
Synonyms: B. angustifolia Blume, Mus. Bot. Lugd. Bat. 1 (1850) 204, Merrill l.c. 350, Masamune l.c. 408; B.
microphylla Griff., J. As. Soc. Beng. 23 (1854) 15, Merrill l.c. 350 (as B. macrophylla), Masamune l.c. 408 (as
B. macrophylla).
Medium-sized tree to 20 m tall, 30 cm diameter; bole without buttresses. Bark dark grey, smooth to
cracking to shallowly fissured; inner bark reddish with dark red latex. Sapwood whitish. Terminal
bud ellipsoid, acute. Twigs swollen at the nodes. Leaves thick-coriaceous; elliptic to oblong,(1.5)
3.510 x (0.8)1.34.4 cm; base broadly cuneate, apex obtuse to rounded; midrib raised above,
lateral veins 817 pairs, very faint on both surfaces, sometimes impressed beneath, intercostal veins
invisible; petioles 0.81.7 cm long, narrowly grooved above. Inflorescences axillary panicles, 26 cm
long. Flowers white to pale yellow. Fruits yellow, red or orange when ripe, broadly ellipsoid, 1.51.8
x 0.81 cm when dried, stipe c. 3 mm long.
Distribution. Myanmar, Andaman Islands, Thailand, Laos, Cambodia, Vietnam, S China Peninsular
Malaysia, Sumatra, Borneo. In Sabah, recorded from Papar and Tawau districts, while in Sarawak,
recorded from Bako National Park and Semengoh Arboretum; also in Brunei and Kalimantan.
Fig. 2. Bouea oppositifolia. A, fruiting leafy twig; B, flowering leafy twig; C, flower in longitudinal
section. (A from SAN 84754, BC from SAN 78049.)
Ecology. Lowland forests including sandy coastal and peat swamp forests. Flowering in
February, March and July and fruiting in February and September.
Uses. Fruits edible.
3. BUCHANANIA Spreng.
(F. Buchanan-Hamilton, 17621829, Scottish botanist)
kasat (Kenyah), kepala tundang, otak udang (Malay)
In Schrader, J. Bot. 2 (1801) 234; Merrill, EB (1921) 348; Ridley, FMP 1 (1922) 518; Masamune,
EPB (1942) 408; Browne, FTSB (1955) 45; Backer & Bakhuizen f., FJ 2 (1965) 147; Burgess, TBS
(1966) 4; Ding Hou, Blumea 24 (1978) 4, FM 1, 8 (1978) 412; Anderson, CLTS (1980) 135; Wong,
DMT (1982) 5; Corner, WSTM 3rd ed. 1 (1988) 110; Kochummen, TFM 4 (1989) 15; Whitmore,
Tantra & Sutisna, CLK 1 (1989) 13; Ng, Mal. For. Rec. 34, 1 (1991) 17; Kessler & Sidiyasa, Trees of
Balikpapan-Samarinda Area (1994) 38.
Small to medium-sized trees with dense bushy crowns. Bark grey-brown to pale whitish,
smooth; inner bark reddish. Sapwood pale. Leaves spirally arranged, simple. Inflorescences
usually axillary panicles. Flowers bisexual; calyx usually 5-lobed (rarely 4- or 6-lobed),
persistent or caducous; petals usually 5 (rarely 4 or 6), imbricate, glabrous; stamens twice the
number of petals, anthers basifixed, sagittate (except in B. splendens); disc cup-shaped with
crenulate upper margin; ovary superior, glabrous or hairy, carpels 46, free, each 1-ovulate,
usually only one fertile. Fruits 1-locular; endocarp thick, woody or bony. Seeds one per fruit;
testa free from endocarp; cotyledons free, plano-convex; germination epigeal, cotyledons
fleshy, hypocotyle elongate, first two leaves opposite.
Distribution. About 25 species; Asia, Malesia, Australia, Micronesia, Melanesia, and Samoa;
4 species in Sabah and Sarawak.
Ecology. Mainly in lowland forests including coastal, peat swamp and limestone.
Key to Buchanania species
1. Leaves sessile or ..............................................................................3. B. sessifolia (in part)
Leaves distinctly stalked.....................................................................................................2
2. Pedicels not articulate. Leaf apex usually obtuse or rounded...................1. B. arborescens
Pedicels articulate. Leaf apex acute, acuminate or cuspidate..............................................3
3. Leaves smaller, 7.513 x 2.55.5 cm, apex acuminate; lateral veins 811 pairs; intercostal
veins distinctly reticulate. Petioles 0.51.5 cm long. Anthers not sagittate (locules not
separate at base)........................................................................................4. B. splendens
Leaves larger, (9)2035(45) x (3)58(10.5) cm, apex acute or cuspidate; lateral veins
9
12 pairs or more; intercostal veins scalariform-reticulate. Petioles absent or 1.55.5 cm

long. Anthers sagittate (locules separate at base).................................................................4
4. Bud scales inconspicuous. Leaves with short but distinct intermediate veins between lateral
veins; lateral veins faint on upper leaf surface; leaf apex cuspidate. Petioles shorter, 1.53 cm
long, with swollen base. Stamen filament papillose. Fruits obliquely subcordate, with
persistent calyx...................................................................................3. B. sessifolia (in part)
Bud scales linear to lanceolate, conspicuous. Leaves without such intermediate veins; lateral
veins raised on both surfaces; leaf apex acute. Petioles longer, 2.55.5 cm long, without
swollen base. Stamen filaments not papillose. Fruits sublentiform (like a doubly convex lens),
without persistent calyx...........................................................................................2. B. insignis
1. Buchanania arborescens (Blume) Blume
Fig. 3A.
(Latin, arbor = tree; tree-like)

Mus. Bot. Lugd. Bat. 1 (1850) 183; Merrill l.c. 348; Masamune l.c. 408; Backer & Bakhuizen f. l.c. 147; Ding
Hou, FM 1, 8 (1978) 415; Burgess l.c. 4; Anderson l.c. 134; Wong l.c. 29; Corner l.c. 111; Kochummen l.c. 16;
Whitmore, Tantra & Sutisna l.c. 13; Ng l.c. 17. Basionym: Caniogeton arborescens Blume, Bijdr. Fl. Ned. Ind.
(1826) 1156. Type: Blume, s.n., Java (holotype L). Synonyms: B. lucida Blume l.c. (1850) 184, Merrill l.c. 348,
Masamune l.c. 409; B. florida Schau. var. lucida (Blume) Engler in A. DC., Mon. Phan. 4 (1883) 188; B.
glaberrima Ridl., Kew Bull. (1933) 195, Masamune l.c. 409.
Small tree, rarely to 28 m tall, 30 cm diameter. Bark grey-brown, smooth; inner bark red or pinkish
brown, laminated. Sapwood yellowish. Twigs dark brown to blackish, with prominent scars of
previous flowering. Bud scales inconspicuous, lanceolate, glabrous or hairy. Leaves generally in
clusters, subcoriaceous to coriaceous; elliptic to oblanceolate or obovate, 424 x 29.5 cm; base
cuneate, apex obtuse, rounded, or rarely acute; midrib flattened above, lateral veins 712 pairs,
faintly visible on both surfaces, intercostal veins reticulate, equally prominent as lateral veins;
petioles 14 cm long, often flattened. Flowers white, glabrous; pedicels not articulate, 24 mm long;
calyx caducous; stamen filaments glabrous, not papillose, narrowed and whitish towards apex,
anthers sagittate. Fruits sublentiform, red tinged green, c. 10 mm diameter.
Distribution. Myanmar, Andamans, Thailand, Indo-China, China, Formosa, Malesia, Solomon
Islands. Common in Sabah and Sarawak; also in Brunei and Kalimantan.
Ecology. Lowland forest to 500 m, including beach and limestone forest, often gregarious by
riverbanks. Flowering and fruiting throughout the year. When in flower the tree crown is conspicuous
with creamy white colour.
Note. A very variable species. There are plenty of collections from Sabah with narrowly acute leaves
and glabrous inflorescences.
2. Buchanania insignis Blume

(Latin, insignis = outstanding, remarkable)
l.c. (1850) 184; Merrill l.c. 348; Masamune l.c. 409; Burgess l.c. 5; Ding Hou, FM 1, 8 (1978) 415; Anderson
l.c. 135; Whitmore, Tantra & Sutisna l.c. 13; Kessler & Sidiyasa l.c. 38. Type: Korthals, s.n. (= L. sheet no.
897.363179), Borneo (holotype L; isotype BO).
10
Fig. 3. Buchanania arborescens (A); B. sessifolia (BC). A, fruiting leafy twig; B, flowering leafy
twig; C, flower in longitudinal section. (A from SAN 93275, BC from S. 23375.)
11
Small to medium-sized tree, to 25 m tall, 40 cm diameter. Bark grey-white, smooth; inner bark
red, laminated. Twigs stout, 812 mm thick, angular or irregularly grooved and densely brown
short-hairy when young. Bud scales conspicuous, linear to lanceolate, 1015 mm long, densely
brown-hairy. Leaves slightly hairy on midrib and veins below when young; oblanceolate or
narrowly obovate, 2445 x 5.510.5 cm; base attenuate, apex acute; midrib raised or flattened
above, lateral veins 2030 pairs, raised on both surfaces, shorter intermediate lateral veins
absent, intercostal veins scalariform-reticulate, visible on both surfaces; petioles 2.55.5 cm
long, without swollen base. Inflorescences 2036 cm long panicles, with many side branches,
yellowish hairy. Flowers white; pedicels distinctly articulate, up to 2 mm long; calyx lobes
broadly ovate with obtuse apices; petals oblong or elliptic; stamen filaments glabrous, not
papillose, upper part narrowed and whitish, anthers sagittate. Fruits sublentiform, without
persistent calyx, 710 mm diameter, green with red tinge when ripe.
Distribution. Borneo, Philippines. In Sabah, known from the Lahad Datu, Mostyn and Sandakan
districts. Not reported from Sarawak; also in Brunei and Kalimantan.
Ecology. Lowland forest to 150 m, including tidal swamps. Flowering in MarchMay. Young
fruits reported in May.
3. Buchanania sessifolia Blume
Fig. 3BC.
(Latin, sessilis = without stalk, folium = leaf; the sessile leaves)

l.c. (1850) 184; Merrill l.c. (1921) 349, PEB (1929) 166; Ridley l.c. 519; Masamune l.c. 409; Backer & Bakhuizen f. l.c. 48;
Burgess l.c. 5; Ding Hou, Blumea 24 (1978) 5, FM 1, 8 (1978) 418; Anderson l.c. 135 (as B. sessilifolia); Wong l.c. 29;
Kochummen l.c. 16; Whitmore, Tantra & Sutisna l.c. 13; Ng l.c. 17; Kessler & Sidiyasa l.c. 39. Lectotype (Ding Hou,
1978): Korthals 1034, Sumatra (L; isolectotype BO).
Small to medium-sized tree, to 20 m tall, 30 cm diameter. Bark grey-brown, smooth; inner bark
reddish. Sapwood pale. Twigs whitish, irregularly grooved, slender, less than 1 cm thick, reddish
brown hairy when young. Bud scales inconspicuous. Leaves glabrous or sparsely reddish brown
hairy below; obovate or narrowly oblanceolate, rarely elliptic, very variable in size, 935 x 310
cm; base attenuate, apex cuspidate, acumen c. 1 cm long; midrib raised above, lateral veins 1222
pairs, distinct below, faint above, with short intermediate veins, intercostal veins scalariformreticulate, visible below, faint above; petioles absent or 1.53 cm long, with swollen base.
Flowers whitish, hairy; pedicels articulate; calyx persistent; stamen filaments papillose,
gradually attenuate towards apex, apical part not whitish, anthers sagittate. Fruits obliquely
subcordate, 1013 x 811 mm, with persistent calyx.
Distribution. Laos, Thailand, Sumatra, Peninsular Malaysia, Borneo. Widely distributed in
Sabah and Sarawak; also in Brunei and Kalimantan.
Ecology. Lowland forest to 100 m. Flowering from February to November and fruiting from
April to December.
Notes. Very variable in petiole length and leaf pubescence.
12
4. Buchanania splendens Miq.

(Latin, splendens = shining; the leaves)
Fl. Ned. Ind. Suppl. (1861) 524; Ding Hou, FM 1, 8 (1978) 413; Anderson l.c. 135; Whitmore, Tantra & Sutisna l.c. 13.
Type: Teijsmann 3715, E Sumatra (holotype L; isotype BO). Synonym: B. fragrans Ridl., Kew Bull. (1933) 195,
Masamune l.c. 409.
Medium-sized tree to 20 m tall, 25 cm diameter. Bark grey-brown, smooth; inner bark

reddish. Sapwood whitish. Bud scales inconspicuous. Leaves thick-coriaceous; elliptic or
oblong, 7.513 x 2.55.5 cm; base cuneate, apex acuminate; midrib raised above, lateral
veins 811 pairs, raised on both surfaces, intercostal veins distinctly reticulate, visible on
both surfaces; petioles 0.51.5 cm long. Flowers white; pedicels articulate; calyx caducous,
lobes triangular; anthers not sagittate. Fruits globose, 35 mm across.
Distribution. Andaman and Nicobar Is., Sumatra, Borneo. Uncommon and of restricted
distribution in Sabah and Sarawak, known respectively from the Lahad Datu and Kuching
areas only; also in Brunei and Kalimantan.
Ecology. Lowland forests. Fruiting in February to May.
4. CAMPNOSPERMA Thwaites
(Greek, kamptein = bent, spermum = seed; the curved seeds)
ketang (Kayan), nyaletang (Kenyah), telatang (Berawan Murut),
terentang (Malay), tetang (Bidayuh)
In Hooker, J. Bot. Kew Misc. 6 (1854) 65; King, J. As. Soc. Beng. 65, 3 (1896) 494; Merrill, EB (1921) 351; Ridley, FMP 1
(1922) 533; Masamune, EPB (1942) 409; Browne, FTSB (1955) 46; Smythies, CST (1965) 1; Burgess, TBS (1966) 6; Ding
Hou, Blumea 24 (1978) 5, FM 1, 8 (1978) 524; Anderson, CLTS (1980) 135; Wong, DMT 30 (1982) 31; Corner, WSTM 3rd
ed. 1 (1988) 111; Kochummen, TFM 4 (1989) 18; Whitmore, Tantra & Sutisna, CLK 1 (1989) 13.
Trees with distinct Terminalia-branching; bole either with small buttresses or with stilt-roots
and pneumatophores (in swampy habitats). Leaves spirally arranged, simple, long- or shortpetioled, usually with minute peltate scales on both surfaces. Inflorescences axillary
panicles. Flowers unisexual (very rarely bisexual), calyx 4-lobed, rarely with 3 or 5 lobes;
petals 4, rarely 3 or 5, imbricate; stamens twice the number of petals, epipetalous ones
shorter than those alternate with them, all fertile, anthers dorso-basifixed, sterile in female
flowers; disc round and flat in male, cup-like in female flowers; ovary superior, subglobose,
2-carpellate, style 1, short or obscure, stigma 1. Fruits incompletely 2-locular with a
vertical, solid or hollow septum. Seeds 1 per fruit; embryo curved, cotyledons free, slightly
plano-convex or flat.
Distribution. Ten species; South and Central America, Madagascar, Seychelles, SE Asia,
Malesia, Micronesia and Melanesia. Three species in Sabah and Sarawak.
13
Ecology. Lowland, including swamp, to lower montane forest to 1100 m. The fruits are eaten by
birds. C. coriaceum grows gregariously.
Uses. The timber is light hardwood, grain shallowly to deeply interlocking, texture fine and even.
Sapwood usually not differentiated from heartwood, not durable on contact with ground;
susceptible to sapstain and fungal infection. Used for making match boxes and packing cases.
Key to Campnosperma species

1. Leaf base without ear-like lobes; petioles 25 cm long; midrib sunken
above........................................................................................................2. C. coriaceum
Leaf base with distinct ear-like lobes; petioles to 5 mm long; midrib flattened above........2
2. Leaf apex emarginate...............................................................................1. C. auriculatum
Leaf apex acute or obtuse.........................................................................3. C. squamatum
1. Campnosperma auriculatum (Blume) Hook. f.
Fig. 4AB.
(Latin, auriculatus = with ear-like appendages; the leaf base)

Fl. Br. Ind. 1 (1876) 41; King l.c. 495; Merrill l.c. (1921) 351; Ridley l.c. 534 (including var. wallichii (King) Ridl.);
Masamune l.c. 409; Ding Hou, Blumea 24 (1978) 5, FM 1, 8 (1978) 529; Smythies l.c. 2; Burgess l.c. 7; Anderson l.c. 235;
Wong l.c. 31; Corner l.c. 112; Kochummen l.c. 18; Whitmore, Tantra & Sutisna l.c. 13. Basionym: Buchanania auriculata
Blume, Mus. Bot. Lugd. Bat. 1 (1850) 185. Lectotype (Ding Hou, 1978): Korthals 779, Sumatra (L; isolectotype BO).
Tree to 36 m tall, 100 cm diameter; crown flat-topped; bole with spreading buttresses to 3 m tall.
Bark greyish, shallowly fissured; inner bark pinkish, with creamy exudate. Sapwood whitish.
Twigs stout, 11.5 cm thick, greyish. Leaves in close spirals, thick-coriaceous; obovate to
oblanceolate, 1636 x 4.514 cm; base narrowly attenuate with a pair of ear-like lobes (auricles)
at the base, apex emarginate; midrib flattened above, lateral veins 1820 pairs, raised on both
surfaces, intercostal veins scalariform-reticulate, faintly visible on both surfaces; petioles obscure
or to 5 mm long. Flowers lemon-yellow, in c. 50 cm long panicles. Fruits subglobose, 68 x 56
mm, green when young, becoming dull reddish when ripe.
Vernacular names. Sabah and Sarawakterentang (Malay).
Distribution. Thailand, Sumatra, including Banka island, Peninsular Malaysia, Borneo. Common
and widely distributed in Sabah and Sarawak; also in Brunei and Kalimantan.
Ecology. Lowland and hill forests to 1000 m, common near valleys.
2. Campnosperma coriaceum (Jack) Hall. f. ex Steenis

(Latin, coriaceus = leathery; the leaf)
Fl. Mal. Bull. 3 (1948) 74; Smythies l.c. 3; Burgess l.c. 7; Ding Hou, Blumea 24 (1978) 5, FM 1, 8 (1978) 530; Anderson l.c.
135; Wong l.c. 31; Corner l.c. 113; Whitmore, Tantra & Sutisna l.c. 14; Kochummen l.c. 20. Basionym: Coelopyrum
coriaceum Jack, Mal. Misc. 2, 7 (1822) 65. Type: Jack, s.n., Sumatra
14
Fig. 4. Campnosperma auriculatum (AB); C. squamatum (CD). A, fruiting leafy twig; B, detail of
leaf lower surface; C, flowering leafy twig; D, flower in longitudinal section. (AB from S. 37161, C
D from S. 40511.)
15
(K). Synonyms: Buchanania macrophylla Blume, Mus. Bot. Lugd. Bat. 1 (1850) 185; C.
macrophylla (Blume) Hook. f. l.c. 41, Merrill l.c. (1921) 351, Masamune l.c. 409.
Tree to 36 m tall, 60 cm diameter; bole with short buttresses, stilt-roots and looped
pneumatophores. Bark brown, cracking to fissured and scaly; inner bark reddish. Sapwood
white. Twigs c. 8 mm thick, densely brown short-hairy. Leaves densely brown short-hairy below,
thick-coriaceous; elliptic, oblong or obovate, 1336 x 4.516 cm; base attenuate down to petiole,
without auricles, apex rounded or emarginate; midrib sunken above, sharply keeled and
channelled below, lateral veins 1432 pairs, raised below, distinct above, intercostal veins
scalariform-reticulate, faintly visible on both surfaces; petioles 25 cm long, densely yellowish
short-hairy. Flowers greenish yellow, in hairy panicles. Fruits ovoid, 1218 x 816 mm, shortly
pointed, black when ripe.
Vernacular name. Sarawakterentang paya (Malay).
Distribution. Sumatra, Peninsular Malaysia, Borneo. Common and widely distributed in Sabah
and Sarawak; also in Brunei and Kalimantan.
Ecology. Peat swamp and fresh-water swamp forests, often gregarious.
3. Campnosperma squamatum Ridl.
Fig. 4CD.
(Latin, squamatus = furnished with scales; the leaf surface)

Kew Bull. (1933) 197; Masamune l.c. 409; Smythies l.c. 3; Burgess l.c 7; Ding Hou, FM 1, 8 (1978) 532; Anderson l.c. 135;
Wong l.c. 31; Corner l.c. 113; Whitmore, Tantra & Sutisna l.c. 14; Kochummen l.c. 20. Type: Haviland & Hose 3196,
Borneo, Sarawak, Kuching (holotype K; isotype L).
Small to medium-sized tree, to 25 m tall, 55 cm diameter; bole with short buttresses. Bark
yellowish grey, smooth to scaly; inner bark pink with creamy exudate. Sapwood whitish. Twigs
grey-white. Leaves oblanceolate or elliptic, 740 x 213 cm; base attenuate and slightly
widening towards the end to form ear-like lobes (sapling leaves very much longer and narrower),
apex acute or obtuse; midrib flattened above, lateral veins 1525 pairs, hardly distinct from the
equally prominent reticulate intercostal veins; petioles to 5 mm long. Flowers greenish yellow, in
hairy panicles. Fruits subglobose, 1217 mm diameter, green speckled white when fresh.
Vernacular name. Sarawakterentang puteh (Malay).
Distribution. Peninsular Malaysia, Borneo. Widely distributed in Sarawak but very uncommon
in Sabah, known from two collections, SAN 51734 from Lamag and SAN 60174 from the Telupid
area; also in Brunei and Kalimantan.
Ecology. Lowlands including fresh-water swamp, peat swamp and kerangas forests to hill forests
to 600 m.
Notes. One collection from Sarawak, S. 24104, has very large leaves, measuring to 30 cm long
and 13.5 cm broad.
16
5. DRACONTOMELON Blume
(Greek, drakon = dragon or snake, melon = fruit)
mengkuang, sengkuang (Iban)
Mus. Bot. Lugd. Bat. 1 (1850) 231; Merrill, EB (1921) 351; Ridley, FMP 1 (1922) 543; Masamune, EPB (1942) 409;
Browne, FTSB (1955) 47; Backer & Bakhuizen f., FJ 2 (1965) 151; Smythies, CST (1965) 5; Burgess, TBS (1966) 9; Ding
Hou, Blumea 24 (1978) 6, FM 1, 8 (1978) 468; Anderson, CLTS (1980) 135; Wong, DMT (1982) 51; Corner, WSTM 3rd
ed. 1 (1988) 113; Kochummen, TFM 4 (1989) 20; Whitmore, Tantra & Sutisna, CLK 1 (1989) 14; Ng, Mal. For. Rec. 34, 1
(1991) 17; Kessler & Sidiyasa, Trees of the Balikpapan-Samarinda Area (1994) 39.
Tall trees with spreading buttresses. Inner bark with scanty white latex. Leaves spirally arranged,
pinnately compound with terminal leaflet, lateral leaflets up to 9 pairs; leaflets opposite,
subopposite or alternate, often with hairy domatia in the axils of lateral veins, base broadly
rounded, symmetric. Inflorescences axillary or terminal panicles. Flowers bisexual; calyx 5lobed; petals 5, imbricate at the apical parts only, puberulous outside or on both surfaces;
stamens 10, those opposite calyx lobes longer, filaments subulate, glabrous, anthers dorsifixed;
disc intrastaminal, discoid or cup-shaped, hairy or glabrous; ovary superior, 5-carpellate, 5locular, hairy to sparsely hairy, carpels free except the basal and apical parts, 14 of them
abortive, styles 5, connate at the apical part, stigma capitate. Fruits 5-locular or appearing 1locular due to abortion, each locule with a distinct operculum; endocarp woody, hard. Seeds with
testa free from endocarp; embryo straight, cotyledons fleshy, free, plano-convex; germination
epigeal.
Distribution. About 8 species, distributed from India to Myanmar, Thailand, Cambodia, China,
Malesia, and Fiji. Two species in Sabah and Sarawak.
Ecology. Lowland primary and secondary forests, often by streams.
Uses. The timber is rather soft and not durable. The coloured heartwood is suitable for making
high quality cabinets and other furniture. This is an important export timber from New Guinea.
However, the timbers from New Guinea and the Philippines are heavier than those from Sabah
and Sarawak. Sometimes D. dao is planted for its edible fruits. The rounded bushy crown of D.
dao is very attractive and so is useful for planting as avenue trees.
Key to Dracontomelon species
Leaflets without domatia; midrib sunken above. Disc glabrous. Fruits globose or broadly
ellipsoid, usually 1-locular...............................................................1. D. costatum
Leaflets with hairy domatia; midrib raised above. Disc hairy. Fruits globose, distinctly 5locular............................................................................................................................2. D. dao
1. Dracontomelon costatum Blume

(Latin, costatus = ribbed; the strongly veined leaves)
17
l.c. (1850) 232; Merrill l.c. (1921) 350, l.c. (1929) 168; Masamune l.c. 409; Burgess l.c. 9; Ding Hou, FM 1, 8 (1978) 473;
Whitmore, Tantra & Sutisna l.c. 14. Type: Korthals 1836, Borneo, Martapura (holotype L; isotype BO).
Tree to 30 m tall, 65 cm diameter; bole with tall buttresses. Bark cracking. Leaves with 47 pairs
of leaflets. Leaflets elliptic to oblong, 722 x 48 cm, without domatia; base almost rounded,
apex acute; midrib sunken above, lateral veins 711 pairs, raised below, faint above, intercostal
veins scalariform-reticulate, visible on both surfaces; petiolules to 0.5 cm long. Flowers pale
yellow, in 70 cm long hairy panicles; disc glabrous. Fruits ovoid or broadly ellipsoid, c. 2 x 1.5
cm, black when ripe, usually 1-locular.
Distribution. Sumatra, Borneo. Widely distributed in Sabah but nowhere abundant, not yet
reported from Sarawak; also in Brunei and Kalimantan.
2. Dracontomelon dao (Blanco) Merr. & Rolfe
Fig. 5.
(from a Philippine plant name dao)

Philip. J. Sc. 3 (1908) Bot. 108; Merrill, Spec. Blanc. (1918) 234; Ding Hou, Blumea 24 (1978) 6, FM 1, 8 (1978) 470;
Anderson l.c. 136; Corner l.c. 113; Kochummen l.c. 20; Whitmore, Tantra & Sutisna l.c. 14; Ng l.c. 17; Kessler & Sidiyasa
l.c. 39. Basionym: Paliurus dao Blanco, Fl. Filip. (1837) 174. Type: Merrill Spec. Blanc. 316, Philippines, Mindanao,
Butuan Subprovince (L). Synonyms: Paliurus edulis Blanco, Fl. Filip. (1837) 173; D. edulis (Blanco) Skeels, US Dept.
Agric. Bur. Pl. Ind. Bull. 261 (1912) 52, Masamune l.c. 410; D. edule (Blanco) Merr., Philip. J. Sc. 10 (1915) Bot. 33; D.
mangiferum Blume l.c. (1850) 231, Merrill l.c. (1921) 351, Masamune l.c. 410; D. sylvestre Blume l.c. (1850) 231, Merrill
l.c. (1921) 351, Masamune l.c. 410; D. brachyphyllum Ridl., Kew Bull. (1933) 202, Masamune l.c. 409.
Tree to 40 m tall, 100 cm diameter; bole with steep buttresses to 4 m high. Bark grey-brown,
smooth to scaly; inner bark pink. Sapwood white. Twigs grey-brown or grey-white, 0.52.5 cm
thick. Leaves clustered towards ends of twigs, each with 59 pairs of subopposite or alternate
leaflets, rachis densely brown short-hairy or glabrous; petioles sometimes flattened or channelled
above with sharp edges. Leaflets glabrous or sparsely hairy below, with hairy domatia in the axils
of lateral veins; elliptic, oblong or ovate, 522 x 2.58 cm; base cuneate or rounded, often
unequal, apex acute with distinct tip; midrib raised above, lateral veins 620 pairs, curving and
joining near margin, raised on both surfaces, intercostal veins scalariform-reticulate, visible
below, faint above; petiolules 3 mm long. Flowers white, in 50 cm long hairy panicles; disc
hairy. Fruits globose, distinctly 5-locular, to 3 cm across, smooth or irregularly angular, edible.
Distribution. Eastern India, Andamans, Myanmar, Thailand, Cambodia, China, Malesia and
Solomon Islands. Common and widely distributed in Sabah and Sarawak; also in Brunei and
Kalimantan.
Ecology. Lowlands to 200 m, common by streams. Seeds germinate within 2867 days. The first
2 leaves opposite, trifoliolate, subsequent leaves pinnately compound, leaflets in early stage with
dentate margins.
18
Fig. 5. Dracontomelon dao. A, leaf; B, detail of lower leaflet surface; C, inflorescence; D, flower with
few sepals, petals and stamens removed; E, fruit. (A, B and E from S. 35022, CD from SAN 89946.)
19
6. DRIMYCARPUS Hook. f.
(Greek, drimus = pungent, karpos = fruit)
In Bentham & Hooker, Gen. Pl. 1 (1862) 424; Ding Hou, Blumea 24 (1978) 6, FM 1, 8 (1978) 520; Kochummen, TFM 4
(1989) 21; Whitmore, Tantra & Sutisna, CLK 1 (1989) 14.
Trees. Leaves spirally arranged, simple, margins thickened. Inflorescences axillary and/or
terminal panicles. Flowers unisexual (very rarely bisexual); calyx 5- or 4-lobed; petals 5 or 4,
imbricate, glabrous except for the sparsely hairy margins; stamen 5 or 4, anthers dorsifixed,
imperfect or aborted in female flowers; disc intrastaminal, slightly concave, 5 or 4-notched;
ovary inferior, abortive and rudimentary in male flower, 1-locular, 1-ovulate; style short,
cylindric, stigmas 3, capitate. Fruits 1-locular, crowned by persistent calyx. Seeds with testa
adnate to the endocarp; cotyledons free, plano-concave.
Distribution. Three species; India, Sikkim, Bhutan, Myanmar, Thailand, Vietnam, Sumatra,
Peninsular Malaysia, Borneo; two species in Sabah and Sarawak.
Key to Drimycarpus species

Leaves thin-coriaceous; intercostal veins faintly visible on both surfaces. Twigs terete,
slender.........................................................................................1. D. luridus
Leaves thick-coriaceous; intercostal veins visible below, very faint to inconspicuous above.
Twigs angular with sharp edges...........................................................2. D. maximus
1. Drimycarpus luridus (Hook. f.) Ding Hou
Fig. 6BE.
(Latin, luridus = dingy brown; perhaps referring to the patches on the twigs)
Blumea 24 (1978) 6, FM 1, 8 (1978) 520; Kochummen l.c. 21; Whitmore, Tantra & Sutisna l.c. 14. Basionym: Semecarpus
lurida Hook. f., Fl. Br. Ind. 2 (1876) 34. Type: Maingay 495, Malacca (K). Synonym: Swintonia lurida (Hook. f.) King, J.
As. Soc. Beng. 65, 3 (1896) 491.
Medium-sized tree to 27 m tall, 40 cm diameter. Bark grey-brown, smooth; inner bark brown
turning purplish on exposure. Twigs terete, slender, dark brown with grey patches. Leaves thincoriaceous; elliptic, oblong or oblanceolate, 836.5 x 210 cm; base cuneate, sometimes
asymmetric, apex acuminate, acumen c. 1 cm long; midrib raised above, lateral veins 1022 pairs,
with short intermediate veins, faintly visible on both surfaces, intercostal veins reticulate, faintly
visible on both surfaces; petioles 1.54 cm long, lower half slightly swollen, rugose on drying.
Inflorescences axillary, 211 cm long; male flowers with distinct pedicels, bisexual flowers
sessile. Fruits transversely oblong, 812 x 1020 mm.
Distribution. Sumatra, Peninsular Malaysia, Borneo. Very uncommon in Sabah, recorded from
Kinabatangan, Kudat and Sandakan districts, and of very restricted distribution in Sarawak,
known from Anap and Ulu Belaga only; also in Brunei and Kalimantan.
Ecology. Lowland to hill forest, at 601000 m.
20
Fig. 6. Drimycarpus luridus (BE); D. maximus (A). A and B, fruiting leafy twigs; C, flowering leafy
twig; D, male flower in longitudinal section; E, female flower in longitudinal section. (A from S.
43483, B from S. 22107, C from SAN 4750, DE after FM 1, 8 (1978) 519, f. 56 b and e.)
21
2. Drimycarpus maximus Kochummen
Fig. 6A.
(Latin, maximus = greatest; the large leaves and fruits)

Sandakania 7 (1996) 81. Type: Othman et al. S. 43483, Borneo, Sarawak, Ulu Sg. Belaga, 7th Division (holotype KEP;
isotypes K, L, MO, SAN, SAR).
Medium-sized tree to 23 m tall, 30 cm diameter. Bark grey, smooth. Twigs brownish, angular
with sharp edges. Leaves thick-coriaceous; oblanceolate, 3435.5 x 8.510 cm; base cuneate to
attenuate, apex acuminate, acumen c. 1 cm long; midrib raised above, strongly keeled below,
lateral veins 2225 pairs, raised below, almost invisible above, with short intermediate veins,
intercostal veins reticulate, visible below, very faint to inconspicuous above; petioles stout, c. 3.5
cm long, strongly rugose on drying. Flowers unknown. Infructescens axillary, 23 cm long.
Fruits transversely oblong, c. 2 x 1.2 cm; stalks c. 0.7 cm long.
Distribution. Endemic to Borneo; so far known only from the type specimen from Sarawak.
Ecology. Ridge mixed dipterocarp forest.
7. GLUTA L.
(Latin, gluten = viscous; the exudate)
rengas (Malay)
Mant. 2 (1771) 293; King, J. As. Soc. Beng. 65, 3 (1896) 480; Merrill, EB (1921) 349; Ridley, FMP 1 (1922) 526;
Masamune, EPB (1942) 410; Browne, FTSB (1955) 48; Burgess, TBS (1966) 12; Backer & Bakhuizen f., FJ 2 (1965) 150;
Ding Hou, Blumea 24 (1978) 8, FM 1, 8 (1978) 446; Anderson, CLTS (1980) 134; Wong, DMT (1982) 70; Corner, WSTM
3rd ed. 1 (1988) 127; Kochummen, TFM 4 (1989) 22; Whitmore, Tantra & Sutisna, CLK 1 (1989) 14; Ng, Mal. For. Rec.
34, 1 (1991) 18; Kessler & Sidiyasa, Trees of the Balikpapan-Samarinda Area (1994) 40. Synonym: Melanorrhoea Wall.,
Pl. As. Rar. 1 (1829) 9, Merrill l.c. (1921) 350, Masamune l.c. 411, Anderson l.c. 137.
Small, medium-sized or large trees, rarely shrubs; bole with buttresses, rarely with stilt-roots
(in swampy habitats). Bark smooth, cracking, fissured, scaly or flaky, often with tar-like
resinous exudate on the surface; inner bark pinkish or reddish, with black sap or with white
or colourless sap which turns black on exposure. Sapwood white. Leaves simple, spirally
arranged or in pseudowhorls (G. sabahana), margin not thickened. Inflorescences axillary
panicles; bracts and bracteoles caducous; pedicels articulate or not. Flowers bisexual; calyx
calyptriform (hood-like) or bursting irregularly at anthesis, caducous; floral axis between the
calyx and ovary (torus) often elongated and enlarged; petals 5, rarely 4 or 8, caducous or
persistent and enlarged in fruit; stamens 5, 10 or many, rarely 4 or 7, inserted on the torus,
filaments filiform, glabrous or hairy, anthers dorsifixed; disc absent; ovary superior, with or
without stalk, glabrous or hairy, 1-locular, style 1, distinct, terminal or lateral, stigma 1,
slightly thicker than style. Fruits stalked or sessile, 1-locular, smooth or rugose, glabrous or
hairy, with or without persistent, enlarged, wing-like petals. Seeds one per fruit; testa adnate
to the endocarp; embryo straight, rarely slightly curved, cotyledons free or incompletely
fused and partly free on one side only; germination hypogeal, cotyledons non-emergent.
22
Distribution. Thirty species; Madagascar, India, Myanmar, Thailand, Indo-China, and Malesia. Ten
Ecology. Lowland including tidal estuary, fresh-water swamp and peat swamp, to lower montane
forest, to 1300 m.
Uses. The timber is medium hardwood; grain shallowly interlocking; texture rather fine and even.
Sapwood white or straw-coloured, sharply differentiated from the red heartwood. Heartwood bright
red with alternating darker coloured layers simulating that of Douglas fir on flat sawn faces, capable
of taking a very smooth, highly polished finish; moderately durable, and used for making excellent
furnitures. However, the irritant sap discourages its wide use.
Key to Gluta species
(based on vegetative and fruit characters)
1. Leaf apex acute (or rarely obtuse).....................................................................................................2
Leaf apex usually rounded or emarginate.........................................................................................6
2. Leaf midrib sunken above.........................................................................................10. G. wallichii
Leaf midrib raised or flattened above...............................................................................................3
3. Trees of tidal rivers, stilt-rooted. Petioles to 5 mm long.............................................9. G. velutina
Trees of inland forests, without stilt-roots. Petioles more than 5 mm long.....................................4
4. Leaves in pseudowhorls............................................................................................7. G. sabahana
Leaves spirally arranged and well-spaced along the twigs..............................................................5
5. Petioles strongly swollen at base. Fruit without wings..............................................3. G. laxiflora
Petioles only slightly swollen at base. Fruits with wings....................................4. G. macrocarpa
6. Leaves hairy below.....................................................................................................8. G. speciosa
Leaves glabrous below....................................................................................................................7
7. Leaves larger, 14.526.5 x810 cm; lateral veins 1526 pairs.................................6. G. rugulosa
Leaves smaller, (3)515(19) x (2)36(8.5) cm; lateral veins to 12 pairs................................8
8. Leaves usually elliptic or only occasionally obovate........................................................5. G. oba
Leaves always obovate....................................................................................................................9
9. Fruits without wings......................................................................................................1. G. aptera
Fruits with long wings................................................................................................2. G. beccarii
Key to Gluta species

(based on flower and fruit characters)
1. Stamens 5 (rarely to 7); calyx bursting irregularly at anthesis........................................................2
Stamens 10 or more; calyx calyptriform (hood-like) at anthesis....................................................5
23
2. Pedicels articulate................................................................................................................3
Pedicels not articulate..........................................................................................................4
3. Fruits with wings..........................................................................................10. G. wallichii
Fruits without wings.......................................................................................9. G. velutina
4. Leaves in pseudowhorls...............................................................................7. G. sabahana
Leaves spirally arranged and well-spaced along the twigs.............................3. G. laxiflora
5. Pedicels articulate................................................................................................................6
Pedicels not articulate..........................................................................................................7
6. Stamens 2028. Petals in fruit only slightly enlarged..............................4. G. macrocarpa
Stamens about 70. Petals in fruit 56 cm long...............................................2. G. beccarii
7. Stamens 10.............................................................................................................5. G. oba
Stamens 40100..................................................................................................................8
8. Stamens c. 40. Fruit with enlarged petals (wings)......................................6. G. rugulosa
Stamens c.100. Fruit without enlarged petals (wings).........................................................9
9. Calyx densely hairy outside............................................................................8. G. speciosa
Calyx glabrous outside except for tuft of hairs at apex......................................1. G. aptera
1. Gluta aptera (King) Ding Hou
Fig. 7AB.
(Greek, a- = without, pteron = wing; the fruit)

Blumea 24 (1978) 12, FM 1, 8 (1978) 452; Kochummen l.c. 24; Whitmore, Tantra & Sutisna l.c. 14. Basionym:
Melanorrhoea aptera King, J. As. Soc. Beng. 65, 3 (1896) 487. Syntypes: Kings collector 3485 (BM, K, L), 3727 (K,
SING), 7636 (SING), Peninsular Malaysia, Perak; Curtis 1567, Penang (K). Synonym: Melanorrhoea tricolor Ridl., Kew
Bull. (1933) 196, Masamune l.c. 412.
Medium-sized to large tree to 30 m tall, 70 cm diameter. Bark grey-brown, smooth. Twigs brown,
youngest ones reddish brown-hairy. Leaves spirally arranged and well-spaced, thick-coriaceous,
glabrous below; obovate, (3)4.519 x (2)2.58.5 cm; base attenuate, apex rounded or emarginate;
midrib flattened (rarely sunken) above, lateral veins 710 pairs, faint to distinct above, intercostal
veins reticulate, faint to distinct below, faint to invisible above; petioles 12 cm long. Flowers white;
pedicels 1023 mm long, glabrous, not articulate; calyx calyptriform, glabrous except for tuft of hairs
at the apex; petals oblanceolate, 1116 x35 mm, densely hairy outside; stamens about 100; ovary
glabrous; style subterminal. Fruits globose to subglobose, 2.53.5 cm diameter, brown, smooth,
without wings; stalks c. 0.5 cm long.
Distribution. Sumatra, Peninsular Malaysia, Borneo. Widely distributed in Sarawak, uncommon in
Sabah; also in Brunei and Kalimantan.
Ecology. Lowland, including peat swamp, to lower montane forest, at 7001200 m. Flowering in
March to October.
24
Fig. 7. Gluta aptera (AB); G. sabahana (CE); G. wallichii (F). A and C, flowering leafy twigs; B and D
flowers in longitudinal section; EF, fruits. (AB from S. 25253, CD from SAN 60402, E from SAN 110836, F
from Ashton BRUN 3309.)
25
2. Gluta beccarii (Engl.) Ding Hou

(Odoardo Beccari, 18431920, Italian explorer and botanist)
Blumea 24 (1978) 13, FM 1, 8 (1978) 452; Anderson l.c. 137; Kochummen l.c. 24; Whitmore, Tantra & Sutisna l.c. 15.
Basionym: Melanorrhoea beccarii Engl., Bot. Jahrb. 1 (1880) 45, Merrill l.c. (1921) 350, Masamune l.c. 411, Anderson l.c.
137. Type: Beccari PB 1484, Borneo, Sarawak (isotype K).
Small to medium-sized tree to 25 m tall, 45 cm diameter; bole with short buttresses. Bark rusty
brown, smooth; inner bark pinkish brown. Sapwood whitish. Terminal buds pale silky hairy.
Leaves spirally arranged and well-spaced, thick-coriaceous, glabrous below; obovate, 4.59.5
x 25 cm; base attenuate, apex emarginate or rounded; midrib flattened above, lateral veins 810
pairs, faint on both surfaces, intercostal veins reticulate, faintly visible below, inconspicuous
above; petioles 0.52 cm long. Flowers white turning to pink; pedicels 38 mm long, sparsely
hairy, articulate; calyx calyptriform, hairy outside; petals hairy outside; stamens c. 70, filaments
hairy; ovary glabrous; style terminal. Fruits bright purplish red when fresh, subglobose, c. 1.5 cm
across; wings 56 cm long.
Distribution. Peninsular Malaysia, Borneo. Common in Sarawak, uncommon in Sabah; also in
Ecology. Fresh-water swamp, peat swamp and ridge forests, to 300 m.
Note. This species can be confused with G. aptera, but in that species the pedicel is not articulate,
the calyx is glabrous and the fruit is without wings.
3. Gluta laxiflora Ridl.

(Latin, laxus = loose, distant, florum = flowers; the laxly arranged flowers)
Kew Bull. (1933) 196; Masamune l.c. 410; Ding Hou, FM 1, 8 (1978) 456; Anderson l.c. 136; Whitmore, Tantra & Sutisna
l.c. 15. Type: Hose 185, Borneo, Sarawak, Baram (holotype K).
Small tree to 20 m tall, 30 cm diameter. Bark dark grey, scaly. Twigs grey-brown. Leaves
spirally arranged and well-spaced; elliptic or oblong, 1019.5 x 37 cm; base cuneate,
sometimes asymmetric, apex acute; midrib raised above, lateral veins 1014 pairs, raised below,
faint above, intercostal veins finely reticulate, very faint on both surfaces; petioles 1.55 cm long,
base strongly swollen and cracked. Flowers in hairy panicles; pedicels c. 4 cm long, not
articulate; calyx bursting irregularly; petals hairy outside; stamens 5, rarely 6; ovary hairy, style
lateral. Fruits obliquely or broadly ellipsoid, 7.59 x 56 cm, brown or reddish brown scurfy,
without wings; stalks obscure.
Distribution. Endemic to Borneo; known only from Sarawak and Brunei.
4. Gluta macrocarpa (Engl.) Ding Hou

(Greek, makros = large, karpos = fruit)
26
Blumea 24 (1978) 14, FM 1, 8 (1978) 454; Kochummen l.c. 28; Whitmore, Tantra & Sutisna l.c. 15. Basionym:
Melanorrhoea macrocarpa Engl. in A. DC., Mon. Phan. 4 (1883) 236, Merrill l.c. (1921) 350, Masamune l.c. 412, Anderson
l.c. 137. Type: Beccari PB 3051, Borneo, Sarawak (isotype K).
Tree to 36 m tall, 55 cm diameter; bole with steep buttresses to c. 1.5 m high. Bark smooth to
rough, grey-brown; inner bark reddish, with white latex soon turning black on exposure.
Sapwood whitish. Young twigs densely rusty brown short-hairy. Leaves spirally arranged and
well-spaced, glabrous (softly hairy below when young); oblanceolate or elliptic, 7.52 x 2.55
cm, rarely larger to 25 cm x9.5 cm; base attenuate, apex acute; midrib flattened above, lateral
veins 1224 pairs, curving and joining near margin, visible on both surfaces, intercostal veins
reticulate, faint; petioles 13.5 cm long, slightly swollen at base. Flowers in hairy panicles;
pedicels 13.5 cm long, articulate; calyx calyptriform, hairy outside; petals densely hairy outside;
stamens 20, rarely to 28, with few staminodes, glabrous; ovary with terminal style. Fruits
purplish black, smooth, globose, 24 cm diameter, with a central stalk and short wings.
Distribution. Peninsular Malaysia, Borneo. Of scattered occurrence in Sabah and Sarawak; also
Ecology. Hill forest at 400900 m. Flowering in August to September and fruiting in October to
February.
5. Gluta oba (Merr.) Ding Hou

(from the vernacular name oba)
Blumea 24 (1978) 14, FM 1, 8 (1978) 454; Whitmore, Tantra & Sutisna l.c. 15. Basionym: Melanorrhoea oba Merr., J. Str.
Br. Roy. As. Soc. 77 (1917) 190, l.c. (1921) 350, Masamune l.c. 412, Anderson l.c. 137. Type: Villamil 316, North Borneo
(isotypes K, SING).
Medium-sized tree to 30 m tall, 35 cm diameter; bole with steep buttresses to 3 m high. Bark
reddish brown with grey patches, cracking and scaly; inner bark reddish. Sapwood whitish.
Twigs dark brown, glabrous. Leaves spirally arranged and well-spaced, coriaceous, drying
chocolate-brown, glabrous; elliptic or occasionally obovate, 815 x 46 cm; base cuneate, apex
usually rounded, very rarely obtuse; midrib flattened above, lateral veins 912 pairs, very faintly
visible on both surfaces, intercostal veins reticulate, faint; petioles 0.71.5 cm long. Flowers in
glabrous panicles; pedicels c. 1 cm long, not articulate; calyx calyptriform, glabrous except the
hairy apical part; petals hairy outside; stamens 10, filaments hairy; ovary glabrous, style terminal.
Fruits subglobose, 34 cm across, dark brown, smooth, without wings; stalks centrally attached,
11.5 cm long.
Distribution. Endemic to Borneo. Common in Sabah and Sarawak, especially on Pulau Sakar in
Sabah; also in Brunei and Kalimantan.
Ecology. Lowand mixed dipterocarp forest. Flowering in February to March and fruiting in April
to June.
6. Gluta rugulosa Ding Hou

(Latin, rugulosus = somewhat wrinkled; the fruits)
27
Blumea 24 (1978) 16; FM 1, 8 (1978) 452; Whitmore, Tantra & Sutisna l.c. 15. Type: Hose 41, Borneo, Sarawak, Baram
(holotype L; isotypes BM, CGE, K).
Tree to 30 m tall, 60 cm diameter; bole with buttresses to 3 m high. Bark brown, scaly; inner
bark pinkish. Sapwood white. Leaves spirally arranged and well-spaced, glabrous below;
obovate, 14.526.5 x 810 cm; base attenuate, apex rounded or rarely bluntly acute; midrib
flattened (very rarely sunken) above, lateral veins 1526 pairs, raised below, faint above, curving
and joining near margin, intercostal veins scalariform, faintly visible below, invisible above;
petioles 12 cm long, base swollen. Flowers hairy; pedicels not articulate; calyx calyptriform,
hairy outside; stamens c. 40; ovary scurfy, style terminal. Fruits globose, c. 3.5 cm across, light
brown, scurfy, rugose, sessile; wings 2.53 cm long.
Distribution. Endemic to Borneo. Uncommon in Sabah and Sarawak; also in Brunei and
Kalimantan.
Ecology. Lowland forests including kerangas.
7. Gluta sabahana Ding Hou
Fig. 7CE.
(of Sabah)
Blumea 24 (1978) 16, FM 1, 8 (1978) 455; Whitmore, Tantra & Sutisna l.c. 15. Type: Sinanggul SAN 40615, Borneo,
Sabah, Tawau (holotype L; isotype SAN).
Medium-sized tree to 30 m tall, 60 cm diameter. Bark dark brown, smooth to scaly. Leaves in
pseudowhorls, coriaceous, glabrous; oblanceolate or elliptic, 1323 x 38 cm; base cuneate to
attenuate, apex acute; midrib raised and prominent above, lateral veins 915 pairs, faintly raised
on both surfaces, intercostal veins reticulate, faint; petioles 0.51.5 cm long, grey-brown and
cracked. Flowers in hairy panicles; pedicels 36 mm long, not articulate; calyx bursting
irregularly, hairy outside; petals whitish, hairy outside; stamens 5, rarely 7; ovary hairy, style
lateral. Fruits broadly ellipsoid, 7.59 x 56.5 cm, brownish scurfy, without wings; stalks
obscure.
Distribution. Endemic to Borneo; so far known from Sabah only where it is common.
Ecology. Lowland forest below 30 m, including swamp. Flowering in February to September and
fruiting in May to December.
8. Gluta speciosa (Ridl.) Ding Hou

(Latin, speciosus = showy; the flowers)
Blumea 24 (1978) 21, FM 1, 8 (1978) 450; Whitmore, Tantra & Sutisna l.c. 16. Basionym: Melanorrhoea speciosa Ridl.,
Kew Bull. (1933) 197, Masamune l.c. 412, Anderson l.c. 138. Type: Haviland 3147, Borneo, Sarawak (holotype K; isotypes
L, SING).
Large tree to 40 m tall, 80 cm diameter; bole with buttresses to 1.5 m high. Bark dark brown,
fissured; inner bark pink. Sapwood pale pink. Leaves spirally arranged and well-spaced,
thick-coriaceous, finely hairy below; oblong to narrowly obovate, 517.5 x 39 cm; base
28
cuneate, apex rounded or emarginate; midrib flattened above, lateral veins 1022 pairs,
raised below, visible above, intercostal veins scalariform-reticulate, distinct below, faint
above; petioles 12 cm long, densely brown short-hairy. Flowers in hairy panicles; pedicels
1022 mm long, not articulate; calyx calyptriform, densely hairy outside; petals white, hairy
outside; stamens c. 100; ovary hairy, style terminal. Fruits subglobose, 23 cm across,
smooth, without wings.
Distribution. Endemic to Borneo; uncommon in Sabah and Sarawak; in Sabah, known from
Ranau districts only from two collections, SAN 22369 and SAN 62741, while in Sarawak, of
scattered distribution; also in Brunei and Kalimantan.
Ecology. Lowland, including swamp, to hill forests to 600 m.
9. Gluta velutina Blume

(Latin, velutinus = velvety hairy; the inflorescences)
Mus. Bot. Lugd. Bat. 1 (1850) 183; Merrill l.c. (1921) 349; Masamune l.c. 410; Burgess l.c. 12; Ding Hou, FM 1, 8 (1978)
461; Anderson l.c. 136; Corner l.c. 128; Kochummen l.c. 30; Whitmore, Tantra & Sutisna l.c. 16. Type: Sine coll., s.n.,
Borneo (L; n.v.).
Large shrub or small tree to 7 m tall; bole with stilt-roots. Bark brownish, smooth. Leaves
spirally arranged and well-spaced; oblanceolate or elliptic, 1319 x 47 cm; base cuneate,
apex acute; midrib flattened above, lateral veins 1518 pairs, faintly visible on both surfaces,
intercostal veins reticulate, faint; petioles to 5 mm long. Flowers in hairy panicles; pedicels
short, c. 1 mm long, articulate; calyx bursting irregularly, hairy outside; petals white,
glabrous; stamens 5; ovary glabrous, style lateral. Fruits subglobose, 4.57.5 cm diameter,
pale brown, with irregular warty ridges, especially towards the base, without wings.
Distribution. Myanmar, Thailand, Vietnam, Sumatra, Peninsular Malaysia, Borneo, Java.
Common in Sabah and Sarawak; also in Brunei and Kalimantan.
Ecology. Edges of tidal rivers, standing in water with submerged trunk except at low tides,
associated with Barringtonia conoidea and Pandanus helicopus.
10. Gluta wallichii (Hook. f.) Ding Hou
Fig. 7F.
(N. Wallich, 17861854, Danish botanist at Calcutta)

Blumea 24 (198) 21, FM 1, 8 (1978) 456; Kochummen l.c. 30; Whitmore, Tantra & Sutisna, l.c. 16; Kessler & Sidiyasa l.c.
41. Basionym: Melanorrhoea wallichii Hook.f., Fl. Br. Ind. 2 (1876) 25, Anderson l.c. 138. Lectotype (Ding Hou, 1978):
Wall. Cat. 980, Singapore (K). Synonyms: Melanorrhoea maingayi Hook.f. l.c. 25, Merrill l.c. (1921) 350, Masamune l.c.
412; Swintonia elmeri Merr., PEB (1929) 167.
Medium-sized to large tree to 36 m tall, 50 cm diameter; bole with short buttresses. Bark
grey-brown, fissured and scaly; inner bark reddish with watery whitish sap turning black on
exposure. Sapwood whitih. Leaves spirally arranged and well-spaced, glabrous, rarely
sparsely brown short-hairy below; oblong, elliptic or narrowly obovate, 8.534 x 414 cm;
29
base broadly cuneate to rounded, apex acute or rarely obtuse; midrib sunken above, lateral
veins 1720 pairs, raised below, faint above, intercostal veins reticulate, distinct below, faint
above; petioles 25 cm long, glabrous or sparsely brown short-hairy, distinctly channelled
above. Flowers in hairy panicles; pedicels 23 mm long, articulate; calyx bursting
irregularly, hairy outside; petals white, densely hairy on both surfaces; stamens 5; ovary
hairy, style lateral. Fruits ovoid or ellipsoid, c. 1.5 x 1 cm, smooth, brownish, with enlarged
bright red wings; stalks obscure.
Distribution. Sumatra, Peninsular Malaysia, Borneo. Common and widely distributed in
Ecology. Lowland, including swamp, to hill forest to 500 m. Flowering from March to
November and fruiting from March to August.
8. KOORDERSIODENDRON Engl.
(S.H. Koorders, 18631919, forester and botanist in the
former Dutch East Indies Forest Service at Bogor, Java)
ranggu (preferred name in ASEAN countries)
In Kooders, Med. Lands Pl. Tuin 19 (1898) 410; Merrill, EB (1921) 350; Masamune, EPB (1942) 410; Smythies, CST
(1965) 3; Ding Hou, FM 1, 8 (1978) 486; Anderson, CLTS (1980) 136; Wong, DMT (1982) 110; Whitmore, Tantra, &
Trees. Inner bark with scanty white latex. Leaves spirally arranged, pinnately compound with
terminal leaflet. Leaflets opposite or subopposite, 1016 pairs, without domatia in the axils of
lateral veins. Inflorescences axillary, paniculate. Flowers bisexual, 5-merous; calyx 5-lobed;
petals 5, imbricate; stamens 10; disc intrastaminal, round and flat, 10-lobed; ovary superior,
subglobose, deeply 5-furrowed (carpels incompletely connate), densely hairy, 5-locular,
usually only one fertile, styles 5, short, stigmas small. Fruits 1(3)-locular, without wings.
Seeds with testa free from the endocarp; embryo straight, cotyledons free, plano-convex.
Distribution. Monotypic; Borneo, the Philippines, Sulawesi, Maluku, and Irian Jaya.
Koordersiodendron pinnatum (Blanco) Merr.
Fig. 8.
(Latin, pinnatus = with parts arranged on each side of a common axis; the compound leaves)
Bull. For. Bur. 1 (1903) 33, Spec. Blanc. (1918) 234, l.c. (1921) 350; Masamune l.c. 410; Smythies l.c. 3; Burgess l.c. 16;
Ding Hou, FM 1, 8 (1978) 486; Anderson l.c. 136; Whitmore, Tantra & Sutisna l.c. 16. Basionym: Helicteres pinnata
Blanco, Fl. Filip. (1837) 384. Neotype (Merrill, 1918): Merrill Spec. Blanc. 744, Philippines, Luzon, Angat, Bulacan
(isoneotype L).
Medium-sized to tall tree to 40 m tall, 90 cm diameter; bole with buttresses to 3 m high. Bark
dark brown or black, fissured. Twigs brownish, rough, with many small lenticels, 1.52 cm
30
Fig. 8. Koordersiodendron pinnatum. A, fruiting leafy twig; B, twig with inflorescences; C, flower in
longitudinal section. (All from SAN 39031.)
31
thick. Leaves 5080 cm long, each with 1016 pairs of leaflets; rachis not swollen at base.
Leaflets chartaceous, ovate-oblong, 320 x 1.55.5 cm; base slightly asymmetric, broadly
cuneate to rounded, apex acuminate, acumen c. 1 cm long; midrib raised above, lateral veins
1024 pairs, distinctly looping, faint on the upper leaflet surface, intercostal veins reticulate,
faintly visible on both surfaces; petiolules 35 mm long. Inflorescences c. 50 cm long,
paniculate. Flowers whitish. Fruits ellipsoid, yellowish when ripe, 2.54 x 1.52.5 cm.
Seeds ellipsoid, compressed.
Distribution. Borneo, Philippines, Sulawesi, Maluku, Irian Jaya. Common and widely
distributed in Sabah (Lahad Datu, Sandakan and Tawau districts); uncommon in Sarawak,
recorded only from northeast border with Sabah; also in Brunei and Kalimantan.
Ecology. Lowland forest, including swamps.
Uses. The timber is moderately hard and moderately heavy with density of 690915 kg/m3
air-dried. The sapwood is white to pale pink and is distinct from the heartwood which is
pinkish brown or reddish brown; texture is fine and even with straight to interlocking grains.
It is suitable for flooring, interior construction, furniture, cabinet making, and door-panels.
9. MANGIFERA L.
(Latin, mango = latinised local name mangga, ferre = bearer; mangga-bearing trees)
asam, machang (Malay)
Sp. Pl. (1753) 200, Gen. Pl. ed. 5 (1754) 93; King, J. As. Soc. Beng. 65, 3 (1896) 466; Merrill, EB (1921) 349; Ridley, FMP
1 (1922) 520; Mukherji, Lloydia 12 (1940) 73; Browne, FTSB (1955) 50; Backer & Bakhuizen f., FJ 2 (1965) 148; Burgess,
TBS (1966) 19; Ding Hou, Blumea 24 (1978) 21, FM 1, 8 (1978) 423; Anderson, CLTS (1980) 136; Wong, DMT (1982)
128; Corner, WSTM 3rd ed. 1 (1988) 115; Kochummen, TFM 4 (1989) 31; Whitmore, Tantra & Sutisna, CLK 1 (1989) 16;
Ng, Mal. For. Rec. 34, 1 (1991) 18; Bompard, PROSEA 2 (1991) 203; Kostermans & Bompard, The mangoes (1993) 1;
Kessler & Sidiyasa, Trees of the Balikpapan-Samarinda Area (1994) 41.
Tall emergent trees; crown dense, rounded; bole usually without buttresses. Bark cracking to
fissured and scaly; inner bark yellowish to brownish, laminated, with droplets of white
creamy or pinkish latex which blackens on drying, often with resinous smell. Twigs terete or
strongly angular. Leaves simple, spirally arranged, sometimes very close and appear
clustered, thick- or thin-coriaceous, glabrous, without peltate scales, margin not thickened;
midrib raised above. Inflorescences terminal or axillary panicles, glabrous or hairy, often
crowded at apex of twigs; pedicels articulate. Flowers bisexual or male; 4-or 5-merous;
calyx 45-lobed; petals free, often with glandular ridges inside, caducous; disc glandular,
pulvinate, cup-shaped, cylindric and stalk-like, or 4- or 5-lobed; stamens 5, rarely 1012,
usually 12 fertile, very rarely 35 fertile, not inserted on the torus, filaments distinct,
anthers 2-locular; staminodes if present very much smaller; ovary superior, one-locular, style
1, usually eccentric; stigma 1. Fruits reniform, globose or cylindrical, ripening green, yellow
with shades of red, brown or black; pulp (mesocarp) fleshy, juicy, sweet to sour, white to pale
to dark yellow, fragrant or stinking, often smelling of turpentine; endocarp hard, fibrous or
not. Seeds one per fruit; embryo filling
32
the seed, cotyledons massive, without endosperm; germination hypogeal or semi-hypogeal, in

some species lasting 8151 days, cotyledons mostly non-emergent, the first 2 leaves opposite,
subsequent leaves in spirals, produced intermittently in flushes.
Distribution. About 70 species; Sri Lanka, India, Myanmar, Thailand, Indo-China, S China,
Malesia, Solomon Islands. Sixteen species in Sabah and Sarawak. A number of species, including
2 introduced ones (M. pentandra and M. odorata), are cultivated for their edible fruits.
Ecology. Widely distributed from lowland to lower montane forest, to 1800 m, but nowhere
abundant.
Uses. Machang is the standard Malaysian name for the timber of Mangifera species. The timber
is a light hardwood with density of 545610 kg/m3 air-dried. The sapwood is not clearly defined
from the heartwood which is light pink-brown to light brown. In some trees, a streaky corewood
is present where the wood is dark brown interspersed with streaks of black colour. Texture
moderately fine and even with straight or interlocking grains. The timber is suitable for light
construction, planking, flooring, plywood manufacture etc. The streaky corewood is in demand as
a decorative timber and is used for high class cabinet work, interior finishing, panelling and
partitioning, sliced decorative veneers and also for furniture manufacture. The most important use
is the edible fruits produced by a few species, mainly M. indica and its cultivars which are
cultivated throughout the tropics and subtropics. Young leaves of some species are used as a
vegetable. Mango leaves and flowers play a significant role in Hindu religious ceremonies. A few
species are used in traditional medicine.
Taxonomy. The genus has been divided by Ding Hou (Blumea 24 (1978) 22) into two sections,
viz., Section Mangifera (characterised by the cup-shaped, papillose disc, which is usually 4- or 5lobed and completely or partly surrounding the ovary, and free stamen filaments) and Section
Limus (characterised by the cylindric or torus-like disc, which is located at the base of the ovary
in bisexual flowers and usually neither lobed nor papillose, and stamen filaments which are
usually connate at base).
Key to Mangifera species

(based mainly on floral and fruit characters)
(M. blommesteinii and M. subsessilifolia are not included due to lack of flowers)
1. Disc cup-shaped, pulvinate or obscure. Stamen filaments free...........................................2

Disc cylindric and stalk-like. Stamen filaments connate...................................................12
2. Disc obscure. Leaves linear, to 4 cm wide...............................................8. M. macrocarpa
Disc distinct. Leaves not linear, wider................................................................................3
3. Fertile stamens 35................................................................................................................
Mangifera pentandra Hook. f.
Fl. Br. Ind. 2 (1876) 14; King l.c 472; Ridley l.c. 522; Mukherji l.c. 81; Ding Hou, Blumea 24 (1978) 28, FM 1, 8
(1978) 426; Corner l.c. 120; Kochummen l.c. 38; Kostermans & Bompard l.c. 36. Lectotype (Ding Hou, 1978):
Maingay 471, Malacca (K).
33
Medium-sized tree to 28 m tall, 90 cm diameter. Leaves spirally arranged, drying

greenish, not papillose below, thin-coriaceous; oblong or elliptic, 1120 x 3.59 cm;
intercostal veins reticulate, distinctly visible on both surfaces; petioles 1.53.5 cm long,
swollen at base. Flowers 5-merous; each petal with 5 yellow ridges; stamens 5, fertile
stamens 35, filaments free; disc distinct, pulvinate; style slightly eccentric. Fruits oblong,
7.510 x 56.5 cm, green when mature, fragrant; pulp pale orange, sweet.
Endemic to Peninsular Malaysia. In Sabah, introduced and cultivated for its fruits.
Fertile stamen one..............................................................................................................4
4. Flowers 5-merous................................................................................................................5
Flowers 4-merous................................................................................................................7
5. Inflorescences pyramidal, to 40 cm long...........................................................6. M. indica
Inflorescences not pyramidal, to 15 cm long.......................................................................6
6. Leaf midrib pinkish (on drying) beneath; lower leaf surface papillose; petioles 12 cm
long.........................................................................................................13. M. rufocostata
Leaf midrib not pinkish (on drying) beneath; lower leaf surface not papillose; petioles 47 cm
long.............................................................................................15. M. swintonioides
7. Leaf intercostal veins very finely reticulate, visible on the lower leaf surface....................8
Leaf intercostal veins not reticulate, visible or invisible on the lower leaf surface...........10
8. Leaf apex rounded with short blunt tip; leaf margin thickened........7. M. khoonmengiana
Leaf apex acute; leaf margin not thickened.........................................................................9
9. Axillary buds persistent. Leaf margin not recurved...................................11. M. parvifolia
Axillary buds caducous. Leaf margin recurved........................................16. M. torquenda
10. Leaves thick-coriaceous, cardboard-like, 59 cm wide..............................9. M. magnifica
Leaves thin-coriaceous, 47 cm wide................................................................................11
11. Inflorescences hairy.......................................................................................5. M. griffithii
Inflorescences glabrous...........................................................................12. M. quadrifida
12. Fertile stamens 5; petals without ridges.......................................................3. M. decandra
Fertile stamens 12; petals with ridges..............................................................................13
13. Petals purplish inside.........................................................................................................14
Petals white to pinkish inside............................................................................................15
14. Petals with 3 ridges each................................................................................10. M. pajang
Petals with one ridge each..................................................................................2. M. caesia
15. Peduncles brick-red. Leaves thick-coriaceous, with invisible intercostal veins. Fruits with
strong turpentine odour....................................................................................4. M. foetida
34
Peduncles pinkish. Leaves thin-coriaceous, intercostal veins distinct. Fruits strongly fragrant but
without turpentine odour....................................................................................
M. odorata Griff.
Notul. Pl. As. 4 (1854) 417; King, l.c. 474; Ridley l.c. 524; Masamune l.c. 411; Mukherji l.c. 122; Backer &
Bakhuizen f. l.c. 149; Ding Hou, Blumea 24 (1978) 26, FM 1, 8 (1978) 437; Anderson l.c. 136; Bompard l.c. 218;
Corner l.c. 120; Kochummen l.c. 37; Whitmore, Tantra & Sutisna l.c. 17; Kostermans & Bompard l.c. 168; Kessler
& Sidiyasa l.c. 42. Type: Griffith 1098, Malacca (holotype K; isotype P).
Tree to 30 m tall, 65 cm diameter. Leaves spirally arranged, drying brownish, thincoriaceous; oblong-lanceolate, 1235 x410 cm; intercostal veins reticulate, distinct;
petioles 37 cm long, swollen at base. Inflorescences pseudoterminal, pyramidal panicles, to
45 cm long; peduncles pinkish. Flowers 5-merous (very rarely 6-merous), fragrant; sepals
dark red outside; petals pale pinkish inside towards apex, each with 35 ridges; stamens 5,
fertile stamen one, filaments connate at base; disc stalk-like; style lateral. Fruits obliquely
ovoid, ellipsoid or oblong, 1013 x69 cm, yellow-green with numerous yellow or brown
dots, strongly fragrant; pulp yellow, sweet.
The species has never been found in the wild. Commonly cultivated in Thailand, Sumatra,
Peninsular Malaysia, Java, Borneo, Sulawesi, and the Philippines. In Sabah and Sarawak,
cultivated for the edible fruits, to 1000 m. Vernacular names: Sabahhuani, kwini, wani
(Malay). Sarawakkwini (Malay).
Key to Mangifera species

(based on vegetative and flower characters)
1. Leaves drying brownish or chocolate-brown......................................................................2

Leaves drying greenish, greenish yellow or pale yellow...................................................10
2. Petioles absent or to 1 cm long only....................................................14. M.
subsessilifolia
Petioles at least 1 cm long....................................................................................................3
3. Leaves in pseudowhorls on the twigs..................................................................................4
Leaves spirally arranged, sometimes in pseudowhorls at apex of twigs only......................6
4. Leaves smaller, to 8.5 x3 cm, apex rounded and emarginate; lateral veins faint; petioles
less than 2 cm long...............................................................................1. M. blommesteinii
Leaves larger, to 38 x15 cm, apex acute; lateral veins raised on both surfaces; petioles
26 cm long.........................................................................................................................5
5. Flowers reddish or pinkish; fertile stamens 5...............................................3. M. decandra
Flowers violet or lilac; fertile stamens 12........................................................2. M. caesia
6. Leaf intercostal veins invisible............................................................................................7
Leaf intercostal veins visible to very conspicuous...............................................................8
7. Petioles 23.5 cm long, slightly swollen at base..............................................4. M. foetida
Petioles 3.59 cm long, strongly swollen at base...........................................10. M. pajang
35
8. Leaves very thick, cardboard-like...............................................................9. M. magnifica

Leaves thinner in texture.....................................................................................................9
9. Leaf intercostal veins very finely reticulate beneath. Leaf margin thickened........................
..........................................................................................................7. M. khoonmengiana
Leaf intercostal veins not so. Leaf margin not thickened.......................................................
..................................................................................M. odorata Griff. (see preceding key)
10. Leaves papillose beneath; midrib pinkish (on drying) beneath...............13. M. rufocostata
Leaves not papillose beneath; midrib not so......................................................................11
11. Leaves linear, 1644 x 24 cm................................................................8. M. macrocarpa
Leaves not linear, wider.....................................................................................................12
12. Leaf intercostal veins invisible or finely reticulate and distinct beneath...........................13
Leaf intercostal veins visible, not reticulate, distinct or faint beneath...............................14
13. Leaf lateral veins 1420 pairs, raised on both surfaces. Leaf margins recurved....................
..................................................................................................................16. M. torquenda
Leaf lateral veins 810 pairs, faint on both surfaces. Leaf margins not recurved..................
...................................................................................................................11. M. parvifolia
14. Leaf intercostal veins faintly visible beneath. Petioles usually almost half the length of
leaf blade.............................................................................................15. M. swintonioides
Leaf intercostal veins distinct beneath. Petioles much less than half the length of
leaf blade..........................................................................................................................15
15. Plants known only through cultivation; fertile stamens 35..................................................
.......................................................................................M. pentandra (see preceding key)
Plants known in wild and in cultivation; fertile stamen 1..................................................16
16. Leaves drying greenish, margins often wavy. Flowers 5-merous, petals yellowish to
pale green.........................................................................................................6. M. indica
Leaves drying to pale yellow or yellowish green, margins not wavy. Flowers 4-merous,
petals whitish.....................................................................................................................17
17. Leaf lateral veins 1012 pairs. Inflorescences hairy......................................5. M. griffithii
Leaf lateral veins 922 pairs. Inflorescences glabrous.............................12. M. quadrifida
1. Mangifera blommesteinii Kosterm.

(L. van Blommestein, a plant collector in Java and Sumatra, c. 1878)
In Kostermans & Bompard l.c. 146. Type: G. Mikil SAN 31986, Borneo, Sabah, Weston Distr. (holotype L; isotype SAN).
Tree to 45 m tall, 110 cm diameter. Bark reddish brown, scaly; inner bark laminated, reddish
brown. Sapwood pale white. Twigs deeply grooved when young. Leaves in pseudowhorls,
36
drying brownish, thick-coriaceous; oblanceolate, 4.58.5 x 1.53 cm; base attenuate, apex
rounded and slightly emarginate; lateral veins 912 pairs, faint below, more distinct above,
intercostal veins invisible below, faintly visible above; petioles 0.51.8 cm long.
Inflorescences (not fully developed) fascicled racemes, to 20 cm long. Flowers unknown.
Fruits (immature) ellipsoid to subglobose, c. 2 x 1.2 cm, smooth.
Distribution. Peninsular Malaysia, Borneo. Very uncommon in Sabah and Sarawak. In
Sabah, known from 3 collections, SAN 31986 (type) from Weston, SAN 39172 from
Sandakan and SAN 49111 from Semporna districts. In Sarawak, known from one collection,
KEP 80068, from Miri.
Ecology. Lowlands, including riverbanks and kerangas, to 120 m.
2. Mangifera caesia Jack
(Latin, caesium = lavender-blue; the flowers)
In Roxburgh, Fl. Ind. ed. Wall. 2 (1824) 441; King l.c. 476; Merrill l.c. (1921) 349; Ridley l.c. 525; Masamune l.c. 410;
Mukherji l.c. 126; Backer & Bakhuizen f. l.c. 15; Kostermans, Reinwardtia 7 (1965) 15; Ding Hou, Blumea 24 (1978) 24,
FM 1, 8 (1978) 438; Anderson l.c. 136; Corner l.c. 117; Kochummen l.c. 33; Bompard l.c. 207; Whitmore, Tantra & Sutisna
l.c. 16; Kostermans & Bompard l.c. 149. Type: Jack, s.n., Sumatra (n.v.).
Tree to 45 m high, 90 cm diameter. Bark brown, shallowly fissured. Twigs stout, 1 cm thick,
with sharp ridges. Leaves usually in pseudowhorls, drying chocolate-brown, thickcoriaceous; elliptic to obovate or oblanceolate, 7.518 x 2.56.5 cm; base attenuate, apex
acute or obtuse; lateral veins 1022 pairs, faintly visible on both surfaces, intercostal veins
reticulate, invisible; petioles 25 cm long, flattened above. Inflorescences pyramidal
panicles, to 30 cm long. Flowers violet or lilac, 5-merous, petals purplish inside, with 1 ridge
each; disc cylindrical and stalk-like, not papillose; stamens 5, one or two fertile, filaments
connate; ovary subglobose, style slightly eccentric. Fruits pale brown or brownish yellow,
ellipsoid or pyriform, 1219 x 610 cm; pulp white or yellowish white, sour to sweet.
Vernacular names. Sabahbeluno (Malay). Sarawakbinjai (Malay), lanyat (Iban).
Distribution. Sumatra, Peninsular Malaysia, Borneo. Widely cultivated in Sabah and
Sarawak.
Ecology. Lowland forest by streams and in swampy areas.
Uses. A well-known fruit tree. The sour fruits are often used as a substitute for tamarind.
3. Mangifera decandra Ding Hou
(Greek, deca =10, andros = male; the flower with 10 stamens)
Reinwardtia 8 (1972) 323, FM 1, 8 (1978) 433; Whitmore, Tantra & Sutisna l.c. 16; Kostermans & Bompard l.c. 142. Type:
Singh SAN 60874, Borneo, Sabah (holotype L; isotypes K, SAN).
37
Medium-sized tree to 30 m tall, 70 cm diameter. Bark reddish brown, fissured and scaly; inner
bark pinkish to orange-brown. Sapwood pale yellow. Twigs stout, to 1.3 cm thick, with sharp
ridges. Leaves in pseudowhorls, drying brownish, coriaceous; elliptic to oblong or obovate, 17
38 x 1215 cm; base cuneate, apex slightly acute; lateral veins 2035 pairs, raised on both
surfaces, intercostal veins invisible; petioles 26 cm long. Inflorescences terminal, pyramidal, to
65 cm long, hairy, with stout peduncles. Flowers 5-merous, reddish or pinkish; petals without
ridges; stamens 10, 5 fertile, filaments connate; disc cylindric, stalk-like; ovary subglobose, style
eccentric. Fruits ellipsoid, 916 x 810.5 cm, irregularly rugose, yellowish to rusty brown when
fresh.
Distribution. Sumatra, Borneo. Widely distributed in Sabah, uncommon in Sarawak; also in
Ecology. Lowland forest, including swamps, to 340 m.
4. Mangifera foetida Lour.

(Latin, foetidus = stinking, strong-smelling; the fruits)
Fl. Cochinch. (1790) 160; King l.c. 474; Ridley l.c. 524; Masamune l.c. 410; Burgess l.c. 20; Smythies l.c. 5; Ding Hou, FM
1, 8 (1978) 435; Anderson l.c. 136; Bompard l.c. 209; Corner l.c. 118; Kochummen l.c. 35; Whitmore, Tantra & Sutisna l.c.
17; Kostermans & Bompard l.c. 159; Kessler & Sidiyasa l.c. 42. Type: Rumph. Herb. Amboin. 1 (1741) t. 28.
Medium-sized tree to 20 m tall, 35 cm diameter. Bark grey-brown, fissured and scaly; inner bark
reddish with creamy to pinkish latex in droplets. Sapwood white. Leaves spirally arranged,
drying brownish, thick-coriaceous; elliptic, oblong or oblanceolate, 1228 x 610 cm; base
cuneate, apex obtuse or rounded; lateral veins 1220 pairs, with short intermediate veins, raised
on both surfaces, intercostal veins very faint to inconspicuous; petioles 23.5 cm long, the base
slightly swollen and rugose. Inflorescences pyramidal, to 30 cm long, peduncles brick-red.
Flowers fragrant, 5-merous; calyx red; petals whitish inside, each with 35 ridges; stamens 5,
fertile stamen one, filaments connate at base; disc cylindric, stalk-like; ovary subglobose. Fruits
subglobose, slightly oblique, to 14 x 10 cm, greenish grey, with strong turpentine odour; pulp
yellow, edible.
Vernacular names. Sabahpachu (Malay). Sarawakbachang (Iban), badut (Bidayuh),
embang (Malay), jabing (Bidayuh), kemantan (Iban), machang (Malay), pangin (Kayan), pawuk
(Bidayuh).
Distribution. Sumatra, Peninsular Malaysia, Java, Borneo. In Sabah and Sarawak, uncommon in
the forest, but often cultivated; also in Brunei and Kalimantan.
Uses. The fruits are edible and so the species is extensively cultivated.
5. Mangifera griffithii Hook. f.
(W. Griffith, 18101845, surgeon at Malacca)
38
Trans. Linn. Soc. 23 (1860) 168; King l.c. 468; Ridley l.c. 52; Mukherji l.c. 103; Ding Hou, Blumea 24 (1978) 25, FM 1, 8
(1978) 430; Anderson l.c. 136; Corner l.c. 118; Kochummen l.c. 35; Whitmore, Tantra & Sutisna l.c. 17; Kostermans &
Bompard l.c. 42. Type: Griffith 1100/1, Malacca (holotype K; isotypes CAL, L, P). Synonyms: M. sclerophylla Hook. f., Fl.
Br. Ind. 2 (1876) 15, Masamune l.c. 411; M. beccarii Ridl., Kew Bull. (1933) 194, Masamune l.c. 410.
Medium-sized tree to 30 m tall, 60 cm diameter. Bark greyish, smooth to cracking. Twigs

grey or dark brown, angular. Leaves spirally arranged, drying pale yellow or yellowish
green, not papillose beneath, thin-coriaceous; elliptic to obovate, 3.515.5 x (1.5)47 cm;
base cuneate, apex acute or obtuse, rarely finely emarginate; midrib not pinkish on drying
beneath, lateral veins 1012 pairs, visible on both surfaces, intercostal veins reticulate,
distinct on both surfaces; petioles 13.5 cm long, strongly swollen at base. Inflorescences
axillary or terminal, hairy. Flowers 4-merous; petals whitish, stamens 4, fertile stamen one,
filaments free; disc cup-shaped; style lateral. Fruits yellow to rose-red, ripening blackish,
ellipsoid, 2.53.5 x 1.52.5 cm; pulp pale orange-yellow.
Vernacular names. Sabahwahab (Malay). Sarawakasam raba (Malay).
Distribution. Sumatra, Peninsular Malaysia, Borneo. Uncommon in Sabah and Sarawak,
sometimes cultivated; also in Brunei and Kalimantan.
Ecology. Lowland forest to 360 m, very rarely to 750 m; often in swampy areas.
6. Mangifera indica L.
(of India)
Sp. Pl. (1753) 200; King l.c. 472; Merrill l.c. (1921) 349; Ridley l.c. 523; Masamune l.c. 411; Mukherji l.c. 83; Ding Hou,
FM 1, 8 (1978) 427; Anderson l.c. 136; Bompard l.c. 211; Corner l.c. 119; Kochummen l.c. 36; Whitmore, Tantra & Sutisna
l.c. 17; Kostermans & Bompard l.c. 88. Type: Drawing in Hermanns Herb. (BM). Synonyms: M. laurina Blume, Mus.
Bot. Lugd. Bat. 1 (1850) 195; M. longipes Griff., Notul. Pl. As. 4 (1854) 419, Masamune l.c. 411; M. applanata Kosterm. in
Kostermans & Bompard, The mangoes (1993) 64, syn. nov.; M. bompardii Kosterm. in Kostermans & Bompard l.c. 174,
syn. nov.; M. orophila Kosterm. in Kostermans & Bompard l.c. 138, syn. nov.; M. rubropetala Kosterm. in Kostermans &
Bompard l.c. 107, syn. nov.
Medium-sized tree to 35 m tall, 90 cm diameter. Bark greyish, shallowly fissured and scaly.
Leaves spirally arranged, drying greenish, not papillose beneath; elliptic, oblong or
lanceolate, 830 x 26 cm; base acute, margin entire, often wavy, apex acute; midrib not
pinkish on drying beneath, lateral veins 1230 pairs, visible on both surfaces, intercostal
veins reticulate, distinct below, faint to distinct above; petioles 1.52 cm long, base swollen.
Inflorescences pseudoterminal or axillary, pyramidal panicles, to 40 cm long, hairy or
glabrous. Flowers 5-merous; calyx hairy or glabrous outside; petals pale green to yellowish;
stamens 5, fertile stamen one, filaments free; disc cup-shaped; style lateral. Fruits very
variable in size and shape, subglobose to oblong, to 30 x 10 cm, ripening green to yellowish
to greenish red; pulp yellowish to orange.
Vernacular names. Sabahmempelam (Malay). Sarawakempelam (Malay).
39
Distribution. India, Myanmar, Thailand, Indo-China, Sumatra, Peninsular Malaysia, Java,

Borneo, New Guinea. Of scattered occurrence in Sabah and Sarawak but extensively
cultivated; also in Brunei and Kalimantan.
Ecology. Lowland to lower montane forest, to 1700 m; cultivated throughout the tropics and
subtropics.
Uses. Cultivated extensively for its edible fruits which can be eaten either raw, ripe or
processed into chutneys, pickles, dried slices, jam, etc. Mango juice is a popular drink.
Taxonomy. Ding Hou (l.c. 427) considered M. laurina as a synonym of M. indica while
keeping M. longipes as a separate species. Kostermans & Bompard (l.c. 120) treated M.
laurina as distinct species with M. longipes as its synonym. It is interesting to note that
Bompard (l.c. 216) recorded the vernacular names empelam, mempelam and manggaayer for M. laurina which are the same vernacular names given to M. indica. By studying
the descriptions given by Kostermans of his new species and the available collections, I have
come to the conclusion that M. applanata, M. bompardii, M. rubropetala, and M. orophila, as
well as M. laurina and M. longipes, are just variants of M. indica. Accordingly, these species
are reduced to synonymy under M. indica. With this reduction, the species occurs naturally
in the whole Indo-Malesian region and is not confined to parts of India and Myanmar as
stated by Kostermans & Bompard (l.c. 96).
7. Mangifera khoonmengiana Kochummen
Fig. 9.
(Wong Khoon Meng, forest botanist at Kepong, Brunei and Sandakan)

Sandakania 7 (1996) 83. Type: K.M. Wong 682, Brunei (holotype KEP; isotypes BRUN, K).
Medium-sized tree to 30 m tall, 40 cm diameter. Bark brownish, smooth to slightly fissured;

inner bark bright yellow with creamy latex which turns blackish on exposure. Sapwood pale
yellow. Twigs brownish or yellowish brown, glossy, slightly angular. Leaves spirally
arranged, drying brownish, coriaceous; elliptic or oblong, 1520 x 69.5 cm; base cuneate,
margin thickened, recurved, apex rounded with short blunt-tip; lateral veins 1315 pairs,
almost at right angles to midrib, faintly raised on both surfaces, intercostal veins finely
reticulate, visible on the lower surface, almost invisible above; petioles 1.52 cm long, base
swollen. Inflorescences terminal, racemose panicles, to 30 cm long, glabrous, with few side
branches; bracteoles lanceolate. Flowers pale green to creamy, 4-merous; pedicels very short,
to 1 mm long; sepals 4, ovate, fleshy; petals with circular brown gland-dots, membranous,
oblong with rounded apex and base, ridges 4 with a number of branches; disc distinct, cupshaped; stamens 4, fertile stamen one, filaments free; ovary globose, style slightly eccentric.
Fruits green when fresh, dark brown on drying, oblong, 2.53.5 x 2 cm, with rounded apex
and base, sessile.
Distribution. Endemic to Borneo. In Sarawak, very uncommon and known from a single
collection, S. 36600, from Miri; not yet reported from Sabah.
Ecology. Lowland forest. In Brunei growing gregariously in swampy areas.
40
Fig. 9. Mangifera khoonmengiana. A, flowering leafy twig; B, open flower viewed from above; C,
flower bud; D, flower bud with one sepal removed; E, fruit. (AD from Wong WKM 882, E from S.
36600.)
41
8. Mangifera macrocarpa Blume

(Greek, makros=large, karpos=fruit)
Bijdr. Fl. Ned. Ind. (1826) 1158; Mukherji l.c. 119; Backer & Bakhuizen f. l.c. 148; Burgess l.c. 20; Ding Hou, Blumea 24
(1978) 26, FM 1, 8 (1978) 439; Kochummen l.c. 37; Whitmore, Tantra & Sutisna l.c. 17; Kostermans & Bompard l.c. 157.
Type: Blume, s.n. (= L. sheet no. 897. 363552), Java, Mt. Seribu (holotype L; isotype BO).
Medium-sized to tall tree to 37 m tall, 70 cm diameter. Bark grey, fissured; inner bark
laminated, yellowish, with white sap. Sapwood yellowish. Leaves spirally arranged, drying
greenish, greenish yellow to pale yellow, not papillose below; linear, 1644 x24 cm; base
cuneate to attenuate, apex acute; midrib not pinkish on drying beneath, lateral veins 2533
pairs, with short intermediate veins, very faint on both surfaces, intercostal veins reticulate,
very faintly visible on both surfaces; petioles 610 cm long. Inflorescences terminal
panicles, pyramidal, to 20 cm long, glabrous. Flowers (only male seen) 5-merous; sepals
ovate-lanceolate; petals white, linear-oblong, each with 13 ridges; stamens 5, 1 fertile,
filaments free; disc obscure. Fruits obliquely, broadly oblong-globose, 812 cm long, with
yellow pulp.
Distribution. Thailand, Sumatra, Peninsular Malaysia, Java, Borneo. Very uncommon in
Sabah, known only from few collections from Beaufort, Kalabakan, Lahad Datu, Tawau,
Tenom and Sandakan districts; not yet reported from Sarawak; also in Brunei and
Kalimantan.
Ecology. Lowland forest to 150 m on hilly areas.
9. Mangifera magnifica Kochummen
(Latin, magnus = large; the tree)
Gard. Bull. Sing. 36 (1983) 189, l.c. (1989) 37; Kostermans & Bompard l.c. 75. Type: Conservator of Forests, Kedah, KEP
20562, Peninsular Malaysia, Kedah, G. Jerai (holotype KEP; isotype SING).
Tree to 40 m tall, 130 cm diameter. Bark grey-brown, smooth to fissured or scaly; inner bark
yellowish brown with white latex which turns to yellow and then black on exposure.
Sapwood pale yellow. Twigs stout, to 8 mm thick, angular. Leaves spirally arranged,
sometimes in pseudowhorls at the ends of twigs, drying brownish, very thick-coriaceous
(cardboard-like); elliptic to oblong, 1317.5 x 59 cm; base broadly cuneate, apex obtuse,
rounded or acute; lateral veins 1517 pairs, prominently raised below, faint above, intercostal
veins reticulate, visible on both surfaces; petioles stout, 23 cm long, basal part swollen and
rugose. Inflorescences terminal and axillary panicles. Flowers fragrant, 4-merous; sepals
yellowish white or with red tinge when fresh; petals white, each with 34 yellow ridges;
fertile stamen one, filaments free; disc pulvinate, obscurely lobed; style lateral. Fruits ovoid
or oblong, 912 x 810 cm, grey-green, often with brown spots; pulp whitish, with a sweet to
acidic taste.
Distribution. Sumatra, Peninsular Malaysia, Borneo. Widely distributed but uncommon in
Sabah and Sarawak.
Ecology. Lowland and hill forests, to 500 m.
42
10. Mangifera pajang Kosterm.

(from the local Kalimantan nameasem pajang)
Reinwardtia 7 (1965) 20; Ding Hou, Blumea 24 (1978) 27, FM 1, 8 (1978) 436; Anderson l.c. 136; Whitmore, Tantra &
Sutisna l.c. 18; Bompard l.c. 209; Kostermans & Bompard l.c. 147. Type: Kostermans 12534, Borneo, Kalimantan (holotype
BO; isotypes K, L, SAN, SING).
Medium-sized tree to 30 m tall, 70 cm diameter. Bark dark brown, smooth to shallowly

fissured; inner bark brownish with creamy latex. Twigs stout, 12 cm thick, angular. Leaves
usually spirally arranged, sometimes in pseudowhorls, drying brownish, thick-coriaceous;
elliptic to oblong or obovate, 1528 x 5.512.5 cm; base cuneate, apex obtuse; midrib usually
finely grooved on the under surface, lateral veins 1325 pairs, raised on both surfaces,
intercostal veins invisible on both surfaces; petioles 3.59 cm long, strongly swollen at base,
upper surface flattened and concave at base, irregularly rugose below. Inflorescences
terminal or appearing in the axils of upper leaves, to 30 cm long. Flowers 5-merous; calyx
glabrous, dark purple; petals purplish inside, pinkish white outside, each with 3 ridges;
stamens 5, fertile stamens 12, filaments free; disc stalk-like; ovary whitish, style eccentric.
Fruits rugose, brownish, globose to ellipsoid, 1012 x 8.510 cm; epicarp thick, and can be
peeled off like a banana skin; pulp deep orange-yellow, sweet.
Vernacular names. Sabahbambangan, embawang (Malay). Sarawakembang (Malay),
bawang, mawang (Bidayuh).
Distribution. Endemic to Borneo. In Sabah and Sarawak, uncommon in the forest, often
cultivated for its fruit; also in Brunei and Kalimantan.
Ecology. Lowland forest to 500 m, often in swampy and riverine habitats.
Uses. The fruit is edible and is in great demand.
Notes. This species is very similar to M. foetida in leaf characters. However, the stouter
twigs, generally larger leaves and longer rugose petioles distinguish it from M. foetida.
11. Mangifera parvifolia Boerl. & Koord.
(Latin, parvus = small, folium = leaf)
In Koorders-Schumacher, Syst. Verz. 2 (1910) 31; Mukherji l.c. 130; Ding Hou, Blumea 24 (1978) 28, FM 1, 8 (1978) 431;
Kostermans & Bompard l.c. 39. Type: Koorders 21218, Sumatra, Sg. Buwatan (BO). Synonym: Mangifera havilandii Ridl.,
Kew Bull. (1933) 194, Masamune l.c. 411, Anderson l.c. 36.
Medium-sized tree to 30 m tall, 40 cm diameter. Bark reddish brown, shallowly fissured.

Twigs reddish brown, smooth; axillary buds globose, persistent. Leaves spirally arranged,
well-spaced, coriaceous, drying greenish, not papillose below; elliptic to narrowly obovate or
oblong 8.512.5 x 4.56.5 cm; base cuneate, margin not thickened or recurved, apex
acuminate, acumen c. 1 cm long; midrib not pinkish on drying beneath, lateral veins 810
pairs, faint on both surfaces, intermediate veins absent or invisible, intercostal veins
43
reticulate, very faintly visible below, invisible above; petioles 1.54.5 cm long, swollen at
base. Inflorescences axillary or pseudoterminal panicles. Flowers 4-merous; petals whitish;
stamens 4, fertile stamen one, filaments free; disc cup-shaped; style lateral. Fruits cylindrical
or ellipsoid, smooth, blue-black, to 6 x 3 cm, with orange pulp.
Vernacular name. Sarawakraba (Malay).
Distribution. Sumatra, Peninsular Malaysia, Borneo. In Sabah and Sarawak, uncommon,
mainly in mixed swamp forests to 300 m.
12. Mangifera quadrifida Jack

(Latin, quadrifidus = 4-cleft; the 4-merous flowers)
In Roxburgh, Fl. Ind. ed. Wall. 2 (1824) 440; King l.c. 471; Merrill l.c. (1921) 349; Ridley l.c. 522; Mukherji l.c. 112; Ding
Hou, Blumea 24 (1978) 28, FM 1, 8 (1978) 429; Anderson l.c. 137; Corner l.c. 120; Kochummen l.c. 38; Whitmore, Tantra
& Sutisna l.c. 18; Kostermans & Bompard l.c. 68. Type: Jack, Herb. Wallich Cat. 8489, Penang (K).
Large tree to 42 m tall. Twigs reddish brown, angular. Leaves spirally arranged, thin-coriaceous,
drying pale yellow or yellowish green, not papillose below; elliptic to oblong to narrowly
obovate, 9.530 x 47 cm; base cuneate, apex acute, or obtuse to rounded; midrib not pinkish on
drying beneath, lateral veins 922 pairs, faintly raised on both surfaces, intercostal veins
reticulate, visible on both side; petioles 1.55.5 cm long, swollen at base. Inflorescences terminal
and axillary panicles, glabrous. Flowers 4-merous; petals whitish, each with 3 ridges; stamens 4,
fertile stamen one, filaments free; disc pulvinate; style eccentric. Fruits ellipsoid or globose, 68
x 56 cm, green, ripening black; pulp orange-yellow.
Vernacular name. Sarawakasam kumbang (Malay).
Distribution. Sumatra, Peninsular Malaysia, Borneo, Sulawesi, the Lesser Sunda Islands and
Maluku; extremely uncommon in Sabah and Sarawak. In Sabah, known from Kudat, Keningau
and Tenom districts, while the only record from Sarawak is from Kuching area; also in Brunei
and Kalimantan.
Ecology. Lowland forest at very low altitude.
13. Mangifera rufocostata Kosterm.

(Latin, rufus = reddish; costa = midrib; the reddish midrib of the leaves)
In Kostermans & Bompard l.c. 116. Type: Kostermans 9799, Borneo, Kalimantan (holotype BO; isotypes BM, L).
Medium-sized to very tall tree to 45 m tall, 120 cm diameter; bole with buttresses to 2.5 m high.
Bark grey, smooth to fissured and scaly; inner bark yellow. Sapwood pale green. Twigs slender,
dark brown. Leaves spiraly arranged, coriaceous, drying greenish, greenish yellow or pale
yellow, papillose and with scattered black dots beneath; ovate to oblong, 817 x 2.56 cm; base
cuneate, apex obtuse or acute; midrib pinkish brown on drying beneath, lateral veins
44
1018 pairs, very faint below, visible above, intercostal veins reticulate, very faintly visible
on both surfaces; petioles 12 cm long. Inflorescences terminal or subterminal racemes, to
15 cm long, glabrous, pedicels articulate. Flowers pale yellow, 5-merous; petals oblong, 45
mm long; stamens 5, fertile stamen one, filaments free; disc distinct, pulvinate. Fruits
obliquely globose, 8.510 cm, brownish green with numerous brown lenticels; pulp whitish,
strongly fibrous.
Distribution. Sumatra, Peninsular Malaysia, Borneo. Widely distributed in Sabah, not yet
reported from sarawak.
Ecology. Lowland to lower montane forest, to 1500 m.
14. Mangifera subsessilifolia Kosterm.

(Latin, subsessilis = almost without stalk, folium = leaf)
In Kostermans & Bompard l.c. 180. Type: Kostermans, s.n., Singapore, Bt. Timah (holotype L; isotype BO).
Medium-sized tree to 25 m tall. Twigs brown, strongly ridged. Leaves in close spirals,
appearing as in pseudowhorls, drying brownish, thick-coriaceous; narrowly lanceolate, 2545
x2.33 cm; base attenuate, margin slightly curled inwards, apex acute with blunt tip; lateral
veins more than 40 pairs, curving near margin, very faint below, raised above, intercostal
veins reticulate, distinct above, inconspicuous below; petioles stout, to 1 cm long.
Inflorescences, flowers and fruits unknown.
Distribution. Sumatra, Peninsular Malaysia, Borneo. Extremely uncommon in Sabah, known
from a single collection (SAN 16792) from Sipitang district, not yet reported from Sarawak.
15. Mangifera swintonioides Kosterm.
(resembling the genus Swintonia)
In Kostermans & Bompard l.c. 80. Type: G.H.S. Wood SAN 16489, Sabah, Tawau (holotype BO; isotypes KEP, SAN,
SING).
Tree to 40 m tall, 70 cm diameter. Bark grey, cracking and scaly; inner bark yellowish brown.
Sapwood white. Twigs pale brown, 4 mm thick, slightly angular. Leaves spirally arranged, more
or less aggregated towards the ends of twigs, thin-coriaceous, drying greenish, not papillose
below; oblong or elliptic, 715 x 2.54 cm; base cuneate, apex acute; midrib not pinkish on
drying beneath, lateral veins 1323 pairs, slightly raised below, very faint above, intercostal veins
reticulate, very faintly visible below, almost invisible above; petioles 47 cm long, strongly
swollen at base, swollen part whitish. Inflorescences pseudoterminal panicles, glabrous. Flowers
5-merous; sepals thin with pale yellow margins; petals whitish, each with 46 ridges; fertile
stamen one, filaments free; disc pulvinate; style eccentric. Fruits oblong, to 11.5 x 10 cm, pale
grey-brown, pulp whitish.
45
Distribution. Sumatra, Peninsular Malaysia, Borneo. Uncommon in Sabah, recorded from Kudat,
Lahad Datu, Sandakan, Sipitang and Tawau districts; not yet reported from Sarawak.
16. Mangifera torquenda Kosterm.

(Latin, torquendus = cut across and twisted; the manner by which ripe fruit is opened for consumption)
Reinwardtia 7 (1965) 21, in Kostermans & Bompard l.c. 60. Type: Kostermans 12533, Borneo, Kalimantan (holotype BO;
isotypes A, BM, K, KEP, L, P, SING).
Tree to 36 m high, 60 cm diameter. Bark greyish, shallowly fissured to scaly; inner bark orange with
white latex. Sapwood pale. Twigs dark brown, slightly angular. Leaves spirally arranged, thickcoriaceous, drying greenish, not papillose below; oblong or elliptic, 12.523(32) x 48(11); base
broadly cuneate to rounded, margin not thickened but recurved, apex acute; midrib not pinkish on
drying beneath, lateral veins 1420 pairs, raised on both surfaces, intercostal veins finely reticulate,
distinctly visible below, faint above; petioles 27 cm long, strongly swollen at base. Inflorescences
pseudoterminal, sparsely hairy at the basal part. Flowers 4-merous, sweet scented, pale yellow; each
petal with 6 ridges, the central ridges with 3 upright branches ending in tiny globose glands; stamens
4, fertile stamen one, filaments free; disc pulvinate, 4-lobed; style eccentric. Fruits globose, c. 7.5
x 6.5 cm, yellowish green with darker dots when fresh; pulp pale yellow, with turpentine smell, with
short fibre, tasting sweet to acidic. Seeds surface grooved.
Vernacular names. Sabahbuniton, bunyitan (Murut). Sarawakkemantan, lemantan, mantan,
rade (Iban).
Distribution. Sumatra, Peninsular Malaysia, Borneo. Uncommon in Sabah and Sarawak. In Sabah,
recorded from Kalabakan, Lahad Datu, Sandakan, Tawau and Tenom areas; in Sarawak, collected
from Lundu, Serian, and Bintulu areas.
Ecology. Lowland and hill forests, to 800 m; also cultivated; also in Kalimantan.
Uses. The fruit is edible and is peeled by making a cross-section and twisting the two halves in
opposite directions.
Taxonomy. Kessler & Sidiyasa (1994), following Ding Hous treatment (1978), considered M.
torquenda a synonym of M. similis Blume. Kostermans & Bompard (1993), however, have given
sufficient reasons to treat these two species as distinct, an opinion accepted in the present account. M.
similis (which does not occur in Sabah and Sarawak) differs from M. torquenda by its conspicuously
raised, reticulate intercostal veins on the upper leaf surface, shorter petioles, clustered flowers in
subracemose, minutely hairy panicles, minutely hairy flowers, strongly prominent petals ridges
topped by larger glands, a compact disc which is not much broader than the ovary, and the seeds
without grooves.
46
10. MELANOCHYLA Hook. f.

(Greek, melanos = black, chulos = sap; the black exudate from bruised parts)
Fl. Br. Ind. 2 (1876) 38; King, J. As. Soc. Beng. 65, 3 (1896) 502; Merrill, EB (1921) 351; Ridley, FMP 1 (1922) 541;
Masamune, EPB (1942) 411; Smythies, CST (1965) 10; Backer & Bakhuizen f., FJ 2 (1965) 154; Burgess, TBS (1966) 25;
Ding Hou, Blumea 24 (1978) 29, FM 1, 8 (1978) 490; Anderson, CLTS (1980) 137; Wong, DMT (1982) 136; Corner,
WSTM 3rd ed. 1 (1988) 128; Kochummen, TFM 4 (1989) 40; Whitmore, Tantra & Sutisna, CLK 1 (1989) 18; Kessler &
Sidiyasa, Trees of the Balikpapan-Samarinda Area (1994) 43.
Small to medium-sized trees; bole with short buttresses or stilt-roots (in swampy habitats). Bark
reddish brown, smooth to cracking; inner bark reddish brown, granular, often with droplets of black
sap. Sapwood whitish. Leaves spirally arranged, simple, usually papillose beneath. Inflorescences
axillary or terminal panicles, rarely in fascicles, with bracts and bracteoles; pedicels articulate.
Flowers unisexual (plants dioecious), 45-merous; hypanthium (receptacle) cup-shaped, hairy
outside, slightly accrescent in fruit; calyx 5- or 4-lobed; petals 5 or 4, imbricate (sometimes only
slightly), hairy outside, densely woolly inside; stamens 5 or 4, filaments free or lower part laterally
adnate to the petals, densely hairy, anthers dorsifixed, imperfect or aborted in female flowers; disc
intrastaminal, rim-like, 5- or 4-lobed; ovary superior, rarely semi-inferior or inferior, 1-locular,
usually densely hairy, style distinct, stigmas 3; ovary in male flowers very small or abortive. Fruits
hairy outside, mesocarp and endocarp full of black resin. Seeds with testa fused to the endocarp;
embryo straight, cotyledons free.
Distribution. Thirty species; Thailand, Sumatra, Peninsular Malaysia, Java, Borneo; 17 species in
Sabah and Sarawak.
Ecology. Mainly in the lowland, including swamp, sandstone and limestone forests, rarely in lower
montane forest to 1350 m.
Uses. The timber is medium hardwood, grain interlocking and slightly wavy, sapwood sharply
differentiated from the heartwood which is dark brown with black streaks. In Sabah and Sarawak, the
timber has no commercial importance at present.
Key to Melanochyla species

1. Leaves glabrous...................................................................................................................2
Leaves hairy, at least on the midrib below.........................................................................11
2. Leaves in close clusters, subsessile; leaf base auricled...............................2. M. auriculata
Leaves well-spaced, with well-developed petioles; base not auricled.................................3
3. Leaves distinctly glaucous below. Inflorescences axillary panicles to 3 cm long.................
........................................................................................3. M. axillaris
Leaves not or only very faintly glaucous below. Inflorescences usually terminal or
appearing from the axils of upper leaves, very much longer...............................................4
47
4. Twigs stout to 1 cm thick. Leaves thick-coriaceous; petioles thick and stout, 34 cm long.
Large tree of lower montane forest, at 13701800 m..................................13. M. montana
Not this combination of characters......................................................................................5
5. Leaf intercostal veins distinctly scalariform.........................................14. M. scalarinervis
Leaf intercostal veins reticulate or scalariform-reticulate...................................................6
6. Leaf base usually almost subcordate or obtuse; papillae in distinct groups on the lower
leaf surface. Fruits with spiny processes...................................11. M. fulvinervis (in part)
Leaf base not so; papillae either obscure or uniformly distributed on lower leaf surface.
Fruits without spiny processes.............................................................................................7
7. Lower half of petioles swollen and whitish and often irregularly rugose on drying. Leaves
not shiny above....................................................................................................................8
Lower half of petioles not swollen. Leaves shiny above......................7. M. caesia (in part)
8. Twigs distinctly whitish. Leaves distinctly papillose beneath.............................................9
Twigs grey-brown. Leaves not or only faintly obcurely papillose beneath...........................
.................................................................................................................1. M. angustifolia
9. Leaf base obtuse; lateral veins sunken above, intercostal veins scalariformreticulate....................................................................................................17. M. woodiana
Leaf base cuneate; lateral veins not sunken above; intercostal veins reticulate.................10
10. Leaves drying reddish brown, faintly glaucous and distinctly papillose beneath..................
.........................................................................................................15. M. semecarpioides
Leaves drying yellowish green, not glaucous and faintly papillose beneath.........................
................................................................................................................8. M. castaneifolia
11. Leaves sparsely hairy below..............................................................................................12
Leaves densely hairy below...............................................................................................16
12. Leaves not or only faintly papillose beneath; leaf base cuneate.............12. M. minutiflora
Leaves distinctly papillose beneath; leaf base subcordate, obtuse or
rounded, rarely cuneate...................................................................................................13
13. Leaf base subcordate or obtuse; papillae on the lower leaf surface in groups. Fruit with
spiny processes...........................................................................11. M. fulvinervis (in part)
Leaf base cuneate or rounded; papillae on the lower leaf surface uniformly distributed or
in groups. Fruit not so........................................................................................................14
14. Papillae in distinct groups on the lower leaf surface..................................5. M. borneensis
Papillae uniformly distributed on the lower leaf surface...................................................15
48
15. Leaves shiny above, glaucous below; intercostal veins scalariform-reticulate......................

.............................................................................................................7. M. caesia (in part)
Leaves not shiny above, not glaucous below; intercostal veins scalariform..........................
........................................................................................................................10. M. elmeri
16. Leaves bullate above; intercostal veins scalariform, distinctly sunken on the upper leaf
surface..............................................................................................................6. M. bullata
Leaves not bullate above; intercostal veins scalariform-reticulate or reticulate, usually not
sunken on the upper leaf surface........................................................................................17
17. Leaves slightly glaucous below..................................................................4. M.beccariana
Leaves not glaucous below................................................................................................18
18. Leaves more than 50 cm long; lateral veins 4044 pairs, sunken on the upper leaf surface.
Inflorescences in axillary clusters, 67 cm long.......................................9. M. condensata
Leaves smaller; lateral veins fewer, very faint on the upper leaf surface. Inflorescences
terminal or axillary, to 30 cm long............................................................16. M. tomentosa
1. Melanochyla angustifolia Hook. f.
Fl. Br. Ind. 2 (1876) 39; King l.c. 506; Ridley l.c. 541; Ding Hou, FM 1, 8 (1978) 496; Kochummen l.c. 41; Whitmore,
Tantra & Sutisna l.c. 18. Type: Maingay 492, Malacca (holotype K; isotype L).
Small to medium-sized tree to 30 m tall, 45 cm diameter; bole with buttresses to 2.5 m high. Bark
grey-brown to reddish, smooth to cracking; inner bark brownish with white latex, soon becoming
black. Sapwood whitish. Twigs grey-brown, strongly grooved, glabrous or sparsely hairy. Leaves
well-spaced, thin-coriaceous; not glaucous below, not shiny above, glabrous; papillae obscure;
elliptic to narrowly obovate, 1527.5 x 3.59 cm; base attenuate and not auricled, apex acuminate,
acumen to 1.5 cm long; midrib raised above with median groove, lateral veins 1224 pairs, curving
and joining near margin, raised below, faint above, intercostal veins scalariform-reticulate, visible
below, faint to inconspicuous above; petioles 24.5 cm long, often swollen at the lower half, strongly
rugose on drying, glabrous or sparsely hairy. Inflorescences terminal hairy panicles, to 30 cm long.
Flowers yellowish, subsessile; calyx lobes triangular; petals oblong, elliptic, lanceolate, or
oblanceolate, flat; stamen filaments free; staminodes small; disc 5- or 4-lobed; ovary globose, stigmas
minute, capitate; pistillodes replaced by tufts of hairs. Fruits ellipsoid, 22.5 x 1.5 cm, ripening
yellowish, apex rounded, surface smooth, sparsely hairy.
Distribution. Peninsular Malaysia, Borneo. Uncommon in Sabah and Sarawak. Recorded from
Beluran, Kota Merudu, Lamag and Sandakan areas in Sabah, and from Anap, Gunung Gading, Kapit
and Tubau areas in Sarawak; also in Kalimantan.
Ecology. Lowland to lower montane forest to 1100 m; on ridges and hillsides. Flowering in April to
November and fruiting in January.
2. Melanochyla auriculata Hook.f.
(Latin, auriculatus = furnished with ear-like appendage; the leaf base)
l.c. 39; King l.c. 505; Ridley l.c. 540; Ding Hou, FM 1, 8 (1978) 495; Corner l.c. 128; Kochummen l.c. 41; Whitmore,
Tantra & Sutisna l.c. 18; Kessler & Sidiyasa l.c. 44. Type: Maingay 1897, Malacca (holotype K).
49
Tree to 30 m tall, 70 cm diameter; bole with steep buttresses to 2 m tall. Bark grey-green, smooth
with horizontal rings; inner bark reddish brown with black latex. Sapwood whitish. Twigs stout, 12
cm thick, strongly grooved, young parts densely covered with reddish brown short hairs. Leaves in
close clusters, glabrous; narrowly obovate, 2046 x 810 cm; base attenuate, auricled, apex acute;
midrib raised above, lateral veins 2025 pairs, raised on both surfaces, intercostal veins scalariformreticulate, visible below, invisible above; petioles absent or stout and to 5 mm long. Inflorescences
terminal panicles, to 6 cm long, axes strongly angular. Flowers white, sessile; calyx lobes ovateoblong; petals oblong, elliptic or lanceolate, slightly longitudinally ridged inside; stamen filaments
free; disc 5- or 4-lobed; ovary subglobose, stigma minute, capitate; pistillodes replaced by tuffs of
hairs. Fruits rusty hairy, rugose, 23.5 x 22.5 cm.
Distribution. Peninsular Malaysia, Borneo. Of scattered occurrence in Sabah, not yet reported from
Sarawak.
Ecology. Lowland, including riverbank, to hill forest, to 400 m. Flowering in May and September,
fruiting in May and June.
3. Melanochyla axillaris Ridl.

(Latin, axillaris = growing in an axil; the flowers)
Kew Bull. (1933) 198; Masamune l.c. 411; Ding Hou, Blumea 24 (1978) 29, FM 1, 8 (1978) 495; Anderson l.c. 137.
Lectotype (Ding Hou, 1978): Haviland 2211, Borneo, Sarawak (K; isolectotypes BM, L, SING).
Medium-sized tree to 24 m tall, 20 cm diameter. Twigs stout, 1.5 cm thick, grey-brown. Bud scales
lanceolate, velvety brown-hairy. Leaves well-spaced; distinctly glaucous below, glabrous on both
surfaces; papillae distinct on the lower leaf surface; oblanceolate, 1867 x 416.5 cm; base attenuate,
not auricled, apex obtuse or acute; midrib flattened above, lateral veins 2137 pairs, intercostal veins
scalariform-reticulate, visible below, faint to invisible above; petioles 38 cm long, lower half
swollen, rusty velvety hairy and strongly rugose on drying. Inflorescences axillary panicles, 13 cm
long; bracts lanceolate, 1.52 mm long. Flowers: hypanthium 34 mm long; calyx lobes triangular or
ovate, c. 2 mm long; petals oblong or obovate-oblong, 45 x 1.21.7 mm; lower parts of stamen
filaments united with the petals; disc 5-lobed; ovary subglobose, rusty hairy, style short, stigmas
minute, capitate. Fruits broadly ellipsoid or ovoid, c. 3 x2 cm, rusty hairy, apex acute, surface
smooth.
Distribution. Endemic to Borneo. Reported from Sarawak only with few collections from Kuching
and Lundu areas.
Ecology. Lowland forest to 450 m. Flowering in April and November and fruiting in January and
April.
4. Melanochyla beccariana Oliv.
Fig. 10AB.

In Hooker, Icon. Pl. 24 (1894) t. 2313; Merrill l.c. 351; Masamune l.c. 411; Ding Hou, FM 1, 8 (1978)
50
Fig. 10. Melanochyla beccariana (AB); M. borneensis (CE); M. fulvinervis (F). A, inflorescence; B, flower in longitudinal
section; C, leafy twig; D, detail of lower leaf surface; EF fruits. (AB from SAN 21255, CE from S. 37898, F from FRI
10599.)
51
497; Anderson l.c. 137; Whitmore, Tantra & Sutisna l.c. 18. Syntypes: Beccari PB 2546, Borneo, Sarawak (K); Haviland
814, Borneo, Sarawak (K).
Medium-sized tree to 24 m tall, 35 cm diameter. Twigs densely reddish brown hairy. Leaves wellspaced; velvety reddish brown hairy below and on midrib above, slightly glaucous below; papillae
uniformly distributed below; oblong or narrowly obovate, (10)2029 x (3)8.512 cm; base cuneate,
apex rounded or with very short tip; midrib raised above, lateral veins 1731 pairs, distinctly raised
below, with 12 short intermediate veins in between each pair, faint and rarely sunken above,
intercostal veins scalariform-reticulate, distinct below, faint above; petioles 0.52.5 cm long, densely
reddish brown hairy, rugose on drying. Inflorescences terminal and axillary panicles, to 33 cm long,
densely short-hairy; bracts linear, to 1.5 cm long. Flowers subsessile; calyx lobes triangular or ovate
oblong; petals ovate-oblong or lanceolate; stamen filaments united laterally with the petals; disc 5lobed; ovary deeply or completely concealed by the receptable; stigma minute, capitate. Fruits
ellipsoid or ovoid, 2.54.5 x 1.22.2 cm, with blunt or slightly acute apex, rusty velvety hairy, surface
smooth.
Distribution. Endemic to Borneo. Uncommon in Sabah and Sarawak; also in Brunei and Kalimantan.
Ecology. Lowland to lower montane forest, to 1500 m. Flowering and fruiting in April and July.
5. Melanochyla borneensis (Ridl.) Ding Hou
Fig. 10CE.
(of Borneo)
Blumea 24 (1978) 31, FM 1, 8 (1978) 497; Anderson l.c. 137; Whitmore, Tantra & Sutisna l.c. 18. Basionym: Nothopegia
borneensis Ridl., Kew Bull. (1933) 197. Type: Beccari PB 241, Borneo, Sarawak (holotype K; isotypes BO, L, S).
Medium-sized tree to 25 m tall, 30 cm diameter. Bark dark grey, smooth. Leaves well-spaced, thickcoriaceous; glabrous above except the sparsely hairy midrib, sparsely hairy below; papillae in
prominent horseshoe-shaped groups on the lower leaf surface; elliptic, oblong or obovate, 928.5
x 5.512 cm; base cuneate to broadly rounded, not auricled, apex cuspidate; midrib prominently
raised above, channelled below, lateral veins 1824 pairs, sunken above, raised below, intercostal
veins reticulate, distinct below, obscure above; petioles stout, rugose on drying, 13.5 cm long, softly
hairy, glabrescent. Inflorescences axillary, to 15 cm long. Flowers sessile, yellowish; calyx lobes
triangular; petals ovate, flat; stamen filaments free; disc 45-lobed; ovary globose, stigma discoid;
pistillodes minute, hairy. Fruits ellipsoid to subglobose, 34 x 22.5 cm, with rounded or pointed
apex, surface soft-hairy, smooth.
Distribution. Endemic to Borneo; so far reported only from the Kuching area (including the
Semengoh Arboretum) in Sarawak.
Ecology. Lowland forest, to 100 m.
6. Melanochyla bullata Ding Hou

(Latin, bullatus = inflated; the protruding leaf upper surface between veins)
52
Blumea 24 (1978) 31, FM 1, 8 (1978) 496; Whitmore, Tantra & Sutisna l.c. 19; Kessler & Sidiyasa l.c.
44. Type: Anderson et al. S. 15436, Borneo, Sarawak, Lundu (holotype L; isotypes BO, SAR, SING).
Small to medium-sized tree to 30 m tall, 80 cm diameter. Bark brown, smooth to scaly. Twigs
greyish. Leaves well-spaced, densely hairy below, bullate above; oblanceolate to narrowly obovate,
13.542 x 59 cm; base cuneate, not auricled, apex rounded with very short tip; midrib flattened
above, lateral veins 3338 pairs, sunken above, raised below, intercostal veins scalariform, sunken
above, raised below; petioles whitish, 23 cm long, lower half flattened. Inflorescences terminal rusty
hairy panicles. Flowers (only male seen) yellow, sessile; calyx lobes triangular; petals ovate or ovateoblong, thickened at the lower two-thirds inside; stamen filaments free; disc obscurely 5-lobed;
pistillodes minute, glabrous. Fruits ovoid, 3.54 x 24.5 cm, velvety hairy, apex pointed, surface
smooth.
Distribution. Endemic to Borneo. Uncommon in Sabah and Sarawak. In Sabah, known from
Sandakan and Tawau districts and in Sarawak, from Lundu area; also in Kalimantan.
7. Melanochyla caesia (Blume ) Ding Hou

(Latin, caesius = lavender-blue; the leaf lower surface)
Blumea 24 (1978) 31, FM 1, 8 (1978) 498; Kochummen l.c. 42; Whitmore, Tantra & Sutisna l.c. 19. Basionym: Semecarpus
caesia Blume, Mus. Bot. Lugd. Bat. 1 (1850) 189. Lectotype (Ding Hou, 1978): Blume, s.n. (= L. sheet no. 897. 3631266),
Java (L; isolectotype BO). Synonym: Semecarpus heterophylla var. caesia (Blume) Engl. in A. DC., Mon. Phan. 4 (1883)
487.
Medium-sized tree to 27 m tall, 30 cm diameter. Bark grey-brown, smooth to scaly; inner bark
pinkish. Sapwood white. Twigs grey-brown, glabrous or rusty brown short-hairy. Leaves wellspaced, thin-coriaceous; glabrous on both surfaces, rarely sparsely hairy beneath, shiny above, faintly
glaucous and with uniformly distributed papillae beneath; oblong or lanceolate, 1020 x 2.54.5 cm;
base cuneate to broadly rounded, not auricled, apex acute; midrib raised above, lateral veins 1822
pairs, looping near margin, raised below, faint above, intercostal veins scalariform-reticulate, very
faint on both surfaces; petioles 0.52 cm long, whitish, rugose and hairy, lower half not swollen.
Inflorescences terminal, 1013 cm long, rusty hairy panicles, with fewer branches in females.
Flowers subsessile; calyx lobes triangular; petals white or yellow, oblong, elliptic or ovate-oblong,
with longitudinal ridges inside; stamen filaments free; staminodes very small; disc 5-lobed; ovary
globose, stigma minute, capitate; pistillodes conical, hairy. Fruits oblong or ellipsoid, 35 x 2 cm,
smooth, densely or sparsely rusty hairy, apex emarginate.
Distribution. Sumatra, Peninsular Malaysia, Borneo. Widely distributed in Sabah; uncommon in
Sarawak, recorded from Anap and Lawas areas only; also in Kalimantan.
Ecology. Lowland and hill forests, including kerangas, to 450 m, often by streams.
8. Melanochyla castaneifolia Ding Hou

(Latin, castaneus = chestnut-coloured, folium = leaf)
53
Blumea 24 (1978) 32, FM 1, 8 (1978) 496; Whitmore, Tantra & Sutisna l.c. 19. Type: Bunang S. 23016,
Borneo, Sarawak (holotype L; isotypes KEP, SAN, SAR).
Small to medium-sized tree to 24 m tall, 30 cm diameter. Twigs grey-white, glabrous. Leaves wellspaced, thin-coriaceous; glabrous, drying to yellowish green, upper surface not shiny; papillae
uniformly distributed on the lower leaf surface, very faint to obscure; elliptic, obovate or oblong, 614
x 25 cm; base cuneate, not auricled, apex acute; midrib raised above, flattened below, lateral veins
615 pairs, faintly raised on both surfaces, looping near margin, intercostal veins reticulate, distinct
below, invisible above; petioles 12.5 cm long, lower half thickened and whitish on drying.
Inflorescences terminal and axillary panicles, 622 cm long, finely hairy. Flowers yellow; calyx
lobes triangular; petals elliptic, oblong or ovate, longitudinally thickened in the central part inside;
stamen filaments free; disc 5-lobed; ovary globose, stigma minute, capitate; pistillodes very small,
glabrous. Fruits unknown.
Distribution. Endemic to Borneo. Uncommon in Sabah and Sarawak; recorded from Lamag and
Sandakan districts in Sabah, and from Kuching and 4th Division in Sarawak; also in Kalimantan.
9. Melanochyla condensata Kochummen

(Latin condensatus = condensed; the inflorescences)
Sandakania 7 (1996) 83. Type: Ashton S. 18323, Sarawak, Ulu Sinrok, Similajau Forest Reserve (holotype KEP; isotypes A,
BO, FHO, K, L, MEL, P, SAN, SING, SAR).
Small tree to 12 m tall, 8 cm diameter. Bark grey-brown, smooth; inner bark pinkish with abundant
black sap. Twigs stout, 11.5 cm thick, yellowish brown appressed hairy. Leaves coriaceous; densely
soft-hairy and not glaucous below, upper surface not bullate; papillae uniformly distributed on the
lower leaf surface; oblanceolate, 5355 x 1213 cm; base attenuate, not auricled, apex rounded or
acute; midrib raised above, lateral veins 4044 pairs, raised below, faintly sunken above, with short
intermediate veins, intercostal veins scalariform, raised below, faint to invisible above; petioles 27
cm long, softly hairy, lower half swollen, rugose on drying. Inflorescences (only male seen)
condensed, arising from leaf axils and from leafless twigs, 67 cm long, hairy; bracts lanceolate, c. 2
mm long, hairy; bracteoles ovate, c. 1 mm long. Flowers (male) sessile; calyx lobes triangular with
pointed tip, hairy, c. 1 mm long; petals oblong, c. 3 mm long, densely hairy inside; stamens free,
filaments densely hairy; disc 5-lobed; pistillodes absent. Infructescences 4.56 cm long. Fruits
oblong, 1.82 x 1.21.5 cm, dark brown velvety hairy, apex pointed, surface smooth.
Distribution. Endemic to Borneo. So far known from the type collection (with fruits) from Sarawak
and one flowering collection from Brunei (Wong WKM 611).
Ecology. Lowland forest. Flowering in January and fruiting in March.
Taxonomy. The short axillary inflorescence is a distinct diagnostic feature of M. condensata, a
character which is present only in M. axillaris, but the latter has glabrous leaves which are glaucous
underneath.
54
10. Melanochyla elmeri Merr.

(A.D.E. Elmer, 18701942, American botanist attached to the Bureau of Sciences, Manila)
PEB (1929) 169; Masamune l.c. 411; Ding Hou, FM 1, 8 (1978) 498; Anderson l.c. 137; Whitmore, Tantra & Sutisna l.c. 19.
Type: Elmer 2133, Borneo, Sabah, Tawau (holotype PNH, destroyed; isotypes BO, K, L, S, SING, NY, UC).
Medium-sized tree to 30 m tall, 40 cm diameter. Bark grey-brown, finely fissured; inner bark
brownish. Twigs rugose, yellowish hairy, glabrous. Leaves thin-coriaceous to coriaceous; upper
surface glabrous, except for the midrib, not shiny, lower surface sparsely hairy especially on midrib
and lateral veins, not glaucous; papillae distinct and uniformly distributed on the lower leaf surface;
narrowly obovate or oblong, 10.523 x 37.5 cm; base cuneate, not auricled, apex cuspidate; midrib
raised above, strongly channelled below, lateral veins 2736 pairs, raised below, very faint to almost
sunken above, intercostal veins scalariform, raised below, very faint to inconspicuous above; petioles
1.54 cm long, glabrous or hairy, channelled to flattened above, rugose at the lower part.
Inflorescences terminal hairy panicles, c. 15 cm long. Flowers whitish, subsessile; calyx lobes ovate
or ovate-oblong; petals oblong or oblong-lanceolate, longitudinally ridged inside; stamens filaments
free, staminodes small; disc 5-lobed; ovary conical, stigma minute, capitate. Fruits ellipsoid, 1.52.5
x 12 cm, rusty hairy, apex pointed, surface smooth.
Distribution. Endemic to Borneo; uncommon in Sabah and Sarawak; also in Brunei and Kalimantan.
11. Melanochyla fulvinervis (Blume) Ding Hou
Fig. 10E.
(Latin, fulvus = yellowish brown, nervus = vein)

Blumea 24 (1978) 32, FM 1, 8 (1978) 493; Anderson l.c. 137; Kochummen l.c. 42; Whitmore, Tantra & Sutisna l.c. 19;
Kessler & Sidiyasa l.c. 44. Basionym: Semecarpus fulvinervis Blume, Mus. Bot. Lugd. Bat. 1 (1850) 189, Merrill l.c. (1921)
35, Masamune l.c. 413. Lectotype (Ding Hou, 1978): Korthals, s.n. (= L. sheet no. 897. 3631228), Borneo, Mt. Parawan
(L).
Medium-sized tree to 24 m tall, 50 cm diameter. Bark chocolate-brown, smooth. Twigs brownish,

hairy. Leaves well-spaced, thin-coriaceous; upper surface shiny, lower surface glabrous or sparsely
hairy on midrib and veins, not glaucous; papillae in distinct groups; oblong, elliptic or oblanceolate,
1129.5 x 4.58 cm; base obtuse to subcordate or rarely cuneate, not auricled, apex acuminate,
acumen c. 2 cm long; midrib raised above, distinctly channelled below, lateral veins 1528 pairs,
raised on both surfaces, intercostal veins scalariform, distinct below, invisible above; petioles 0.51.5
cm long, densely short-hairy, channelled above. Inflorescences terminal hairy panicles, to 18 cm
long. Flowers white, subsessile; calyx lobes triangular; petals ovate or lanceolate, flat; stamen
filaments free, staminodes in female flowers usually large; disc 5-lobed; ovary conical, stigma
discoid; pistillodes small, hairy. Fruits ellipsoid or subglobose, 34 x 2.53 cm, rusty hairy, with long
insect-gall-like processes.
55
Distribution. Peninsular Malaysia, Borneo. Uncommon in Sabah and Sarawak; also in Kalimatan.
Ecology. Lowland to hill forest to 750 m.
Notes. S. 32352 (in fruit) and S. 32752 (male flowers) are different from the rest in having glabrous
leaves with cuneate base and 1.5-cm-long petioles.
12. Melanochyla minutiflora Ding Hou

(Latin, minutus = very small, floris = flower)
Blumea 24 (1978) 33, FM 1, 8 (1978) 497; Whitmore, Tantra & Sutisna l.c. 19. Type: Charington SAN 17810, Borneo,
Sabah (holotype L).
Small tree to 15 m tall, 30 cm diameter. Bark grey-brown, scaly; inner bark brown. Twigs densely
yellowish brown short-hairy. Leaves well-spaced, drying to chocolate-brown; lower surface not
papillose, sparsely hairy on midrib and veins; elliptic to oblong, 10.516.5 x 3.55 cm; base cuneate,
not auricled, apex acute; midrib raised above, lateral veins 1115 pairs, distinct below, faint above,
intercostal veins reticulate, distinct below, faint to inconspicuous above; petioles 0.71.3 cm long,
densely short-hairy, lower half swollen and greyish white on drying. Inflorescences terminal and
axillary hairy panicles, 56 cm long. Flowers (only male seen) yellow, sessile; calyx lobes triangular;
petals ovate or ovate-oblong, slightly longitudinally thickened at lower part inside; stamen filaments
free; disc 5-lobed; pistillodes absent, replaced by tufts of hair. Fruits subglobose, c. 1.2 cm diameter,
smooth, thinly velvety yellowish hairy, apex rounded.
Distribution. Endemic to Borneo. Uncommon, known from Sabah only from three collections, SAN
17810 (type), SAN 36562 from Lamag and SAN 24997 from Tawau areas.
13. Melanochyla montana Kochummen

(Latin, montanus = of mountains)
Sandakania 7 (1996) 86. Type: Mikil SAN 44330, Borneo, Sabah, Tambunan, Trus Madi FR (holotype SAN; isotype L).
Tree to 45 m tall, 200 cm diameter; bole with buttresses to 2.5 m high. Bark dark grey, smooth; inner
bark reddish brown with black sap. Sapwood reddish brown. with black sap, Sapwood reddish brown.
Bud scales stiff, lanceolate. Twigs stout, 1.2 cm thick. Leaves well-spaced, very thickly-coriaceous;
glabrous, faintly glaucous below; papillae uniformly distributed on the lower leaf surface; oblong to
oblanceolate or obovate, 1832 x 718 cm; base cuneate, not auricled, apex rounded or emarginate;
midrib prominently raised above, ahannelled below, lateral veins 1722 pairs, raised on both surfaces,
intercostal veins scalariform-reticulate, visible below, invisble above; petioles very stout, 34 cm
long, lower half strongly swollen and prominently rugose on drying. Flowers and fruits unknown.
56
Distribution. Endemic to Borneo. Known only from a few collections from Sabah and Sarawak from
lower montane forests where it appears to be one of the tallest trees. In Sabah, recorded from Ranau,
Sipitang and Tambunan districts (SAN 44330, SAN 75893, SAN 132820), and in Sarawak, from
Kelabit Highlands (S. 20060).
Ecology. Lower montane forest at 13701800 m.
Notes. Although this species is described from sterile material, it is sufficiently distinct.
14. Melanochyla scalarinervis Kochummen

(Latin, scalaris = ladder-like, nervus = veins; the leaf intercostal veins)
Sandakania 7 (1996) 88. Type: Ilias S. 37722, Borneo, Sarawak, Semengoh Arboretum (holotype SAR; isotypes MO, SAN).
Medium-sized tree to 22 m tall, 15 cm diameter. Bark blackish, smooth; inner bark with black sap.
Twigs 7 mm thick, grey, with minute lenticels. Leaves well-spaced, coriaceous; not glaucous below,
glabrous on both surfaces; papillae uniformly distributed on the lower leaf surface; oblong, 2125
x 6.57 cm; base cuneate, not auricled, apex blunt with short point; midrib raised above, lateral veins
c. 30 pairs, prominent below, faintly raised above, intercostal veins scalariform, raised below,
invisible above; petioles 1.52.5 cm long, whitish, rugose on drying. Inflorescences and flowers
unknown. Infructescences terminal, c. 30 cm long, velvety hairy, with very short branches, each of
which ends in a single fruit. Fruits ellipsoid, 22.5 x 1.52 cm, densely velvety hairy, pointed, with a
shallow cup-like swelling at the base, surface smooth.
Distribution. Endemic to Borneo; so far known from the type only from the Semengoh Arboretum in
Sarawak.
Ecology. Lowland forest. Fruiting noted in March.
Notes. M. scalarinervis appears rather similar to M. bullata, but the latter has bullate hairy leaves with
sunken lateral and intercostal veins.
15. Melanochyla semecarpoides Ding Hou

(Latin, semecarpoides = resembling Semecarpus)
Blumea 24 (1978) 33, FM 1, 8 (1978) 495; Whitmore, Tantra & Sutisna l.c. 19. Type: Sibat S. 21799, Sarawak, Ulu
Mayong, Kakus (holotype L; isotype SAR).
Small tree to 10 m tall, 10 cm diameter. Twigs whitish, glabrous. Leaves well-spaced, thincoriaceous; glabrous, drying reddish brown; upper surface not shiny, lower surface faintly glaucous;
papillae distinct and uniformly distributed on the lower leaf surface; oblong or oblanceolate, 1225
x 510 cm; base cuneate, not auricled, apex acute; midrib raised above, lateral veins 1116 pairs,
distinct below, faint above, intercostal veins reticulate, distinct below, very faint above; petioles 1.2
2.5 cm long, lower half swollen, whitish, rugose on
57
drying. Inflorescences terminal, male to 20 cm long, female shorter, minutely hairy. Flowers:
females slightly larger than males; calyx lobes triangular; petals triangular, longitudinally ridged
inside; stamens free; disc 5-lobed in male flower, entire in female flower; ovary hairy, globose, style
c. 1 mm long. Fruits ellipsoid, 1015 x 7 mm, apex pointed, surface rugose and finely hairy.
Distribution. Endemic to Borneo; known from three collections only (S. 21799, S. 22541, S. 43854)
from the Kapit district, Sarawak.
Ecology. Mixed dipterocarp forest, at 150200 m.
16. Melanochyla tomentosa Hook. f.

(Latin, tomentosus = covered with matted hairs; the leaf lower surface and twigs)
Fl. Br. Ind. 2 (1876) 38; King l.c. 503; Ridley l.c. 539; Ding Hou, FM 1, 8 (1978) 498; Kochummen l.c. 43. Type: Maingay
1803, Malacca (holotype K).
Medium-sized tree to 20 m tall, 37 cm diameter. Bark reddish brown, smooth; inner bark reddish with
black sap. Sapwood white. Young twigs velvety yellowish hairy. Leaves coriaceous, not bullate;
lower surface not glaucous, densely velvety hairy; papillae distinct and uniformly distributed on the
lower leaf surface; elliptic, oblong or obovate, 5.519 x 2.59 cm; base cuneate, not auricled, apex
rounded or acute; midrib raised above, lateral veins 1222 pairs, raised below, faint above,
intercostal veins scalariform-reticulate, raised below, invisible to sunken above; petioles 13 cm long,
velvety hairy, base swollen. Inflorescences terminal, to 30 cm long, hairy. Flowers (only male seen)
sessile; calyx and petals dark brown to yellowish hairy; calyx lobes triangular; petals elliptic to
elliptic-lanceolate, flat; stamen filaments free; disc 5-lobed; pistillodes reduced to tufts of hairs.
Fruits ellipsoid, globose or ovoid, 1.53 cm diameter, velvety hairy, surface smooth.
Distribution. Sumatra, Peninsular Malaysia, Borneo; very uncommon in Sabah and Sarawak. In
Sabah, known from a single collection, SAN 57370, from Semporna district; and in Sarawak, recorded
from Mulu National Park only (S. 39356).
Ecology. Lowland mixed dipterocarp forest, at 150250 m.
17. Melanochyla woodiana Kochummen

(G.H.S. Wood, 19271957, first forest botanist in Sabah)
Sandakania 7 (1996) 88. Type: G.H.S. Wood SAN 16969, Borneo, Sabah, Beaufort (holotype SAN; isotypes A, BO, BRI, K,
KEP, L, SING).
Medium-sized tree to 20 m tall. Twigs whitish, densely brown short-hairy. Leaves well-spaced,
coriaceous; glabrous, upper surface not shiny, lower surface not glaucous; papillae distinct and
uniformly distributed on the lower leaf surface; elliptic or oblong, 17.519 x 910 cm; base broadly
cuneate to obtuse, not auricled, apex cuspidate; midrib raised above, lateral veins c. 18 pairs, raised
below, very faint to sunken above, intercostal veins scalariform-reticulate,
58
raised below, invisible above; petioles 1.52.5 cm long, densely short-hairy, lower half swollen,
whitish and rugose on drying. Inflorescences and flowers unknown. Infructescences terminal, c. 30
cm long, with a number of long side-branches, finely brown short-hairy. Fruits ovoid or ellipsoid,
1.72 x 11.2 cm, apex sharp-pointed, surface rugose and densely brown short-hairy.
Distribution. Endemic to Borneo; so far known from the type only from Sabah.
Ecology. Lowland mixed dipterocarp forest, at c. 120 m. Fruiting noted in July.
11. PARISHIA Hook. f.

(C.S. Parish, 18221897, English botanist
in British Burma, now Myanmar)
layang-layang (Malay), upi (Iban)
Trans. Linn. Soc. 23 (1860) 169; Ridley, FMP 1 (1922) 535; Browne, FTSB (1955) 51; Burgess, TBS (1966) 27; Ding Hou,
FM 1, 8 (1978) 541; Anderson, CLTS (1980) 38; Wong, DMT (1982) 117; Corner, WSTM 3rd ed. 1 (1988) 121;
Kochummen, TFM 4 (1989) 44; Whitmore, Tantra & Sutisna, CLK 1 (1989) 20; Kessler & Sidiyasa, Trees of the
Balikpapan-Samarinda Area (1994) 45.
Medium-sized to tall emergent dioecious trees; bole with tall spreading buttresses. Bark smooth to
shallowly fissured or scaly; inner bark pink or reddish, often with pale stripes, laminated or soft, with
abundant white sap which darkens on drying. Sapwood whitish. Leaves spirally arranged, trifoliolate
or pinnately compound with opposite or subopposite leaflets and terminal leaflets. Inflorescences
axillary and/or terminal panicles. Flowers unisexual; calyx 4-lobed, enlarged and persistent in fruit;
petals 4, imbricate, glabrous or sparsely hairy outside; stamens 4, filaments long, glabrous, anthers
dorsifixed or dorso-basifixed, abortive in female flowers; disc intrastaminal, hairy, pulvinate or flat,
4-lobed or 4-grooved; ovary superior, densely hairy, 1-locular, style 3(rarely 4)-lobed, pistillodes very
small. Fruits 1-locular, densely brown-hairy, with an enlarged persistent calyx with wing-like lobes.
Seeds with testa fused to the endocarp; embryo straight, cotyledons free, plano-convex.
Distribution. Seven species; Andaman Islands, Myanmar, Thailand, Malesia; 6 species in Sabah and
Sarawak.
Ecology. Lowland, including swamp, to lower montane forest, to 1500 m. The trees are deciduous
and flowers appear before or during the onset of new leaves.
Uses. The timber is known in the ASEAN region as lelayang. The wood is moderately hard with
density of 545755 kg/m3 air-dried; the heartwood is red-brown to light brown; and texture is fine and
even with interlocking grain. Not durable and susceptible to powder post beetles. It is suitable for
light construction, general planking, pallets, packing crates as well as core veneers for plywood
manufacture.
59
Key to Parishia species

1. Leaves trifoliolate; leaflets 1015 cm wide; intercostal veins strongly reticulate, distinct
on both surfaces...........................................................................................6. P. trifoliolata
Leaves pinnately compound; leaflets less than 10 cm wide; intercostal veins weakly
reticulate, usually very faint................................................................................................2
2. Leaflets concave, reddish-brown hairy below; lateral veins sunken above...........................
..................................................................................................................1. P. dinghouiana
Not this combination of characters......................................................................................3
3. Leaflets with asymmetric rounded or subcordate base; petiolules very short, to
2 mm long......................................................................................................2. P. insignis
Leaflets with symmetric cuneate base; petiolules well-developed, much longer.................4
4. Leaves with 2 pairs of leaflets.......................................................................4. P. paucijuga
Leaves with more than 2 pairs of leaflets.............................................................................5
5. Enlarged calyx lobes shorter than the fruits.......................................................5. P. sericea
Enlarged calyx lobes very much longer than the fruits..................................3. P. maingayi
1. Parishia dinghouiana Kochummen

(Ding Hou, b. 1921, botanist at the Rijksherbarium, Leiden)
Sandakania 7 (1996) 91. Type: Jugah & Sibat S. 26300, Borneo, Sarawak, Kuching (holotype SAR; isotypes A, BO, K, L,
SING).
Small tree to 15 m tall, 30 cm diameter. Twigs stout, 1.52 cm thick, brown-hairy. Leaves pinnately
compound; rachis flattened above, hairy. Leaflets glabrous above, reddish-brown hairy below,
concave; oblong, 1018.5 x 67.5 cm; base broadly cuneate, apex acute; midrib raised above, lateral
veins 1521 pairs, sunken above, raised below, with short intermediate veins, intercostal veins weakly
reticulate, faint below, invisible above; petiolules 3 mm long, densely hairy. Inflorescences and
flowers unknown. Infructescences terminal, c. 70 cm long, densely hairy, axes stout, 11.5 cm thick.
Fruits (immature) ovoid, c. 7 x 5 mm, velvety hairy; calyx tube c. 2 cm long, 2.5 cm wide, calyx lobes
c. 5.5 x 1.5 cm, with distinct reticulate veins.
Distribution. Endemic to Borneo; so far known from the type only from Sarawak.
Ecology. Lowland forest by streams.
2. Parishia insignis Hook. f.
(Latin, insignis = remarkable)
Trans. Linn. Soc. 23 (1860) 170; Ridley l.c. 536; Burgess l.c. 27; Ding Hou, FM 1, 8 (1978) 545; Anderson l.c. 138; Corner
l.c. 121; Kochummen l.c. 45; Whitmore, Tantra & Sutisna l.c. 20; Kessler & Sidiyasa l.c. 45. Type: Griffith 1037, Burma
(holotype K; isotype SING). Synonym: P. borneensis Ridl.,
60
Kew Bull. (1933) 20, Masamune l.c. 412; P. lowei Ridl. l.c. (1933) 201, Masamune l.c. 412.
Emergent tree to 40 m tall, 60 cm diameter; bole with tall buttresses. Bark brownish, cracking; inner
bark reddish, laminated. Sapwood whitish. Twigs stout, 1.51.8 cm thick, with scattered whitish
lenticels, youngest ones angular. Leaves pinnately compound, each with 210 pairs of opposite or
subopposite leaflets; basal part of rachis flattened above, sparsely hairy to hairy. Leaflets glabrous or
sparsely hairy below, coriaceous; oblong or ovate to lanceolate, 6.515 x 3.56.5 cm; base obtuse to
subcordate, asymmetric, apex acute; midrib raised above, lateral veins 1017 pairs, raised below,
faint above, intercostal veins weakly reticulate, faint but visible on both surfaces; petiolules very
short, to 2 mm long, or leaflets sessile. Inflorescences axillary panicles, to 54 cm long, rusty hairy;
bracts triangular, lanceolate or narrowly elliptic, 23 mm long, minutely hairy outside, glabrous
inside; pedicels 25 mm long. Flowers: calyx 2.54.5 mm long, minutely hairy on both surfaces,
lobes triangular, unequal, 23 mm long; petals broadly ovate to ovate-oblong or elliptic, 35 x 1.53
mm; stamens 2.54 mm long, anthers ovoid, c. 0.75 mm long; staminodes c. 1.5 mm long; disc fleshy,
flat, round or slightly 4-angular, or shallowly cup-shaped, hairy, 22.7 mm diameter; ovary conical, c.
1.5 mm diameter, style c. 1.5 mm long, stigma capitate. Fruits ovoid, 1.52 x 0.71 cm, velvety
hairy, with a distinct beak; calyx lobes, 1015 x 11.5 cm; stalks to 1.5 cm long.
Vernacular name. Sarawakupi bunga (preferred name).
Distribution. Andaman Is., Myanmar, Thailand, Peninsular Malaysia, Sumatra, Borneo. Uncommon
in Sabah and Sarawak; also in Kalimantan.
Ecology. Lowland forest, including swamp forest.
3. Parishia maingayi Hook. f
Fig. 11.
(A.C. Maingay, 18361869; 18621868 magistrate in charge of the jail in Malacca)

Fl. Br. Ind. 2 (1876) 30; King l.c. 493; Ridley l.c. 535; Burgess l.c. 27; Ding Hou, FM 1, 8 (1978) 543; Anderson l.c. 138;
Corner l.c. 122; Kochummen l.c. 44; Whitmore, Tantra & Sutisna l.c. 20. Type: Maingay 488, Malacca (holotype K; isotype
L).
Medium-sized to emergent tree to 36 m tall, 95 cm diameter; bole with steep buttresses to 1.5 m tall,
or with stilt-roots in swampy habitats. Bark dark brown, shallowly fissured to scaly; inner bark
reddish, with abundant white latex. Sapwood whitish. Twigs slender to very thick, 0.52 cm thick,
glabrous, often lenticellate. Leaves pinnately compound, each with 48 pairs of leaflets; rachis
slightly swollen at base. Leaflets coriaceous, often shiny above, very rarely sparsely reddish-brown
hairy below; elliptic, ovate, or oblong, 416.5 x 1.86 cm; base cuneate to rounded, symmetrical,
apex acute, rarely rounded or obtuse and emarginate; midrib raised above and prominent beneath,
rarely flattened and faint above, lateral veins 1012 pairs, usually very faint to almost invisible on
both surfaces, intercostals veins weakly reticulate, very faint and hardly distinguishable from the
lateral veins; petiolules 48 mm long, rugose on drying, sometimes faintly channelled above.
Inflorescences axillary or terminal panicles, hairy, to 50 cm long, with many long branches,
sometimes smaller ones with few short branches; axes occasionally very thick, almost as thick as the
twig; bracts ovate, 23.5 mm long, minutely hairy outside; pedicels 1.53 mm long. Flowers white;
calyx 26 mm long, densely appressed hairy on both surfaces, lobes triangular, unequal, 0.61 mm
long; petals
61
Fig. 11. Parishia maingayi. A, leafy twig; B, detail of lower leaflet surface; C, inflorescence; D, flower in longitudinal
section; E, fruit. (AB from SAN 15031, CD from Ladi BRUN 5108, E from SAN 16328.)
62
oblanceolate or narrowly oblong, 58 x 12 mm, sparsely hairy outside; stamens 34 mm, inserted in
the grooves at the lower half of the disc, anthers ovoid, c. 1 mm long, staminodes c. 2 mm long; disc
pulvinate and 4-grooved, c. 1.2 mm diameter in male flowers, fleshy and shallowly cup-shaped, c. 2.5
mm diameter in female flowers; ovary conical, c. 1.5 mm diameter, style c. 1 mm long, stigmas
capitate. Fruits ovoid or broadly ellipsoid, 1.52.5 x 11.5 cm, velvety hairy, gradually narrowed into
a beak; enlarged calyx lobes hairy on both surfaces, red when fresh, oblong, 613.5 x 11.8 cm.
Vernacular name. Sarawakupi paya (preferred name).
Distribution. Sumatra, Peninsular Malaysia, Borneo and the Philippines. Common in Sabah and
Sarawak; also in Brunei and Kalimantan.
Ecology. Lowland, including swamp and kerangas, to lower montane forest, to 1450 m. Flowering in
February to June and fruiting in March to June.
Taxonomy. There is considerable variation in the shape, texture and pubescence of leaflets, the
thickness of the twigs, and branching as well as thickness of axes of the inflorescences. Two varieties
are recognised.
Key to varieties
Leaflet apex acute, not emarginate; midrib prominently raised above..........................................
var. maingayi
Synonyms: P. elmeri Merr., PEB (1929) 168, Masamune l.c. 412; P. polycarpa Ridl., Kew
Bull. (1933) 200, Masamune l.c. 412.
Distribution: as for the species.
Leaflet apex obtuse or rounded-emarginate; midrib not prominent, flattened above....................
var. minor (Ridl.) Kochummen
Sandakania 7 (1996) 94. Basionym: Parishia minor Ridl. l.c. (1933) 201, Masamune l.c. 412.
Known from the Kuching area in Sarawak.
4. Parishia paucijuga Engl.

(Latin, pauci = few, jugum = pair; the few pairs of lealets)
In A. DC., Mon. Phan. 4 (1883) 309; Ridley l.c. (1922) 536; Ding Hou, FM 1, 8 (1978) 544; Anderson l.c. 138; Corner l.c.
122; Kochummen l.c. 46; Whitmore, Tantra & Sutisna l.c. 20. Type: Maingay 496, Malacca (holotype K).
Medium-sized tree to 30 m tall, 20 cm diameter. Leaves pinnately compound, each with two pairs of
leaflets; rachis finely hairy, flattened above at the base. Leaflets coriaceous; oblong, 1015 x 57 cm;
base cuneate, symmetric, apex rounded with short point; midrib raised above, lateral veins c. 20 pairs,
faint above, intercostal veins weakly reticulate, very faintly visible above; petiolules to 7 mm long,
finely hairy, rugose on drying. Inflorescences to 35 cm long, hairy, with many branches; bracts
triangular, 25 mm long, usually densely hairy outside; pedicels absent or c. 0.6 mm long. Flowers
white; calyx 34.5 mm long, densely hairy on both
63
surfaces, lobes triangular, 23 mm long; petals obovate-oblong or oblong, 36 x 1.52 mm, sparsely
hairy outside; stamens inserted at the base of the disc, 33.5 mm long, anthers oblong, 0.71 mm
long; staminodes c. 2 mm long; disc pulvinate, 4-grooved and 1.52 mm diameter in male flowers,
shallowly cup-shaped and 2.53 mm diameter in female flowers; ovary conical, c. 2 mm diameter,
style c. 1.7 mm long, stigmas capitate. Fruits ellipsoid, 46 x 2 cm, apex acute; calyx lobes hairy on
both surfaces, shorter than the fruit.
Distribution. Sumatra, Peninsular Malaysia, Borneo. In Sarawak, so far known only from two
collections (S. 404 and S. 15508) from Lundu area, not yet reported from Sabah; also in Brunei and
Kalimantan.
Notes. S. 15508 from Sarawak has young fruits with the calyx lobes longer than the fruits.
5. Parishia sericea Ridl.

(Latin, sericeous = with silky long appressed hairs)
l.c. (1933) 201; Masamune l.c. 412; Burgess l.c. 27; Ding Hou, FM 1, 8 (1978) 544; Whitmore, Tantra & Sutisna l.c. 20.
Type: Haviland 2273, Borneo, Sarawak Kuching (holotype K).
Tree to 25 m tall, 40 cm diameter; bole with short buttresses. Bark scaly. Leaves pinnately compound,
each with 47 pairs of leaflets; rachis minutely hairy. Leaflets coriaceous, shiny above, faintly hairy
below; ovate-oblong to lanceolate, 6.517 x 36.5 cm; base cuneate to rounded, symmetric, apex
obtuse or acute; lateral veins 1420 pairs, raised and prominent below, visible or obscure above,
intercostal veins weakly reticulate, faintly visible on both surfaces; petiolules to 1 cm long.
Inflorescences to 44 cm long, hairy, with long branches; bracts ovate to lanceolate, c. 2 mm long,
densely minute-hairy outside; pedicels absent or to 0.7 mm long. Flowers (only male seen): calyx c. 3
mm long, hairy on both surfaces, lobes triangular, c. 1.5 mm long; petals oblanceolate, glabrous, 5
x11.5 mm; stamens c. 3 mm long, inserted at the base of the disc, anthers ovoid, c. 0.7 mm long;
disc pulvinate, 4-grooved, 1.51.7 mm diameter. Fruits ellipsoid, 55.7 x 1.7 cm; calyx tube c. 0.7
cm long; calyx lobes oblong or lanceolate, 23 x 0.51 cm, shorter than the fruit, brown to dark
brown hairy on both surfaces.
Distribution. Endemic to Borneo. Locally common, recorded from Lahad Datu and Ranau districts in
Sabah, and Bintulu and Kuching areas in Sarawak; also in Brunei and Kalimantan.
6. Parishia trifoliolata Kochummen

(Latin, tri = three, foliolum = leaflet; leaf with three leaflets)
Sandakania 7 (1996) 91. Type: Yii S. 42974, Borneo, Sarawak, Samunsam Wildlife Sanctuary (holotype KEP; isotypes K, L,
SAN, SAR).
Small tree 12 m tall, 10 cm diameter. Twigs brown, c. 8 mm thick. Leaves trifoliolate; rachis
64
glabrous, with many close grooves. Leaflets thick-coriaceous, drying to yellowish green; oblong, 17
21 x 1015 cm; base rounded, apex rounded with a very short tip; midrib sunken above at least along
the upper half, lateral veins 1416 pairs, faintly visible on both surfaces, intercostal veins strongly
reticulate, distinct on both surfaces; petiolules 11.5 cm long, rugose. Inflorescences and flowers
unknown. Infructescences terminal, more than 40 cm long, with long branches. Fruits (immature)
ovoid, c. 7 x 6 mm, velvety hairy; calyx tube c. 1 cm long; calyx lobes dark red when fresh, dark
brown velvety hairy on both surfaces, c. 3 cm long.
Distribution. Endemic to Borneo; so far known only from the type specimen from the Samunsam
Wildlife Sanctuary in Sarawak.
Ecology. Lowland kerangas forest.
Taxonomy. This species differs from the other species of the genus by its trifoliolate leaves, large
leaflets with distinct reticulate intercostal veins and the dark brown velvety hairy calyx lobes.
12. PENTASPADON Hook. f.

(Greek, penta = five, spadon = eunuch;
the presence of five sterile stamens in the flowers)
pelajau (Iban), pelong (Malay)
Trans. Linn. Soc. 23 (1860) 168; Merrill, EB (1921) 351; Ridley, FMP 1 (1922) 537; Masamune, EPB (1942) 413; Browne,
FTSB (1955) 52; Ding Hou, FM 1, 8 (1978) 520; Anderson, CLTS (1980) 138; Wong, DMT (1982) 177; Corner, WSTM
3rd ed. 1 (1988) 122; Kochummen, TFM 4 (1989) 46; Whitmore, Tantra & Sutisna, CLK 1 (1989) 20; Ng, Mal. For. Rec.
34, 1 (1991) 18.
Large deciduous trees. Leaves spirally arranged, clustered at the ends of twigs, pinnately compound
with terminal leaflet, with hairy domatia in the axils of lateral veins. Inflorescences axillary,
paniculate. Flowers bisexual, calyx 5-lobed; petals 5, imbricate; fertile stamens 5, sterile stamens 5,
inserted outside the disc, anthers basifixed or dorsifixed; disc cup-shaped, grooved or wavy outside;
ovary superior, 1-locular, style short, stigma subglobose or 2-lobed. Fruits 1-locular, ovoid. Seeds
one per fruit; testa free from endocarp; germination epigeal, in P. motleyi seeds germinate within 12
30 days, cotyledons emergent, fleshy; the first two leaves opposite and divided into 35 leaflets with
dentate margins.
Distribution. Six species; Thailand, Vietnam, Sumatra, Peninsular Malaysia, Borneo, Maluku, New
Guinea and Solomon Islands; one species in Sabah and Sarawak.
Ecology. Lowland and hill forests.
Uses. The timber, known by the ASEAN standard name as pelajau, is suitable for interior finishing,
panelling, partitioning, moulding, flooring, etc. It has been successfully used as a peeler-log for
plywood manufacture. It is fairly hard and fairly heavy with density of 480835
65
Fig. 12. Pentaspadon motleyi. A, flowering leafy twig; B, part of inflorescence; C, flower with 1 sepal and 2 petals removed;
D, fruit. (A from SAN 27312, BC from Hamid FMS 13252, D from SAN 25959.)
66
kg/m3 air-dried, easy to saw and cross-cut, and moderately durable under exposed conditions; the
sapwood is not clearly differentiated from the heartwood. The oil obtained from P. motleyi is used to
cure certain skin diseases; and the fruits are edible after boiling.
Pentaspadon motleyi Hook. f.
Fig. 12.
(J. Motley, civil engineer and plant collector in Labuan and Banjarmasin in the 1850s)
l.c. 168; Merrill l.c. (1921) 351; Ridley l.c. (1922) 538; Masamune l.c. 413; Burgess, TBS (1966) 30; Ding Hou, FM 1, 8
(1978) 524; Anderson l.c. 138; Corner l.c. 122; Kochummen l.c. 47; Whitmore, Tantra & Sutisna l.c. 20; Ng l.c. 18. Type:
Motley, s.n., Borneo (holotype K).
Medium-sized tree to 30 m tall, 60 cm diameter; bole with spreading buttresses. Bark grey- brown,
smooth to scaly; inner bark pink, laminated, with droplets of white sap which turns to brown resin
after a few days. Sapwood white. Twigs grey-brown. Leaves each with 45 pairs of subopposite
leaflets and a terminal leaflet, the basal pair always smaller. Leaflets thin-coriaceous, glabrous, lower
surfaces often with hairy domatia in the axils of lateral veins; elliptic-oblong or ovate, 4.513 x 35.5
cm; base obtuse, apex acute; midrib raised above, lateral veins 810 pairs, raised below, faint above,
with short intermediate veins; leaflets of seedlings with dentate margins; petiolules 12 mm long.
Inflorescences to 30 cm long. Flowers white; calyx lobes broadly ovate; petals obovate to oblong;
anthers bent towards the centre. Fruits ovoid, sharply pointed, green with brown spots when fresh,
2.54 x 1.82.5 cm; edible.
Vernacular names. Sarawakbuah pusit (Penan), emplangau (Iban), japing (Kenyah), lakacho
(Murut), pelasih (Berawan), pelajau (Iban), perajau (Bidayuh), umit (Iban), uping (Kayan).
Distribution. Sumatra, Peninsular Malaysia, Borneo, Maluku, New Guinea, Solomon Islands. Widely
distributed in Sabah and Sarawak, but less common in Sarawak; also in Brunei and Kalimantan.
Ecology. Lowland forest, usually by rivers, to 300 m. Flowering twice a year along with new foliage.
13. PLEIOGYNIUM Engl.

(Greek, pleio = more than usual, gyne = female organs;
the number of carpels and styles)
In A. DC., Mon. Phan. 4 (1883) 255; Ding Hou, FM 1, 8 (1978) 474; Whitmore, Tantra & Sutisna, CLK 1 (1989) 20.
Dioecious trees. Leaves spirally arranged, pinnately compound, usually with terminal leaflet. Leaflets
opposite, entire. Inflorescences axillary panicles. Flowers unisexual; calyx 5 (rarely 4 or 6)-lobed;
petals 5 (rarely 4 or 6), imbricate; stamens 10 (rarely 812), twice the number of petals, anthers
abortive or imperfect in female flowers; disc annular, pulvinate; ovary 512-locular, style 512,
stigmas spathulate. Fruits 512-
67
Fig. 13. Pleiogynium timorense. A, flowering leafy twig; B flower buds; CD, flowers. (All from Kostermans 28246.)
68
locular, with 512 seeds; endocarp hard, woody. Seeds with testa free from the endocarp; embryo
slightly curved, cotyledons free, plano-convex.
Distribution. Two to three species; Malesia, Australia, Pacific islands; one species in Sabah.
Pleiogynium timorense (DC.) Leenh.
Fig. 13.
(of Timor island)

Blumea 7 (1952) 159; Ding Hou, FM 1, 8 (1978) 474; Whitmore, Tantra & Sutisna l.c. 21. Basionym: Icica timorensis DC.,
Prod. 2 (1825) 78. Type: Sine coll., s.n., Timor (G, n.v.).
Tree to 36 m tall, 75 cm diameter; bole with buttresses to 2.5 m high. Bark dark grey to grey-brown,
fissured or scaly. Leaves each with 36 pairs of leaflets. Leaflets glabrous or sparsely hairy, lower
surface with hairy domatia in the axils of lateral veins; elliptic to oblong or lanceolate, rarely ovate or
obovate, 3.513.5 x 26 cm; base asymmetric, cuneate, apex acute; lateral veins 811 pairs, faint on
both surfaces, intercostal veins reticulate; petiolules 0.51 cm long. Inflorescences: males to 30 cm
long, with branches to 8.5 cm long, females usually to 3.5 cm long. Flowers greenish yellow; calyx
lobes suborbicular, 0.50.7 mm long; petals ovate-oblong, 1.53 x 12.5 mm; stamens 23 mm long,
those opposite calyx lobes usually longer than those opposite petals, anthers 0.50.6 mm across;
staminodes 0.61 mm long; disc 1.51.7 mm diameter; ovary subglobose, c. 1 mm diameter, glabrous,
styles c. 0.5 mm long; pistillodes 0.61 mm across. Fruits broadly ovoid, 1.51.7 x 1.52 cm, red to
dark brown when ripe, with distinct scars of styles near the apex.
Distribution. Borneo, Lesser Sunda Islands, Philippines, Maluku, New Guinea, Australia, Fiji,
Solomon Islands, and Pacific Islands. In Borneo, known only from a single collection (NBFD 9227)
from Sabah.
14. RHUS L.
(Greek, rhous = an old plant-name for Rhus coriaria L.)
Gen. Pl. ed 5 (1754) 129; Engler, Bot. Jahrb. 1 (1881) 378; Merrill, EB (1921) 351; Ridley, FMP 1 (1922) 538; Brizicky, J.
Arn. Arb. 44 (1963) 62; Backer & Bakhuizen f., FJ 2 (1965) 153; Ding Hou, Blumea 24 (1978) 34, FM 1, 8 1978) 534;
Kochummen, TFM 4 (1989) 48.
Small trees, scandent shrubs or woody climbers. Leaves spirally arranged, pinnately compound, each
with 121 leaflets, or very rarely simple (R. borneensis); lateral leaflets in compound leaves opposite.
Inflorescences terminal or axillary panicles or racemes. Flowers unisexual or bisexual; calyx 5lobed; petals 5, imbricate; stamens 5, anthers dorsifixed,
69
imperfect or sterile in female flowers; disc intrastaminal, shallowly cup-shaped, cup-shaped or round
and flat; ovary 1-locular, superior, abortive in male flowers, style short, distinct or obscure, stigmas 3,
free or united, capitate or obscure. Fruits 1-locular, without wings. Seeds with testa adherent to or
free from the endocarp; embryo straight, cotyledons free, flat.
Distribution. About 200 species; widely distributed in temperate zones of both hemispheres with a
few species extending to the tropics and subtropics. Two species are recorded from Sabah and
Sarawak, one of which is endemic to Borneo.
Ecology. From sea-level to 2400 m.
Uses. There are no recorded local uses of the indigenous species. However, the galls of R. chinensis, a
native of China, are medicinal and imported to Malaysia. R. verniciflua is cultivated in China and
Japan for its lacquer.
Key to Rhus species

Leaves simple. Shrub....................................................................................................................
R. borneensis Stapf
FMK (1894) 142; Merrill l.c. (1921) 35; Masamune l.c. 413; Ding Hou, FM 1, 8 (1978) 541.
Synonym: Toxicodendron borneense (Stapf) Gillis, Rhodora 73 (1971) 164.
Shrub or small tree to 3 m high. Inflorescences axillary, paniculate. Fruits (young)
subglobose, 57 mm diameter.
Endemic to Borneo; uncommon, recorded from Sabah and Sarawak. Montane forest, at 1200
2300 m.
Leaves pinnately compound; scandent shrub or woody climber, rarely small tree......R. nodosa
Rhus nodosa Blume
Fig. 14.
(Latin, nodosus = knotty; probably referring to the twigs)

Bijdr. Fl. Ned. Ind. (1826) 1164; Backer & Bakhuizen f. l.c. 153; Ding Hou, FM 1, 8 (1978) 539. Type: Blume, s.n., Java,
Mount Salak (holotype L). Synonyms: R. perakensis Scort. ex King, J. As. Soc. Beng. 65, 3 (1896) 500, Masamune l.c. 413;
Toxicodendron nodosum (Blume) Gillis l.c. 168.
Scandent shrub or woody climber, rarely small tree to 9 m tall. Twigs strongly lenticellate. Leaves
copper-red when young, each with (1)23(5) pairs of leaflets; rachis 410 cm long. Leaflets thincoriaceous; lower surface minutely hairy, early glabrescent, domatia present in the axils of lateral
veins; ovate, elliptic or elliptic-lanceolate, 415 x 26 cm; base cuneate, attenuate or obtuse, margin
entire, apex attenuate-acuminate; lateral veins 1223 pairs, intercostal veins reticulate and rather faint
on both surfaces; petiolules absent or to 0.7 cm long, of terminal leaflets absent to 2.3 cm long.
Inflorescences axillary or pseudoterminal, to 40 cm long. Flowers creamy or yellowish; calyx lobes
triangular, 0.50.7 mm long; petals ovate-elliptic, 23 x 11.5 mm, glabrous; stamens 1.52 mm long,
anthers ovoid, c. 1 mm long; disc shallowly cup-shaped, c. 1 mm diameter; ovary subglobose, c. 1
mm diameter. Fruits subglobose, 57 x 58 mm.
70
Fig. 14. Rhus nodosa. A, fruiting leafy twig; B, part of inflorescence; C, flower with one sepal and one petal removed. (AB
from S. 43015, C from S. 16547.)
71
Distribution. Sumatra, Peninsular Malaysia, Borneo. In Sarawak, uncommon, so far recorded only
from Baram, Bau, Kelabit Highlands, the Kuching area and the Hose Mountains; not yet reported
from Sabah.
15. SEMECARPUS L. f.
(Greek, sema = mark, karpos = fruit; named after
the marking nut-tree, Semecarpus anacardium)
rengas (Malay)
Suppl. Pl. (1782) 25; Hooker f., Fl. Br. Ind. 2 (`1876) 30; Merrill, EB (1921) 351; Ridley, FMP 1 (1922) 54; Masamune,
EPB (1942) 413; Ding Hou, Blumea 24 (1978) 35, FM 1, 8 (1978) 499; Anderson, CLTS (1980) 138; Corner, WSTM 3rd
ed. 1 (1988) 131; Kochummen, TFM 4 (1989) 50; Whitmore, Tantra & Sutisna, CLK 1 (1989) 21. Synonym:
Melanocommia Ridl., Kew Bull. (1933) 198, Masamune l.c. 411.
Small trees or shrubs, sometimes unbranched, often spiny when young with irritant creamy or black
sap. Leaves simple, spirally arranged and well-spaced, in close spirals or in pseudowhorls, often
papillose beneath, margin not thickened. Inflorescences paniculate or racemose, terminal, extraaxillary or axillary, rarely borne on leafless stems (not in Borneo), pedicels articulate. Flowers
unisexual (plants dioecious), rarely bisexual; female flowers usually larger than males; calyx 5- or 4lobed, petals 5 or 4, imbricate or valvate; stamens 5 or 4, filaments subulate, glabrous, anthers
dorsifixed, stamens in female flowers shorter, sterile; disc intrastaminal, often 5- or 4-notched, round,
flat, dish-shaped, cup-shaped or funnel-shaped; ovary superior, rarely semi-inferior, 1-carpellate, 1locular, usually densely hairy, styles 3, often hairy near the base, stigmas transversely oblong or
subreniform; pistillodes minute or absent or replaced by a tuft of hairs. Fruits 1-locular, often
laterally compressed, seated on a distinct fleshy hypocarp (formed by calyx and floral axis). Seeds
with testa free from the endocarp; cotyledons free, plano-convex.
Distribution. About 75 species; India, Sri Lanka, Myanmar, Thailand, Indo-China, Formosa, Malesia,
Australia, Micronesia, Solomon Islands, New Caledonia and Fiji; 19 species in Sabah and Sarawak.
Ecology. Lowland to montane forest, to 1950 m, including swampy and limestone habitats.
Uses. No reported use in Sabah and Sarawak. The pericarp is filled with black irritant resin, but the
fleshy fruit of certain species such as S. anacardium can be eaten after roasting. The black sap of S.
anacardium, when mixed with lime water or alum, is used for a water-resistant marking-ink.
Key to Semecarpus species
1. Leaves glabrous (sometimes only very sparsely hairy below)..............................................2

Leaves distinctly hairy (at least below)................................................................................17
2. Leaf midrib sunken above......................................................................................................3
Leaf midrib raised or flattened above.....................................................................................6
72
3. Leaf lateral veins more than 20 pairs; leaf margin recurved; midrib below and petiole pale
yellow on drying..................................................................................14. S. minutipetalus
Leaf lateral veins less than 20 pairs; leaf margin not recurved; midrib and petiole not pale
yellow on drying..................................................................................................................4
4. Leaves distinctly glaucous below........................................................9. S. glaucus (in part)
Leaves not or only slightly glaucous below.........................................................................5
5. Leaf lateral veins faint on the lower surface..............................................8. S. euodiifolius
Leaf lateral veins raised on the lower surface..............................6. S. cuneiformis (in part)
6. Papillae distinct and in groups on the lower leaf surface.........................18. S. sandakanus
Papillae obscure or distinct but uniformly distributed on the lower leaf surface.................7
7. Leaves spirally arranged, well-spaced.................................................................................8
Leaves in pseudowhorls or in close spirals........................................................................10
8. Leaf apex rounded; lateral veins to 10 pairs..........................................15. S. paucinervius
Leaf apex acute or obtuse; lateral veins more than 10 pairs.................................................9
9. Young stem not spiny. Hypocarp disc-like...........................................10. S. heterophyllus
Young stem spiny. Hypocarp not disc-like................................4. S. bunburyanus (in part)
10. Leaves narrow, the length more than 4 times the width.....................................................11
Leaves broader, the length less than 4 times the width......................................................13
11. Petioles short, to 1 cm long; leaf base obtuse...........................................2. S. angustifolius
Petioles longer; leaf base attenuate....................................................................................12
12. Leaves to 2 cm wide; petioles to 1.5 cm long, base slightly swollen..............13. S. lineatus
Leaves wider than 3.5 cm; petioles 4.56 cm long, base prominently swollen.....................
...................................................................................................................16. S. pulvinatus
13. Leaves sessile............................................................................4. S. bunburyanus (in part)
Leaves with distinct petioles.............................................................................................14
14. Inflorescences or infructescences extra-axillary. Leaf intercostal veins finely reticulate.
Trees of limestone habitats............................................................................5. S. calcicolus
Inflorescences or infructescences axillary or terminal. Leaf intercostal veins laxly
reticulate. Trees of non-limestone habitats........................................................................15
15. Leaf margin wavy. Inflorescence axes angular.............................................1. S. angulatus
Leaf margin entire. Inflorescence axes not angular (or not known)...................................16
16. Leaves glaucous below. Hypocarp cup-shaped, enclosing more than three-quarters of the
fruit ...............................................................................................................7. S. cupularis
Leaves not or only rarely glaucous below. Hypocarp not cup-shaped, usually not
73
enclosing the fruit.....................................................................4. S. bunburyanus (in part)

17. Leaf margin wavy, recurved...............................................................19. S. trengganuensis
Leaf margin plane..............................................................................................................18
18. Leaf midrib sunken above.................................................................................................19
Leaf midrib raised or flattened above................................................................................22
19. Leaves in close spirals, 4385 cm long, sessile or with very short petioles...........................
.........................................................................................................11. S. impressicostatus
Leaves well-spaced, shorter, petioles well-developed.......................................................20
20. Leaves velvety hairy below, margin pale whitish on drying.................17. S. rufovelutinus
Leaves sparsely hairy below, margin not so.......................................................................21
21. Leaves glaucous below.......................................................................9. S. glaucus (in part)
Leaves not glaucous below..........................................................6. S. cuneiformis (in part)
22. Leaves well-spaced. Inflorescences axillary................................................3. S. borneensis
Leaves in pseudowhorls. Inflorescences terminal.............................................................23
23. Leaves densely tomentose below, margin whitish. Fruit with cup-shaped hypocarp.............
.............................................................................................................12. S. kinabaluensis
Leaves sparsely, often rough-hairy below, margin not whitish. Fruit without
cup-shaped hypocarp..............................................................4. S. bunburyanus (in part)
1. Semecarpus angulatus Kochummen
(Latin, angulatus = angular; the inflorescence axes)
Sandakania 7 (1996) 94. Type: Fedilis & Sumbing SAN 91435, Borneo, Sabah, Kalabakan (holotype SAN).
Small tree to 6 m tall, 5 cm diameter. Twigs stout, c. 1.7 cm thick, grey-brown, rugose. Leaves in
close spirals, coriaceous; faintly glaucous and sparsely hairy below; papillae dense, distinct and
uniformly distributed on the lower leaf surface; lanceolate, 2831.5 x 10.512.5 cm, the length less
than 4 times the width; base attenuate, slightly asymmetric, margin wavy, apex acute; midrib raised
above, lateral veins 1115 pairs, raised below, faint above, intercostal veins laxly reticulate, raised
below, faint above; petioles 89 cm long, basal part swollen, concave above and rugose on drying.
Inflorescences terminal, in clusters, 1519 cm long, with few short branches, finely hairy, axes
angular. Flowers (only female seen) sessile; bracteoles triangular, hairy, c. 1 mm long; calyx lobes
deltoid, c. 1 mm high, c. 2 mm broad, hairy outside; petals imbricate, elliptic, c. 5 x 2.5 mm, glabrous,
with a number of black veins inside, apex blunt; disc prominent, c. 0.5 mm high, saucer-shaped with
entire rim, glabrous; staminodes very small; ovary broadly conical, c. 3 x 2 mm, densely hairy, styles
c. 1 mm long, hairy, stigmas peltate. Fruits unknown.
Distribution. Endemic to Borneo; so far known only from the type from Sabah.
74
2. Semecarpus angustifolius Kochummen

Sandakania 7 (1996) 94. Type: Zainudin 5060, Sabah, Telupid, Bukit Tawai Forest Reserve (holotype SAN).
Small tree; stem spiny. Twigs c. 1 cm thick, brown, shallowly fissured, glabrous. Leaves in
pseudowhorls, glabrous, faintly glaucous below; papillae distinct, uniformly distributed on lower leaf
surface; narrowly oblong, 5264 x 3.84.4 cm, the length more than 4 times the width; base obtuse,
apex acute; midrib raised above, lateral veins 3236 pairs, curving and joining near margin, visible on
both surfaces, intercostal veins reticulate, visible below, faint to invisible above; petioles to 1 cm long,
rugose. Inflorescences and flowers unknown. Fruits ellipsoid, c. 1 x 0.5 cm, glabrous, apex obtuse;
hypocarp glabrous or sparsely scaly, funnel-shaped, c. 3 mm long; fruit stalks c. 2 mm long.
Distribution. Endemic to Borneo; so far known only from the type specimen from Sabah.
Ecology. Lowland forest, at c. 200 m.
3. Semecarpus borneensis Merr.

(of Borneo)
J. Str. Br. Roy. As. Soc. 86 (1922) 323; Masamune l.c. 413; Ding Hou, Blumea 24 (1978) 35, FM 1, 8 (1978) 507. Type:
Agama 1061, Borneo, Sabah, Kudat , Rosop (holotype PNH, destroyed; isotype A).
Small tree to 12 m tall, 15 cm diameter. Young twigs tomentose. Leaves in well-spaced spirals,
glabrous above except the midrib, hairy below; papillae distinct, uniformly distributed on the lower
leaf surface, and veins; elliptic, oblong or oblanceolate, 720 x 1.58 cm, the length usually less than
4 times the width; base broadly rounded to cuneate, margin plane, apex acute; midrib flattened above,
lateral veins 1016 pairs, raised below, faint above, intercostal veins reticulate, distinct below, faint
above; petioles 12.5 cm long, lower part swollen and rugose on drying. Inflorescences axillary
panicles, to 20 cm long, hairy; bracts linear, 23 mm long. Flowers (only male seen) subsessile; buds
subglobose; calyx lobes triangular; petals hairy outside. Fruits (young) broadly ellipsoid, c. 7 x 6
mm, with obtuse apex, hypocarp obconical.
Distribution. Endemic to Borneo. Common in the Ranau district, with a few collections from Beluran
(Labuk Sugut district), Lamag (Kinabatangan district) and Marudi area in Sabah; not yet reported
from Sarawak.
Ecology. Hill to lower montane forest, including riverbanks, at 6001500 m.
Notes. SAN 69337 and SAN 92476, collected from riverbanks, have narrowly linear leaves, 1.52 cm
wide.
4. Semecarpus bunburyanus Gibbs

(Mr. Bunbury, probably an officer of the British North Borneo administration)
75
J. Linn. Soc. Bot. 42 (1914) 67; Merrill l.c. (1921) 351; Masamune l.c. 413; Ding Hou, FM 1, 8 (1978) 512. Type: Gibbs
3964, N Borneo, Sabah, Mt. Kinabalu (holotype BM). Synonym: S. scaberulus Merr., PEB (1929) 169, Masamune l.c. 414.
Small tree to 15 m tall, 20 cm diameter, often a treelet, sometimes unbranched; young stem spiny.
Twigs very stout to 2 cm thick or slender to 0.5 cm thick. Leaves in pseudowhorls or in well-spaced
spirals; upper surface glabrous, lower surface glabrous, sparsely rough-hairy or densely tomentose,
only sometimes glaucous; papillae distinct to obscure, uniformly distributed on the lower leaf
surface; obovate, oblong or oblanceolate, 15100 x 3.522 cm, the length less than 4 times the width;
base attenuate, margin plane, not whitish, apex acute or obtuse; midrib raised above, lateral veins 10
35 pairs, raised on both surfaces, intercostal veins reticulate, distinct below, visible above; petioles
absent or to 8 cm long. Inflorescences terminal or axillary panicles, to 35 cm long, with many side
branches, sparsely hairy, axes not angular; bracts and bracteoles of variable length and shape;
pedicels very short to 1.5 mm long. Flowers whitish, buds oblong; calyx lobes triangular, sparsely
hairy outside; petals valvate, elliptic or oblong, usually glabrous outside with several longitudinal
veins which are more distinct on the inner surface; stamens c. 2.5 mm long, anthers ovoid-oblong, 1
1.5 mm long, staminodes slightly smaller; disc in male flowers flat and often glabrous, in female
flowers cup-shaped, glabrous; ovary conical, hairy, styles to 1 mm long. Fruits subglobose, 0.72
x 0.61.5 cm, glabrous or sparsely hairy, apex rounded; hypocarp funnel-shaped or obconical or
cylindrical.
Distribution. Borneo and the Philippines. Common and widely distributed in Sabah, very uncommon
in Sarawak; also in Brunei and Kalimantan.
Notes. There is extremely wide variation in twig thickness, leaf arrangement, leaf size, pubescence
and petiole length. Even though there are a large number of collections of this species from Sabah,
mature fruits are lacking and only very few collections have female flowers.
5. Semecarpus calcicolus Kochummen

(Latin, calx = chalk or lime, colus = dweller; the limestone habitat)
Sandakania 7 (1996) 97. Type: Dewol & Rahman SAN 90206, Borneo, Sabah, Kota Belud, Mantanani Besar Island
(holotype SAN).
Small tree to 15 m tall, 20 cm diameter. Bark grey-white; inner bark pale. Sapwood white. Twigs
stout, c. 1.5 cm thick, lenticellate, hollow. Leaves in close spirals, glabrous on both surfaces; papillae
obscure on the lower leaf surface; oblanceolate, 2634 x 78.5 cm, the length less than 4 times the
width; base attenuate, apex attenuate-acuminate; midrib raised above, lateral veins 2830 pairs,
distinct below, faint above, intercostal veins finely reticulate, visible on both surfaces; petioles c. 1 cm
long, drying black and rugose. Inflorescences and flowers unknown. Infructescences extra axillary,
c. 12 cm long. Fruits broadly oblong, 2.12.5 x 22.2 cm, glabrous, green when fresh, apex blunt and
slightly depressed in the middle; hypocarp obconical, c. 0.7 cm high.
76
Distribution. Endemic to Borneo; so far known only from the type from Mantanani Besar Island in
Sabah.
Ecology. On limestone.
6. Semecarpus cuneiformis Blanco

(Latin, cuneiformis = wedge-shaped; the leaf base)
Fl. Filip. (1837) 220; Merrill, Spec. Blanc. (1918) 235; Ding Hou, FM 1, 8 (1978). Neotype (Merrill, 1918): Merrill Spec.
Blanc. 283, Philippines, Luzon, Manila (L). Synonym: S. lanceolatus Ridl., Kew Bull. (1933) 199, Masamune l.c. 413.
Small tree to 15 m tall, 20 cm diameter. Leaves spirally arranged, well-spaced; lower surface
distinctly or sparsely hairy to almost glabrous, not glaucous; papillae distinct and uniformly
distributed on lower leaf surface; oblong or oblanceolate, 9.528 x28.5 cm, the length less than 4
times the width; base cuneate, margin plane, not whitish on drying, not recurved, apex acute or obtuse
or emarginate; midrib sunken above, not pale yellow on drying, lateral veins 1518 pairs, raised
below, very faint to invisible above, intercostal veins reticulate, distinct below, very faint above;
petioles 0.52.5 cm long, not pale yellow on drying. Inflorescences usually terminal, to 30 cm long,
hairy. Flowers: buds subglobose; calyx lobes triangular; petals hairy outside with distinct longitudinal
veins; ovary densely hairy. Fruits subglobose, c. 7 mm diameter; hypocarp funnel-shaped.
Distribution. Borneo, Philippines, Sulawesi, Lesser Sunda Islands, Formosa. In Borneo, recorded
from the eastern and southern parts of Sabah; not yet reported from Sarawak; also in Brunei and
Kalimantan.
Ecology. Lowland forest below 100 m.
7. Semecarpus cupularis Kochummen

(Latin, cupularis = cup-shaped; the hypanthium of the fruit)
Sandakania 7 (1996) 97. Type: Ilias S. 41671, Borneo, Sarawak, Kapit (holotype SAR; isotypes K, L, SAN).
Small tree to 10 m tall, 10 cm diameter; stem with scattered spines. Twigs glabrous, stout, 12.2 cm
thick, yellowish grey, shallowly fissured. Leaves in close spirals; glabrous, drying to greenish grey
above, glaucous below; papillae uniformly distributed on the lower leaf surface; obovate, 2852 x 8
19 cm, the length less than 4 times the width; base attenuate, margin plane, apex acute; midrib raised
above, lateral veins c. 30 pairs, curving and joining near margin, raised on both surfaces, intercostal
veins laxly reticulate, visible on both surfaces; petioles c. 1.5 cm long, flattened above, yellowish on
drying. Inflorescences and flowers unknown. Infructescences terminal, more than 20 cm long, axes
stout, glabrous. Fruits green, ripening to orange, transversely oblong, 1.42 x 1.21.4 cm, glabrous,
apex blunt with shallow depression in the middle; hypocarp cup-shaped, enclosing more than threequarters of the fruit.
77
Distribution. Endemic to Borneo. Very uncommon, known from two collections, S. 41671, the type,
and S. 37203, from Kapit District, Sarawak.
Ecology. Lowland forest, at about 250 m.
8. Semecarpus euodiifolius Kochummen

(Latin, Euodia = a genus in the Rutaceae, folium = leaf; leaves resembling that of Euodia)
Sandakania 7 (1996) 100. Type: Mikil SAN 28100, Borneo, Sabah, Beaufort (holotype KEP; isotype SAN).
Medium-sized tree to 23 m tall, 20 cm diameter. Bark brownish, smooth; inner bark pale yellow.
Twigs pale whitish, glabrous, 23 mm thick. Leaves spirally arranged, well-spaced, thin-coriaceous;
glabrous, slightly glaucous below; papillae uniformly distributed on the lower leaf surface; ellipticoblong, 7.519.5 x 3.56.5 cm; base attenuate, margin not recurved, apex acute with sharp tip; midrib
sunken above, not pale yellow on drying, lateral veins 1114 pairs, looping and joining near margin,
faint on both surfaces, intercostal veins reticulate, visible on both surfaces; petioles 1.25 cm long,
swollen near the base, not pale yellow on drying. Inflorescences in terminal panicles or in clusters, 9
16 cm long, densely hairy; sometimes male and female flowers on the same inflorescence; bracts
linear-lanceolate, c. 1.5 mm long. Flowers: hypanthium funnel-shaped in male flower, cup-shaped
and longer and 46 mm wide in female flower, densely hairy outside; calyx lobes triangular, c. 0.5
mm long; petals oblong, glabrous, c. 2.5 x 1 mm, with 810 distinct black veins, apex acute; stamens
c. 1.5 mm long; pistillodes densely hairy; staminodes present; ovary conical, densely hairy. Fruits
globose, 1.21.5 x 1.21.4 cm, densely hairy, apex slightly depressed; hypocarp cup-shaped,
enclosing almost half of the fruit.
Distribution. Endemic to Borneo. Uncommon, recorded only from Beaufort and Kuala Penyu
districts in Sabah (SAN 28072, SAN 28100 (type), SAN 58436, and SAN 102258).
Ecology. Lowland forest. Flowering from February to May, fruiting in February.
9. Semecarpus glaucus Engl.
Fig. 15.
(Latin, glaucus = with a pale bloom; the lower leaf surface)

In A. DC., Mon. Phan. 4 (1883) 478; Merrill l.c. (1921) 352; Masamune l.c. 413; Ding Hou, FM 1, 8 (1978) 513; Anderson
l.c. 138; Whitmore, Tantra & Sutisna l.c. 21. Syntypes: Beccari PB 2875 & 3318, Borneo, Sarawak (FI).
Small to medium-sized tree to 25 m tall, 20 cm diameter. Bark grey-brown, smooth. Twigs grey or
grey-brown, densely brown short-hairy when young. Leaves spirally arranged, well-spaced; lower
surface distinctly or slightly glaucous, uniformly covered with distinct papillae, upper surface
glabrous, except for the midrib and lateral veins which are tomentose; elliptic to oblong or
oblanceolate, 726.5 x 2.513 cm, the length less than 4 times the width; base cuneate, margin plane,
not recurved, not pale whitish, apex acute; midrib usually sunken above, not pale yellow on drying,
lateral veins 717 pairs, looping and joining near margin, raised below, visible above, intercostal
veins reticulate, visible on both surfaces; petioles
78
Fig. 15. Semecarpus glaucus. A, flowering leafy twig; B, part of inflorescence; C, male flower; D, female flower; E, fruit.
(AB from S. 41896, CD from FM 1, 8 (1978) 500, f. 47 c and d, E from S. 34664.)
79
0.54.8 cm long, not pale yellow on drying. Inflorescences terminal panicles, 1135 cm long, hairy;
bracts linear, 1.53 mm long. Flowers: buds oblong or ellipsoid; pedicels to 2.5 mm long; calyx lobes
triangular; petals white, hairy outside with several longitudinal veins; ovary velvety hairy. Fruits
(young) broadly ellipsoid, c. 1 x 0.7 cm, hairy, glabrescent, with almost rounded apex; hypocarp
discoid.
Distribution. Endemic to Borneo. Widely distributed in Sabah and Sarawak but nowhere abundant;
also in Kalimantan.
Ecology. Lowland forest to 450 m, in a variety of habitats including peat swamp, fresh-water swamp,
and limestone.
Note. Very close to S. rufovelutinus from which it can be distinguished by the reddish brown leaves
which are sparsely hairy and glaucous below.
10. Semecarpus heterophyllus Blume

(Greek, hetero = different, phullon = leaves; the different shapes of the leaf)
Mus. Bot. Lugd. Bat. 1 (1850) 187; Ridley, FMP 5 (1925) 302; Backer & Bakhuizen f. l.c. 154; Ding Hou, FM 1, 8 (1978)
512; Anderson l.c. 139; Whitmore, Tantra & Sutisna l.c. 21. Type: Blume, s.n., Java (holotype L).
Medium-sized tree to 22 m tall, 60 cm diameter; young stem not spiny. Bark grey-brown, smooth.
Leaves spirally arranged, well-spaced, variable in shape, coriaceous; glabrous, slightly glaucous
below; papillae distinct and uniformly distributed on the lower leaf surface; elliptic, obovate or
oblanceolate, 1122 x 58 cm; base broadly attenuate, apex acute; midrib raised above, lateral veins
1020 pairs, raised on both surfaces, intercostal veins reticulate, raised below, visible above; petioles
0.56 cm long. Inflorescences terminal panicles, rarely in the axils of upper leaves. Flowers sessile;
calyx lobes triangular; petals imbricate, hairy outside, with 9 distinct veins. Fruits subglobose, 1.72
x 1.52 cm, glaucous, apex rounded; hypocarp discoid.
Distribution. Sumatra, Peninsular Malaysia, Borneo, Java, Lesser Sunda Islands. In Sabah, recorded
only from Mt. Kinabalu (RSNB 4264 and RSNB 7095); not yet reported from Sarawak; also in
Kalimantan.
Ecology. Lower montane forest, at 13501800 m.
11. Semecarpus impressicostatus Kochummen
(Latin, impressus = sunken, costus = vein; the sunken midrib on the upper leaf surface)
Sandakania 7 (1996) 102. Type: Kulip & Goh SAN 137045, Borneo, Sabah, Tongod (holotype SAN).
Small unbranched tree to 20 m tall, 10 cm diameter. Twigs very stout, c. 2 cm thick, rugose. Leaves
in close spirals, coriaceous; glabrous above, rough hairy below; papillae uniformly distributed on the
lower leaf surface; oblanceolate, 4385 x 1122 cm, the length less than 4 times the width; base
attenuate, margin plane, pale white on drying, apex rounded, bluntly pointed or emarginate; midrib
sunken above, prominent below, to 8 mm broad, lateral veins
80
3050 pairs, raised below, faint above, looping and joining near margin, intercostal veins reticulate,
raised below, visible above; petioles absent or to 5 mm long. Inflorescences terminal, 7592 cm long,
axes stout, to 7 mm thick, rusty brown-hairy; bracts lanceolate, c. 1.5 mm long, woolly; bracteoles
lanceolate, woolly, c. 0.5 mm long. Flowers (only male seen): pedicels c. 1 mm long, pale yellow in
bud, densely hairy; calyx lobes c. 0.5 mm long, triangular with blunt apex; petals oblong, c. 2 x 1 mm,
glabrous on both surfaces, with distinct black veins inside; stamen filaments c. 2 mm long; disc
densely hairy; pistillodes absent. Fruits obliquely oblong, c. 1.7 x 1.5 cm, with blunt apex, sparsely
hairy; hypocarp c. 5 mm long, slightly obconical.
Distribution. Endemic to Borneo. In Sabah, known only from Beluran (Pensiangan district),
Sandakan district and Tongod (Kinabatangan district) (SAN 137045, SAN 79352, SAN 131008); not
yet recorded in Sarawak.
12. Semecarpus kinabaluensis Kochummen

(of Mt. Kinabalu)
Sandakania 7 (1996) 102. Type: Aban & Saikeh SAN 74108, Borneo, Sabah, Kinabalu Park (holotype SAN; isotypes K, L).
Small tree to about 10 m tall; stem spiny. Twigs 0.51 cm thick, grey-brown, shallowly fissured,
glabrous. Leaves in pseudowhorls; glabrous above, densely tomentose below; papillae distinct and
uniformly distributed on the lower leaf surface; obovate, 2237.5 x 8.515.5 cm, the length less than
4 times the width; base attenuate, margin plane, pale whitish, apex attenuate and sharp-pointed;
midrib raised above, lateral veins 1828 pairs, raised below, visible above, intercostal veins reticulate,
raised below, visible above; petioles 0.53.5 cm long, rugose on drying, swollen at base.
Inflorescences terminal panicles, to 33 cm long, with many side-branches, hairy; bracts broadly
triangular, c. 1 x 1 mm, hairy; bracteoles lanceolate, c. 1.5 mm long, hairy. Flowers (only male seen)
sessile; calyx lobes triangular, c. 0.5 mm long; petals yellow when fresh, imbricate, glabrous, outer
surface rugose, oblong, c. 2 x 0.5 mm; stamens c. 2 mm long, anthers oblong, c. 1 mm long; disc 5lobed, glabrous; pistillodes absent. Fruits subglobose, c. 1.2 x 1.3 cm, glabrous, apex with short
point; hypocarp cup-shaped, 34 mm high.
Distribution. Endemic to Borneo. So far recorded only from the Kinabalu Park in Sabah, where it
appears to be common.
Ecology. Lower montane forest, at 12001800 m.
Notes. In addition to the type, the following collections belong to this species: SAN 56516, SAN
76483; RSNB 4167, RSNB 4339, and Kokawa & Hotta 4392.
13. Semecarpus lineatus Kosterm.

(Latin, lineatus = marked with fine lines; the twigs)
81
Blumea 33 (1988) 348. Type: Binideh SAN 55161, Borneo, Sabah, Lahad Datu (holotype L; isotype SAN).
Small tree to 5 m tall, 10 cm diameter. Twigs slender, shallowly longitudinally furrowed. Leaves in
pseudowhorls, glabrous; papillae obscure; linear, 2043 x 12 cm, the length more than 4 times the
width; base attenuate, apex acuminate; midrib raised above, lateral veins 2023 pairs, with short
intermediate veins, intercostal veins reticulate, lax; petioles 11.5 cm long, base slightly swollen.
Inflorescences panicles, terminal or in the axils of upper leaves, 1520 cm long. Flowers light green;
sepals triangular, 0.250.5 mm long; petals c. 2.5 mm long. Fruits unknown.
Distribution. Endemic to Borneo; so far known only from Mt. Silam in Lahad Datu area, Sabah.
14. Semecarpus minutipetalus Kochummen

(Latin, minutus = very small, petalus = petal)
Sandakania 7 (1996) 105. Type: Ilias S. 39169, Borneo, Sarawak, Miri (holotype KEP; isotypes K, L, SAN, SAR).
Small tree to 12 m tall, 12 cm diameter. Twigs pale whitish, glabrous. Leaves spirally arranged, wellspaced; glabrous, faintly glaucous below; papillae distinct on the lower leaf surface except on the
midribs and veins; narrowly obovate to oblong, 18.531.5 x 5.510 cm; base attenuate, margin pale
whitish, recurved, apex acute; midrib sunken above, on drying pale yellow below, lateral veins 2025
pairs, looping and joining near margin, distinct below, faint to invisible above, intercostal veins
reticulate, faintly visible on both surfaces; petioles 3.56 cm long, pale yellow on drying, swollen at
base, narrowly grooved above. Inflorescences short panicles, arising from the axils of upper leaves,
to 8 cm long, densely hairy; bracts linear-lanceolate, c. 1 cm long, densely hairy; bracteoles to 2 mm
long. Flowers (only immature male flowers seen): calyx lobes c. 2 mm long, oblong, hairy outside
with glabrous margins and with 3 black ridges inside; petals narrowly oblong, apex obtuse, very much
smaller than calyx lobes, c. 0.2 mm long, with a single ridge on the inside; stamens 5; pistillodes
absent. Fruits unknown.
Distribution. Endemic to Borneo; so far recorded only from the type specimen from Miri in Sarawak.
Ecology. Lowland forest, at about 120 m.
15. Semecarpus paucinervius Merr.

(Latin, pauci = few; nervis = veins; the leaf)
Philip. J. Sc. 7 (1912) Bot. 286; Ding Hou, Blumea 24 (1978) 36, FM 1, 8 (1978) 510; Whitmore, Tantra & Sutisna l.c. 21.
Type: Foxworthy Bur. Sci. 750, Borneo, Sabah, Tawau (isotype BO).
82
Small tree to 20 m tall, 60 cm diameter. Twigs grey-brown, glabrous. Leaves well-spaced, spirally
arranged, coriaceous, glabrous; papillae uniformly distributed on the lower leaf surface; obovate or
oblong, 512.5 x 25.5 cm; base attenuate, apex rounded; midrib flattened above, lateral veins 510
pairs, raised below, faint above, intercostal veins reticulate, distinct below, faint above; petioles 0.5
1.7 cm long. Inflorescences panicles, often terminal; bracts lanceolate. Flowers: buds globose; calyx
lobes triangular; petals sparsely hairy outside, with several distinct veins; ovary hairy. Fruits
maturing red, subglobose, c. 8 mm diameter, apex rounded; hypocarp funnel-shaped.
Distribution. Borneo, the Philippines. In Borneo, recorded only from Kota Belud and Kudat districts
in Sabah.
Ecology. Lowlands including seashores and riverbanks.
16. Semecarpus pulvinatus Kochummen

(Latin, pulvinatus = cushion-shaped; the swollen petiole base)
Sandakania 7 (1996) 105. Type: Dewol SAN 124556, Borneo, Sabah, Kota Kinabatangan (holotype SAN).
Small tree to 8 m tall, 20 cm diameter. Bark grey, smooth. Twigs c. 7 mm thick, brownish, shallowly
fissured. Leaves in pseudowhorls; glabrous on both surfaces, faintly glaucous below; papillae
obscure on the lower leaf surface; narrowly elliptic to oblanceolate, 2030 x 3.56.5 cm, the length
more than 4 times the width; base attenuate, apex acute; midrib raised above, lateral veins c. 35 pairs,
looping and joining near margin, raised below, visible above, intercostal veins reticulate, faintly
visible on both surfaces; petioles 4.56 cm long, lower 11.5 cm prominently swollen. Inflorescences
terminal panicles, c. 45 cm long, with many side branches, sparsely hairy; bracts lanceolate, c. 0.5 mm
long, hairy. Flowers (only male seen) sessile; calyx lobes triangular, c. 0.5 mm long, hairy outside;
petals valvate, glabrous on both surfaces, oblong, c. 3 x1.5 mm, with acute apex and 67 distinct
veins; stamen filaments c. 2 mm long, anthers oblong, c. 1.5 mm long; disc pulvinate, glabrous;
pistillodes absent. Fruits globose, 1.52 x 11.2 cm, glabrous apex emarginate; hypocarp saucershaped.
Distribution. Endemic to Borneo; known only from 2 collections from Kinabatangan and Tawau
districts in Sabah (SAN 89703, and SAN 124556 respectively).
Ecology. Lowland forest. Flowering in February and fruiting in September.
17. Semecarpus rufovelutinus Ridl.

(Latin, rufus = reddish, velutinus = velvety; indumentum on the lower leaf surface)
Kew. Bull. (1933) 199; Masamune l.c. 414; Ding Hou, FM 1, 8 (1978) 513; Anderson l.c. 139; Whitmore, Tantra & Sutisna
l.c. 22. Type: Hose 654, Borneo, Sarawak, Miri (holotype K; isotypes BM, L).
Synonym: Melanocommia borneensis Ridl. l.c. (1933) 198, Masamune l.c. 411.
83
Small tree to 10 m tall, 10 cm diameter. Bark grey, smooth; inner bark brownish. Sapwood whitish.
Twigs grey-brown, soft-hairy when young. Leaves in spirals, well-spaced; glabrous above, reddish
velvety hairy below, drying to grey-brown; papillae distinct, uniformly distributed on the lower leaf
surface; elliptic, oblong or obovate, 1032 x 611 cm, the length less than 4 times the width; base
attenuate, margin plane, pale whitish on drying, apex acute; midrib sunken above, lateral veins 1012
pairs, raised below, visible above, intercostal veins reticulate, raised below, faintly visible above;
petioles 0.53.5 cm long, hairy. Inflorescences often terminal panicles, to 34 cm long, velvety hairy;
bracts lanceolate, 14 mm long. Flowers: buds oblong; pedicels to 1 mm long; calyx lobes triangular;
petals hairy outside, with distinct longitudinal veins; ovary subglobose, velvety. Fruits subglobose,
11.5 x 1.51.7 cm, velvety hairy, apex rounded; hypocarp discoid or cupular.
Distribution. Peninsular Malaysia, Borneo. Common and widely distributed in Sabah, uncommon in
Sarawak; also in Kalimantan.
Ecology. Lowland and hill forests, including limestone, to 500 m.
Notes. Often confused with S. glaucus but it is easily distinguished by the reddish velvety hairy leaves
which are not glaucous below.
18. Semecarpus sandakanus Kochummen

(of Sandakan in Sabah)
Sandakania 7 (1996) 108. Type: Saikeh SAN 87801, Borneo, Sabah, Sandakan (holotype KEP; isotype SAN).
Small tree to 5 m tall. Twigs slender, grey, glabrous. Leaves in close spirals, thin-coriaceous;
glabrous, glaucous below; papillae distinct and in groups on the lower leaf surface; elliptic, 2630.5
x 810 cm; base attenuate, apex acuminate, acumen 22.5 cm long; midrib raised above, lateral veins
2025 pairs, looping near margin, distinct below, faintly raised above, intercostal veins reticulate,
faintly visible below, invisible above; petioles 23 cm long, channelled above, yellowish on drying,
not swollen at base. Inflorescences slender terminal panicles, to 10 cm long, with few side-branches,
sparsely hairy; bracts linear, c. 4 mm long, hairy; bracteoles linear-lanceolate, c. 1.5 mm long, hairy.
Flowers (only male seen): calyx lobes triangular; petals imbricate, oblong, c. 2.5 x 1 mm, glabrous,
with distinct veins; stamens c. 1 mm long; disc pulvinate, c. 1 mm diameter; pistillodes absent. Fruits
unknown.
Distribution. Endemic to Borneo; so far known only from the type from Sandakan in Sabah.
19. Semecarpus trengganuensis Kochummen

(of Trengganu, a state in Peninsular Malaysia)
Gard. Bull. Sing. 36 (1984) 196, l.c. (1989) 52. Type: Chan FRI 16876, Peninsular Malaysia, Bukit Bauk FR, Trengganu
(holotype KEP; isotypes A, K, L, SING).
84
Shrub or small tree to 5 m tall. Twigs grey or grey-brown. Leaves in spirals, well-spaced; rough hairy
below; papillae uniformly distributed on the lower leaf surface; elliptic to obovate, 1133 x 311 cm,
the length less than 4 times the width; base attenuate, margin wavy, recurved, apex acute; midrib
flattened or faintly sunken above, lateral veins 1317 pairs, looping and joining near margin, raised
below, faint above, intercostal veins reticulate, raised below, faint above; petioles 1.53 cm long,
densely brown short-hairy. Inflorescences terminal and axillary panicles, up to 12 cm long, slender,
with few short branches, hairy; bracts lanceolate, hairy. Flowers: buds (male) oblong; calyx lobes
triangular. Fruits (young) oblong to subglobose, c. 10 x 8 mm, hairy; hypocarp funnel-shaped, hairy.
Distribution. Peninsular Malaysia, Borneo. Common in Sabah, uncommon in Sarawak.
Note. This is the first record of its occurrence in Borneo.
16. SOLENOCARPUS Wight & Arn.

(Greek, solenos = tube, karpos = fruit)
Prod. Fl. Pen. Ind. Or. 1 (1834) 171; Kostermans, Found. Useful Pl. Asia 1 (1991) 69. Synonym: Spondias (non L.) Ding
Hou, p.p. (quoad S. indica & S. philippinensis), FM 1, 8 (1978) 485.
Trees or lianas. Leaves in close spirals, pinnately compound with opposite or subopposite leaflets and
terminal leaflets. Inflorescences terminal and/or axillary panicles. Flowers bisexual, 5-merous; calyx
5-lobed; petals 5, valvate in bud, reflexed; disc saucer-shaped; stamens 10, filaments free, anthers
versatile, dorsifixed, 2-locular, longitudinally dehiscent; ovary superior, 1-carpellate, 1-locular, style
distinct, stigmas 2-lobed, ovule one, pendulous. Fruits with thin mesocarp; endocarp smooth and free
from fibers.
Distribution. Two species (S. indica and S. philippinensis); S India to New Guinea. One species (S.
philippinensis) in Sabah and Sarawak.
Ecology. Lowland and hill forests, also in mangroves.
Taxonomy. Airy Shaw & Forman (Kew Bull. 21 (1967) 2) reduced Solenocarpus Wight & Arn.,
along with a few other genera, to Spondias L., characterised by simple, pinnately or bipinnately
compound leaves, a 1- or 45-carpellate ovary, and variable size and structure of the endocarp.
Kostermans (l.c. 1991) recognised 6 distinct genera, including Solenocarpus, instead of one as
delimited by Airy Shaw & Forman. In recognising the fact that the number of carpels in each ovary is
an important generic character in the Anacardiaceae, I concur with Kostermans in treating
Solenocarpus as a separate genus from Spondias. An additional character which can be used to
distinguish Solenocarpus from Spondias is the smooth fruit endocarp in the former.
Solenocarpus philippinensis (Elmer) Kosterm.
Fig. 16.
(of the Philippines)
85
Fig. 16. Solenocarpus philippinensis. A, fruiting leafy twig; B, part of inflorescence; C, flower with one petal removed. (A
from SAN 85111, BC from S. 18265.)
86
New & Crit. Mal. Pl. 3 (1955) 1, l.c. (1991) 70. Basionym: Pegia philippinensis Elmer, Leafl. Philip.
Bot. 8 (1919) 3100. Type: Elmer 13467, Philippines, Mindanao, Agusan Prov., Cabadbaran, Mt.
Urdaneta (K, L). Synonyms: Phlebochiton philippinensis (Elmer) Merr., En. Philip. Fl. Pl. 2 (1923) 472;
Spondias philippinensis (Elmer) Airy Shaw & Forman, Kew Bull. 21 (1967) 15; Pentaspadon telianthera
Ridl., Kew Bull. (1933) 199, Masamune l.c. 413.
Tree to 45 m tall, shrub, or large woody climber, or epiphytic shrub. Leaves compound, each
with 14 pairs of leaflets; rachis minutely hairy to sparsely hairy. Leaflets sparsely hairy on
both surfaces, to subglabrous; elliptic, ovate or oblong, 613.5 3.56 cm; base obtuse, apex
acute; lateral veins 79 pairs, visible on both surfaces, intercostal veins reticulate, distinct on
both surfaces; petiolules c. 3 mm long. Inflorescences lax, to 25 cm long, appearing before
the new leaves or accompanied by young leaves; bracts lanceolate, 11.5 mm long. Flowers
white; pedicels 1.52 mm long; calyx lobes triangular, c. 0.5 mm; petals valvate in bud,
elliptic-lanceolate to obovate-oblong, 2.53 1 mm, patent; stamens 1.53.5 mm, anthers
ellipsoid, c. 1 mm; disc c. 1 mm diameter; styles to 1 mm long. Fruits 1-locular, oblong,
slightly curved, c. 1 0.5 cm, yellowish when ripe; style scar lateral; endocarp woody, smooth
with faintly raised reticulation at the apical end, without envelope of fibres.
Distribution. Sumatra, Borneo, Philippines, New Guinea. Common and widely distributed in
Sabah and Sarawak.
17. SWINTONIA Griff.
(G. Swinton, a government officer in Bengal, c. 1840)

merpauh (Malay), pitoh (Iban)
Proc. Linn. Soc. Lond. 1 (1846) 283; Merrill, EB (1921) 349; Ridley, FMP 1 (1922) 532; Masamune,
EPB (1942) 414; Browne, FTSB (1955) 53; Burgess, TBS (1966) 33; Ding Hou, Blumea 24 (1978) 38,
FM 1, 8 (1978) 440; Anderson, CLTS (1980) 139; Wong, DMT (1982) 233; Kochummen, TFM 4 (1989)
44; Whitmore, Tantra & Sutisna, CLK 1 (1989) 22; Ng, Mal. For. Rec. 34, 1 (1991) 19.
Medium-sized to large emergent trees; bole with steep buttresses. Bark grey-brown, reddish
brown or dark brown, smooth to minutely dippled or scaly; inner bark yellowish or pinkish
with creamy to yellowish latex in droplets which darkens on exposure. Sapwood pale. Leaves
spirally arranged, well-spaced or in close clusters and appearing as in pseudowhorls, simple,
often glaucous below; petioles swollen at base. Inflorescences axillary and terminal panicles.
Flowers male and bisexual or bisexual only; calyx 5-lobed, floral axis between calyx and
stamens elongated or (in Sabah and Sarawak) not; petals 5, imbricate in bud, persistent,
usually enlarged and wing-like in fruit; disc extrastaminal, consisting of 5 gland-like lobes,
confluent with the base of filaments or alternating with them; stamens 5, filaments filiform or
subulate, glabrous, anthers dorsifixed; disc lobed, lobes confluent with filament base; ovary
superior, 1-carpellate, 1-locular, style distinct, cylindric, stigmas minute, capitate; pistillodes
very small, hairy. Fruits 1-locular, subtended by the persistent petals which are usually very
much enlarged and wing-like. Seeds with the testa fused to the endocarp; embryo straight,
cotyledons free, plano-convex; germination hypogeal or semi-hypogeal, in S. schwenkii seeds
87
germinate within 542 days after sowing, seedling leaves simple, entire, the first two opposite, subsequent
ones spirally arranged.
Distribution. About 13 species; Andamans, Myanmar, Thailand, Cambodia, Laos, Vietnam, Sumatra,
Peninsular Malaysia, Borneo and the Philippines; 6 species in Sabah and Sarawak.
Ecology. Lowlands, including fresh-water and peat swamps, to hill and lower montane forests, to 1300 m.
Uses. The timber, which is medium hardwood (density 640-880 kg/nr' air-dried) and known by the
standard Malaysian name merpauh, is suitable for light construction indoors, interior finishing, panelling,
partitioning and flooring. It has been successfully tried for rotary-cut veneers.
Key to Swintonia species

1. Leaves in close clusters (in pseudowhorls)............................................................................... 2
Leaves well-spaced ..................................................................................................................5
2. Leaf midrib sunken above; lateral veins 20-25 pairs..................................... 5. S. sarawakana
Leaf midrib raised or flattened above; lateral veins to 20 pairs ................................................ 3
3. Leaf midrib sharply keeled beneath. Fruits ovoid-oblong or ellipsoid...............6. S. schwenkii
Leaf midrib not keeled beneath. Fruits globose....................................................................... 4
4. Enlarged petals in fruits shorter than the fruits. Petals sparsely hairy on both surfaces
....................................................................................................................... 4. S. minutalata
Enlarged petals in fruits much longer than the fruits. Petals glabrous except for sparse
hairs on the apical parts ...................................................................................2. S. foxworthyi
5. Leaves distinctly glaucous below, without domatia or pustules ............................. 3. S. glauca
Leaves faintly glaucous below, often with domatia in the axils of lateral veins or with
pustules on the lower surface.................................................................................. 1. S. acuta
1. Swintonia acuta Engl.
Fig. 17A-D, I.
(Latin, acutus - pointed; the leaf apex)
Bot. Jahrb. 1 (1880) 44; Merrill I.e. (1921) 349; Masamune I.e. 414; Smythies I.e. 13; Burgess I.e. 33; Ding Hou, FM 1, 8 (1978)
444; Anderson I.e. 139; Whitmore, Tantra & Sutisna I.e. 22. Type: Beccari PB 2753, Borneo (holotype FI; isotype K).
Synonym: S. schwenkii var. beccarii Engl. I.e. 44.
Medium-sized to large tree to 30-40 m tall, 40-70 m diameter; bole with buttresses to 3 m high. Bark
grey-brown to blackish, smooth to shallowly fissured; inner bark pinkish with colourless exudate.
Sapwood pale. Twigs black, slender, 2-3 mm thick. Leaves well-spaced, thin-coriaceous; faintly
glaucous below, often with dome-like domatia in the axils of lateral veins 'or small circular pustules all
over the lower surface; elliptic, lanceolate or oblong, 7-22
88
Fig. 17. Swintonia acuta (AD and I); S. minutalata (EH). A, flowering leafy twig; B, part of inflorescence; C, male
flower; D, female flower; E, male flower; F, female flower; G, young fruit; H and I, mature fruits. (AB from S. 45131, CF
and H from FM 1, 8 (1978) 441, f. 13, G from S. 40590, I from SFN 35665.)
89
TREEFLORAOFSABAHANDSARAWAKVOL.2(1996)
x 2-7 cm; base attenuate, apex acute; midrib raised above, lateral veins 10-21 pairs, distinct on both
surfaces, intercostal veins reticulate, very faint on both surfaces; petioles 1.5-5 cm long, slightly
swollen at base. Inflorescences paniculate, to 27 cm long, sparsely hairy; pedicels to 2 mm long.
Flowers white; calyx divided almost to the base; petals sparsely hairy at the apical part on both
surfaces; ovary globose or subglobose. Fruits ellipsoid, 1-2 x 0.7-1.7 cm, enlarged petals reddish when
fresh, 4-6 x 0.7-1.5 cm, base narrowed.
Vernacular name. Sarawakpitoh ayer (Iban).
Distribution. Borneo and the Philippines. Widely distributed in Sarawak, less common in Sabah; also in
Ecology. Lowland forest, rarely to 750 m, mainly by streams.
2. Swintonia foxworthyi Elmer
(F.W. Foxworthy, 1877-1950, American botanist who served in Manila and Kepong)
Leafl. Philip. Bot. 5 (1913) 1751; Ding Hou, FM 1, 8 (1978) 443; Whitmore, Tantra & Sutisna I.e. 22. Syntypes: Elmer 12982
& 13155, Philippines, Palawan, Puerto Princesa (BO, L).
Tree 30-40 m high, to 50 cm diameter; bole with buttresses to 1.5 m high. Bark reddish brown or dark
brown, smooth to scaly; inner bark with creamy latex which soon becoming black on exposure. Twigs
slender, 3-1 mm thick, grey-brown. Leaves in close clusters, faintly glaucous below; elliptic to narrowly
obovate, 6-13.5 x 2-5.5 cm; base attenuate, sometimes distinctly asymmetric, apex acute, rarely obtuse;
midrib flattened above, not keeled below, lateral veins 8-17 pairs, visible below, faint to invisible above,
intercostal veins reticulate, faintly visible on both surfaces; petioles 2-4 cm long, swollen at base.
Inflorescences paniculate, to 19 cm long. Flowers white; calyx divided almost to the base, lobes broadly
ovate; petals glabrous except for sparse hairs on both surfaces at apical parts; ovary subglobose. Fruits
globose or subglobose, 1.3-1.7 cm diameter; enlarged petals elliptic or oblanceolate, 5.5-8 x 1.5-2.5 cm,
with obtuse base, much longer than the fruits.
Distribution. Sumatra, Borneo, and the Philippines. Very uncommon in Sabah, but more widely
distributed in Sarawak; also in Brunei and Kalimantan.
Ecology. Lowland to upper hill mixed dipterocarp forest, to 1100 m, mainly on ridges.
3. Swintonia glauca Engl.
(Latin, glaucus = with a pale bloom; the lower leaf surface)

l.c. 44; Merrill I.e. (1921) 349; Masamune I.e. 414; Ding Hou, FM 1, 8 (1978) 442; Anderson I.e. 139; Whitmore, Tantra &
Sutisna l.c. 22. Type: Beccari PB 366, Borneo (holotype FI).
Tree to 33 m tall, 65 cm diameter; bole with buttresses to 1.5 m high. Bark grey-brown, scaly or dippled.
Twigs slender, c. 3 mm thick. Leaves well-spaced, without domatia or pustules, distinctly glaucous
below; oblong, elliptic or lanceolate, 6-15 x 3-6 cm; base broadly obtuse,
90
margin faintly recurved, apex acute; midrib raised above, lateral veins 8-16 pairs, with few short
intermediate veins between each pair, very faint on both surfaces, intercostal veins reticulate, visible
above; petioles 2.5-7 cm long. Inflorescences paniculate, to 30 cm long, hairy. Flowers white; calyx
divided almost to the base, lobes oblong or elliptic; petals densely hairy on both surfaces; ovary
subglobose. Fruits ellipsoid, 1.7-2.5 x 0.7-1.5 cm; enlarged petals 5.5-6.5 x 1.0-2.5 cm.
Vernacular name. Sarawakpitohpaya (Iban).
Distribution. Sumatra, Borneo. Not yet reported from Sabah, in Sarawak widely distributed; also in
Ecology. In lowland fresh-water, peat swamp, and kerangas forests.
4. Swintonia minutalata Ding Hou
(Latin, minutus = very small, alatum = wing; the small fruit wings)
Fig. 17E-H.
Blumea 24 (1978) 38, FM 1, 8 (1978) 443; Whitmore, Tantra & Sutisna I.e. 22. Type: Rosli S. 14966, Sarawak, Semengoh
Arboretum (holotype L; isotype K).
Tree to 35 m tall, 50 cm diameter; bole with buttresses to 1 m high. Bark smooth. Twigs grey-brown, c. 1
cm thick. Leaves in close clusters, not or slightly glaucous below; elliptic to oblong or lanceolate, 8.522.5 x 2-6.5 cm; base cuneate, apex acute; midrib raised above, not keeled below, lateral veins 11-20
pairs, very faintly visible on both surfaces, intercostal veins invisible or very faintly visible on both
surfaces; petioles 2-6.5 cm long, strongly swollen at base. Inflorescences 22-26 cm long, hairy panicles;
calyx divided almost to the base, lobes suborbicular; petals sparsely hairy on both surfaces; ovary
globose. Fruits globose, 1-1.5 cm diameter; enlarged petals dark red when fresh, elliptic to oblong, 0.71.2 cm long, very much shorter than the fruit.
Distribution. Endemic to Borneo. In Sarawak, known from Semengoh Arboretum and Bako National
Park; in Sabah, recorded from Beluran (Pensiangan district) Nabawan (Keningau district), Sandakan and
Tawau districts; also in Kalimantan.
Ecology. Lowland to hill mixed dipetrocarp forest, to 800 m. Flowering in January-April and fruiting in
June-November.
5. Swintonia sarawakana Kochummen

(of Sarawak)
Sandakania 7 (1996) 111. Type: Othman S. 41497, Borneo, Sarawak, Kapit (holotype SAR; isotypes K, KEP, L, P, SAN).
Medium-sized tree to 30 m tall, 50 cm diameter. Bark reddish brown, smooth. Twigs brownish, c. 5
mm thick. Leaves in close clusters, thin-coriaceous; elliptic, 6.5-11.5 x 2.5-5 cm; base attenuate, apex
acute; midrib sunken above, not keeled below, lateral veins 20-25 pairs, raised and distinct below, flat
and seemingly sunken on the upper surface, intercostal
91
veins reticulate, very faint below, sunken above; petioles slender, 4-4.5 cm long.
Inflorescences terminal panicles, to 23 cm long, with a number of long branches, minutely
hairy. Flowers (only male seen) pale green; bracts ovate, c. 1 mm long; pedicels c. 0.5 mm long;
calyx subglabrous, divided to more than half of its length, lobes triangular with an acute tip;
petals obovate, c. 2.5 x 1 mm, densely hairy on both surfaces, base attenuate, apex obtuse;
filaments dilated at base; pistillodes glabrous. Fruits unknown.
Distribution. Endemic to Sarawak; so far known from the type only from Bukit Entuboh, Ulu
Balleh, Kapit.
Ecology. Upper hill to lower montane forest, at c. 1300 m.
6. Swintonia schwenkii (Teijsm. & Binn.) Teijsm. & Binn. ex Hook.f. (Heinrich
Schwenk, 1809-1856, German soldier and plant collector)
Fl. Br. Ind. 2 (1876) 26; Merrill I.e. (1921) 350; Ridley I.e. (1922) 533; Masamune I.e. 414; Ding Hou, Blumea 24 (1978) 39,
FM 1, 8 (1978) 443; Smythies I.e. 13; Burgess I.e. 33; Anderson I.e. 139; Kochummen I.e. 57; Whitmore, Tantra & Sutisna
I.e. 23. Basionym: Anauxanopetalum schwenkii Teijsm. & Binn. in Miquel, J. Bot. Neerl. 1 (1861) 368.
Tree to 36 m tall, 40 cm diameter; bole with buttresses to 3 m high. Bark grey-brown or

reddish brown, shallowly dippled. Twigs 3-5 mm thick. Leaves in close clusters, thincoriaceous, glaucous below; elliptic or oblong, 6.5-12.5 x 2.5-4 cm; base attenuate, apex acute;
midrib raised above, sharply keeled below, lateral veins 12-16 pairs, visible below, faint above,
intercostal veins reticulate, very faint; petioles 1.5-3.5 cm long, swollen at the base.
Inflorescences panicles, to 22 cm long, hairy. Flowers white; calyx divided up to half of its
length; petals densely hairy on both surfaces; ovary subglobose. Fruits ovoid-oblong or
ellipsoid, 1.7-2 x 0.7-1 cm; enlarged petals linear-oblong, 5.5-7 x 0.7-1 cm.
Distribution. Myanmar, Thailand, Cambodia, Sumatra, Peninsular Malaysia, Borneo.
Uncommon in Sabah and Sarawak; also in Kalimantan.
Ecology. Lowland mixed dipterocarp forest, to 200 m.
92
BORAGINACEAE
A.L. Dayang Awa
Universiti Malaysia Sarawak,
Kota Samarahan, Sarawak, Malaysia
King, J. As. Soc. Beng. 74, Extra No. (1906) 487; Ridley, FMP 2 (1923) 438; Merill, EB (1921) 510;
Johnston, J. Arn. Arb. 16 (1935) 1, l.c. 32 (1951) 1; Masamune, EPB (1942) 637; Backer &
Bakhuizen f., FJ 2 (1965) 457; Burgess, TBS (1966) 58; Kazmi, J. Arn. Arb. 51 (1970) 133, 367, l.c.
53 (1971) 110, 334, 486, 666; Keng, OFMSP (1978) 270; Anderson, CLTS (1980) 154; Cockburn, TS
2 (1980) 20; Whitmore & Tantra, CLS (1986) 33; Corner, WSTM 3rd ed. 1 (1988) 197; Ng, TFM 4
(1989) 58; Whitmore, Tantra & Sutisna CLK 1 (1989) 33.
Herbs, shrubs, trees, or rarely lianas; parts often scabrid- or hispid-hairy. Leaves alternate or
spirally arranged, rarely opposite or in rossettes, simple, without stipules; blade mostly entire
or dentate, rarely deeply lobed. Inflorescences spikes, cymes or racemes, often
dichotomously branched and composed of 1 or more scorpioid or helicoid cymes, uncoiling
as the flowers open, very rarely a solitary flower. Flowers mostly bisexual, radially or
bilaterally symmetrical, mostly 5-merous; calyx tubular, cup- or bell-shaped with (4)5(8)
teeth or lobes, usually persistent in fruit; corolla gamopetalous, 47-lobed, funnel-shaped or
bell-shaped; stamens as many as corolla lobes, inserted on the corolla tube, alternating with
corolla lobes, filaments slender, often hairy at the base, anthers 2-locular, splitting
lengthwise, basifixed or subdorsifixed, introrse; disc if present annular; ovary superior,
entire, 4-lobed or rarely 2-lobed, carpels 2, style 1, terminal or arising from the middle part of
ovary lobes, generally simple, stigmas mostly capitate, bilobed or 4-lobed, ovules usually 2 in
each carpel, placentation axile. Fruit of 4 nutlets or a 14-seeded nut, a drupe, or a samara.
Seeds with or without endosperm; embryo straight or curved, radicle short.
Distribution. A large family of about 113 genera and 2400 species, distributed throughout
the world. In Sabah and Sarawak, represented by 7 genera with 9 species.
Ecology. Found in wide variety of habitats, from lowland to high altitude, in secondary and
primary forests.
Uses. In Sabah and Sarawak, currently of no economic importance, but elsewhere a number
of species are cultivated as ornamentals, notably those of Anthusa, Borago, Cerinthe,
Cynoglossum, Echium and Symphytum. The glutinous mesocarp of Cordia species has been
used for glue.
Key to genera
1. Trees or shrubs....................................................................................................................2
Herbs...................................................................................................................................5
93
TREE FLORA OF SABAH AND SARAWAK VOL. 2
2. Fruit a samara................................................................................................4. Pteleocarpa

Fruit a drupe........................................................................................................................3
3. Leaves covered with dense silvery hairs on both surfaces...................................1. Argusia
Leaves more or less glabrous...............................................................................................4
4. Fruits partly or completely enclosed by an enlarged and hardened calyx. Seed usually one
per fruit. Style twice forked, giving 4 ultimate stigmas.........................................2. Cordia
Fruits not as above. Seeds usually 24 per fruit. Style once forked, giving rise to 2
stigmas..................................................................................................................3. Ehretia
5. Diffused or tufted herbs. Fruits of 4 angular-rounded nutlets................................................
Trigonotis Steven
Bull. Soc. Nat. Mosc. 24 (1851) 603; Johnston, J. Arn. Arb. 18 (1937) 1, l.c. 21
(1940) 56, l.c. 33 (1952) 68; Kazmi, J. Arn. Arb. 51 (1970) 401. Synonym:
Havilandia Stapf, FMK (1894) 209.
About 35 species, distributed in Asia and Melanesia.
Perennial pubescent herbs. Flowers solitary in the axils of leaves; calyx deeply 5lobed or 5-partite, strigose, not or only slightly enlarged in fruit; corolla with 5
rounded lobes; stamens 5, included, inserted at the middle of corolla tube; style short,
stigmas capitate. Fruits subsessile or stalked.
In Borneo, 1 species (T. borneensis), so far known from Sabah (Mt. Kinabalu) only.
Erect herbs. Fruits not as above...........................................................................................6

6. Fruit a drupe, 2- or 4-lobed or furrowed, glabrous.................................................................
Heliotropium L.
Sp. Pl. (1753) 130; Merrill l.c. 511; Masamune l.c. 637; Ridley l.c. 441; Henderson,
Mal. Wild Fls. (1950) 310; Backer & Bakhuizen f. l.c. 461; Kazmi l.c. (1970) 149;
Cockburn l.c. 20.
About 300 species, in tropical and temperate zones of both hemispheres.
Erect scabrid herbs, rarely shrublets to 60 cm tall. Leaves spirally arranged or the
lower ones opposite; base of blade running down the petiole forming a narrow wing.
Flowers bisexual, in terminal scorpioid inflorescences; calyx 5-lobed or 5-partite;
corolla accrescent at anthesis, limb 5-angular or 5-lobed; stamens included, anthers
mucronate or obtuse; ovary 4-lobed, 4-locular, style short, stigmas annular.
In Sabah and Sarawak, only H. indicum is found, mainly in open waste ground in the
lowland and lower montane regions, to 1500 m.
Fruit a depressed pyramidal or ovoid-globose structure, covered with short-hooked

bristles...................................................................................................................................
Cynoglossum L.
Sp. Pl. (1753) 134; Backer & Bakhuizen f. l.c. 463; Kazmi l.c. (1971) 342; Cockburn
l.c. 20.
About 60 species, widely distributed throughout the world, except in the arctic
regions.
Perennial, biennial, rarely annual, erect soft-hairy herbs. Leaves entire, lower ones in
rossettes, with rather long petioles, upper ones spirally arranged, with short petioles or
subsessile. Flowers bisexual, 5-merous, mostly in abracteate often paniculate
scorpioid or helicoid inflorescences; calyx deeply divided, after anthesis enlarged and
spreading or curved; corolla tube short, throat scaly, lobes patent, rounded or obtuse;
94
BORAGINACEAE (DAYANG AWA)
stamens included; ovary deeply 4-partite, style short, stigmas broad.

In Sabah and Sarawak, 1 weedy species (C. lanceolatum), found mainly in shifting
cultivation areas, especially in the mountain region up to 3000 m.
1. ARGUSIA Boehm.
(name coined from an apparently deliberate misspelling of the river Argun in Manchuria)
In Ludwig, Defin. Gen. Pl. (1760) 507. Synonym: Messerschmidia L. ex Hebenstr., Nov. Comm.
Acad. Sci. Imp. Petrop. 8 (1763) 315, t. 11; Johnston, J. Arn. Arb. 16 (1935) 161, l.c. 32 (1951) 118;
Backer & Bakhuizen f., FJ 2 (1965) 461; H. Heine, F1. New Caledon. & Depend. 7 (1976) 108;
Cockburn, TS 2 (1980) 21; Ng, TFM 4 (1989) 64.
Herbs, shrubs or small trees. Leaves spirally arranged, often crowded at the ends of stout
branches, entire. Flowers bisexual, 5-merous, sessile or pedicellate, in terminal, ebracteate,
forked scorpioid cymes often combined into branched panicles or corymbs; calyx bell-shaped,
5-lobed, densely hairy; corolla tube cylindrical or bell-shaped, lobes spreading, conduplicate
in bud; anthers on very short filaments, mucronulate at apex; stigma a thick disc bearing 2
stout sterile lobes, with a receptive ring-like zone around the base. Fruits drupaceous, dry
when mature, breaking up into 2-seeded nutlets; mesocarp vesicular and corky.
Distribution. Three species, of which one occurs in Sabah. A. sibirica is distributed from
Japan through northern parts of China and across continential Asia (between 4055 N
latitude) to Rumania and Central Russia; A. gnaphalodes is restricted to the West Indies, and
A. argentea is a tropical strand species widely distributed in the Indian and Pacific Oceans.
Ecology. Confined to exposed and saline habitats, often constituting characteristic
components of strand vegetation on small offshore islands. In A. argentea and A.
gnaphalodes, the corky mesocarp facilitates fruit dispersal by oceanic currents.
Argusia argentea (L. f.) H. Heine
Fig. 1.
(Latin, argenteus = silvery; the indumentum)

l.c. 109. Basionym: Tournefortia argentea L. f., Sp. Pl. Suppl. (1781) 133; Ridley l.c. 440. Type:
Konig, s.n., Ceylon, 1777 (LINN). Synonyms: Messerschmidia argentea (L. f.) Johnston l.c. (1935)
164, Backer & Bakhuizen f. l.c. 461, Cockburn l.c. 21, Ng l.c. 64; Tournefortia arborea Blanco, Fl.
Filip. (1837) 129.
Shrub or much-branched small tree to 10 m tall. Young branches, leaves and inflorescences
covered with appressed silky white hairs. Leaves somewhat fleshy when fresh, clustered at
the ends of stout branches; broadly oblanceolate, 1020 x 48 cm; base attenuate, apex
obtuse to somewhat rounded; midrib sunken or grooved on the upper leaf surface, lateral
veins 34 pairs, intercostal veins obscurely reticulate; petioles 525 mm long. Inflorescence
a panicle, 918 cm across. Flowers numerous, initially crowded in compact groups but later
displayed in crowded, two-ranked, elongated pairs of scorpioid cymes 210 cm long; calyx
1.52 mm long, lobes fleshy, oblong to suborbicular, densely appressed hairy outside,
glabrous inside, not much enlarged in fruit; corolla white, lobes 47 mm broad, ovateorbicular, margin entire
95
Fig. 1. Argusia argentea. A, flowering leafy twig; B, open flower; C, flower in longitudinal section. (AC from
Mitchell M 26.)
96
or subdenticulate, base abruptly narrowed, outer surface strigose down the middle, tube
shorter than the lobes, bell-shaped, 1.52 mm long; filaments inconspicuous, inserted at 0.4
0.8 mm above the base of the corolla tube, anthers ovate-oblong, c. 1.5 mm long, half-exerted
from the open corolla throat, apex acute, thickened, mucronulate; ovary subglobose, 0.81.5
mm diameter, glabrous, stigmas as long as broad, sessile, composed of a thick stigmatic disc
bearing 2 stout oblong lobes. Fruits when mature dry, smooth, glabrous, depressed globose,
58 mm diameter, splitting into plano-convex halves, each half containing 2 fertile and a
small interposed sterile cavity.
Distribution. A common constituent of strand vegetation of small offshore islands, from the
Ryukyu southeastwards to the South Pacific and northern Australia, and southwestwards to
the Indian Ocean and Africa. In Sabah, it is found on Pulau Selingan and Pulau Langkayan
off Sandakan and Pulau Sipadan off Semporna; so far no record from Sarawak.
2. CORDIA L.
(Valerius Cordus, 15151544, German botanist)
Sp. Pl. (1753) 190; King, J. As. Soc. Beng. 74, Extra No. (1906) 488; Ridley, FMP 2 (1923) 438;
Johnston, J. Arn. Arb. 16 (1935) 3, l.c. 21 (1940) 336, l.c. 24 (1948) 227, l.c. 30 (1949) 85, 111, l.c.
32 (1951) 2; Burkill, EPMP 1 (1935) 660; Masamune, EPB (1942) 637; Backer & Bakhuizen f., FJ 2
(1965) 458; Burgess, TBS (1966) 58; Kazmi, J. Arn. Arb. 51 (1970) 139; Cockburn, TS 2 (1980) 21;
Whitmore & Tantra, CLS (1986) 33; Corner, WSTM 3rd ed. 1 (1988) 197; Ng, TFM 4 (1989) 60;
Trees, shrubs, or climbers (not in Borneo). Leaves entire, dentate, or crenate-lobed, spirally
arranged, alternate, or subopposite. Inflorescences laxly paniculate, glomerate, capitate, or
spicate cymes. Flowers usually 5-merous, bilaterally symmetrical, polygamous, usually male
and hermaphrodite; calyx obconical, bell-shaped or tubular, 45-toothed, usually persistent
in fruit; corolla white, yellow, orange or reddish, salverform, subrotate, funnelform or
subtubular, lobes 48; stamen as many as corolla lobes, inserted on the corolla tube,
alternating with the corolla lobes, filaments often hairy at base, anthers sagittate or hastate,
shortly exserted; ovary glabrous, 4-locular, with 1 ovule in each locule, style terminal,
dichotomous, of a simple column at base then dividing into 2 branches, which in turn fork to
produce the 4 ultimate branches each bearing a capitate or clavate stigma. Fruit a 14seeded drupe with bony endocarp and watery or glutinous mesocarp, or rarely without fleshy
mesocarp and hence a nut surrounded by or seated on the persistent calyx. Seeds without
endosperm.
Distribution. About 250 species, throughout the tropics in both hemispheres, centering in
America. Only 3 species are found in Borneo (Sabah, Sarawak, Brunei & Kalimantan), two
native (C. dichotoma and C. subcordata) and one (C. cylindristachya) introduced weed from
tropical America.
Ecology. In Sabah and Sarawak, found mainly in open forests or thickets at low to medium
altitude. Species found elsewhere in semi-evergreen forests are deciduous.
Uses. C. gerascanthus produces one of the best timbers in Jamaica. In Myanmar, wood of C.
97
fragrantissima is used for face powder due to its pleasant smell. In other species, though the
wood is easy to work with and seasons well, it is prone to insect attack. Cordia timber is not
commercially important because the trees have poor form and short bole, and large trees are
often hollow. The fruit exudate of C. dichotoma is very mucilaginous and has been used for
glue, though the adhesiveness is not permanent. The leaves of certain species are used to treat
wounds.
Key to Cordia species
1. Low,
bushy,
weedy
or
planted
shrubs.
Inflorescences
unbranched
spikes.............................
C. cylindristachya Roem. & Schult.
Syst. 4 (1819) 459; Henderson, Mal. Wild Fls. 1 (1959) 311; Corner l.c. 198; Ng l.c. 60.
Leaves scabrid, elliptic, 510 x 1.53 cm; base rounded-attenuate, margin dentate,
apex acute; lateral veins 58 pairs, intercostal veins coarsely scalariform, distinct on
both surfaces. Flowers c. 1 cm across, white, crowded in a terminal spicate cyme, the
distal flowers open first. Fruit a fleshy drupe, yellowish and turning red when ripe.
Native of tropical America. In Sabah and Sarawak, common weeds especially in
open, rather damp, low lying areas.
Trees. Inflorescences dichotomously branched panicles ....................................................2
2. Trees of inland areas, coastal hills, or inner fringes of mangroves. Flowers small; corolla
tube less than 1 cm long, white, green or cream, 46-lobed. Persistent calyx not enlarged
after anthesis, not enclosing the fruit, shallowly cup-shaped .....................1. C. dichotoma
Trees of sandy or rocky seashores. Flowers big; corolla tube 35 cm long, orange, 67lobed. Persistent calyx enlarged after anthesis, enclosing the fruit leaving only a small
opening at the apex....................................................................................2. C. subcordata
1. Cordia dichotoma Forster
Fig. 2.
(Latin, dichotomous = forked; the inflorescence)

Prod. 18 (1786) 110; Masamune l.c. 637; Johnston l.c. (1951) 8; Backer & Bakhuizen f. l.c. 459; Kazmi l.c.
(1970) 141; Cockburn l.c. 21; Corner l.c. 198; Ng l.c. 60. Type: Sine coll., s.n., New Caledonia (BM).
Synonyms: C. suaveolens Blume, Bijdr. Fl. Ned Ind. 14 (1826) 843; C. griffithii C. B. Clarke in Hooker f., F1.
Br. Ind. 4 (1885) 139; C. premnifolia Ridley, J. Roy. As. Soc. Str. Br. 68 (1915) 12.
Small tree to 13 m tall, 30 cm diameter. Bark smooth to vertically cracked, becoming thick
and fissured on old trees. Leaves membranous to coriaceous; ovate or orbicular, 416 x 13
10 cm; base attenuate, acute, rounded or cordate, margin entire, wavy or obscurely dentate,
apex acuminate; midrib slightly raised or grooved on the upper leaf surface, lateral veins 36
pairs, intercostal veins transverse-reticulate; petioles 1.54.5 cm long, slender or stout.
Inflorescences terminal, 411 cm long, forking repeatedly and bearing 10many flowers.
Flowers male or bisexual, on separate trees; pedicels articulate; calyx 48 mm long, opening
by bursting irregularly at apex, not enlarged after anthesis; corolla white, cream or green,
tube less than 1 cm long, with 46 spreading lobes, each up to 1 cm long and wide; stamens
46, filaments 45 mm long, anthers 23 mm; ovary 4-locular, style 56 mm long. Fruits
white,
98
Fig. 2. Cordia dichotoma. A, fruiting leafy twig; B, flower; C, flower in longitudinal section; D, ovary; E,
stamens. (A from SAN 92453, BE from fresh specimen.)
99
pale green, pink, orange or yellow, ovoid, 12.5 x 0.51.5 cm; persistent calyx shallowly cupshaped, 0.41 cm deep, with a wavy rim; mesocarp mucilagineous; endocarp stony. Seed 1
per fruit.
Vernacular names. Sabahpetudang (Pensiangan), potudung (Dusun Ranau), siopnama
(Ranau), tomudong (Dusun Bundu Tuhan).
Distribution. Wide-ranging from India to South China, across Malesia to NE Australia and
New Caledonia. In Sabah, it is found from Sandakan towards the western side of the state. In
Sarawak, it is known from a cultivated tree at Bukit Antu, Kuching.
Ecology. Sporadic but locally common at the back of mangroves, coastal hills below 100 m,
inland forest along rivers and on limestone.
Uses. The fruit rind contains a sticky gum which has been used in Peninsular Malaysia for
glue (Burkhill l.c. 660)
2. Cordia subcordata Lam.
(Latin, subcordatus = almost heart-shaped; the leaf base)
Tab. Encycl. 1 (1891) 421; King l.c. 488; Ridley l.c. 439; Masamune l.c. 637; Johnston l.c. (1951) 3;
Backer & Bakhuizen f. l.c. 458; Cockburn l.c. 21; Ng l.c. 62; Corner l.c. 199; Whitmore, Tantra &
Sutisna l.c. 33. Type: Commerson, s.n., ex insulis Praliniis (P). Synonym: C. rumphii Blume,
Bijdr. Fl. Ned. Ind. 14 (1826) 843.
Small tree to 15 m tall, 30 cm diameter; crown dense, bushy, round. Bark brown or grey,
shallowly fissured and flaky. Leaves membranous to thin-coriaceous; upper surface sparsely
tomentose, in older leaves frequently dotted with groups of mineralized epidermal cells,
lower surface more or less villulose or tomentulose along the midrib and lateral veins;
broadly ovate-elliptic, 620 x 515 cm; base acute, rounded or subcordate, occasionally
asymmetric, margin entire, apex acute to acuminate with a sharp point; midrib slightly raised
below, lateral veins 45 pairs, intercostal veins reticulate, fine and distinct on the lower leaf
surface; petioles 26 cm long. Inflorescences terminal, laxly cymose, 35 cm long, 14times forked, 520-flowered, often displaced to pseudoterminal position opposing the leaves.
Flowers: pedicels 210 mm long; calyx tubular, 1020 mm long, 48 mm diameter,
shallowly 3-lobed, glabrous or sparingly strigose, enlarged after anthesis; corolla trumpetshaped, orange, tube 35 cm long, with 57 spreading rounded lobes of 23.5 cm across;
filaments attached above the middle of the flaring corolla tube, anthers 23.5 mm long; ovary
4-locular, style 23 cm long. Fruit a nut, ovoid, obovoid or subglobose, 23 x 1.52.5 cm,
completely and tightly enclosed by the enlarged tin-walled persistent calyx, green ripening
yellow; mesocarp firm, corky, dry at maturity; endocarp bony, angular, much ridged and
roughened, containing 4 fertile cavities and a central sterile one, usually only two 1-seeded
locules developing.
Distribution. Hainan, Indo-China, India, Malesia, islands of the South Pacific and Indian
oceans and along the east coasts of Africa. In Sabah, known from Kuala Penyu and
Sandakan. There is no record from Sarawak.
Ecology. A plant of strand vegetation, the fruits are dispersed by ocean currents.
100
Uses. No recorded use in Sabah. However, the timber is used for making some musical
instruments in the Phillippines, and for house posts in Peninsular Malaysia (Burkill l.c. 670).
3. EHRETIA L.
(G. D. Ehret, 17081770, German botanical artist)
Syst. ed. 10 (1759) 936; King, J. As. Soc. Beng. 74, Extra No. (1906) 493; Ridley, FMP 2(1923) 441;
Johnston, J. Arn. Arb. 32 (1951) 19; Backer & Bakhuizen f., FJ 2 (1965) 459; Cockburn, TS 2 (1980)
21; Ng, TFM 4 (1989) 62.
Trees or erect or rarely scandent shrubs. Leaves spirally arranged, usually distinctly stalked.
Flowers mostly in axillary or terminal, many-flowered cymes, corymbs, or panicles, bisexual;
calyx small, deeply divided into 5 segments, not enclosing the corolla in bud, persistent;
corolla white or yellowish, tube bell-shaped or elongate, lobes obtuse, spreading or recurved;
stamens inserted on corolla tube, exserted, filaments filiform, anthers ovoid or ellipsoid;
ovary entire, 4-ovulate, style terminal, divided into 2 branches above the midpoint, stigmas
capitate or elongate. Fruit a drupe, subglobose, at maturity breaking apart into 2-seeded or 1seeded pyrenes.
Distribution. About 50 species, distributed mainly in tropical Asia, Central America, and
Africa. In Sabah, one species.
Ecology. Lowland forest, coastal areas of offshore islands, and on limestone hills. In Sabah,
has been reported from Ranau, Lahad Datu and Labuk Sugut districts. No record from
Sarawak.
Uses. A few species yield a good timber which is used in light-construction, and some
species produce edible fruits (Burkill l.c. 905). However, due to its small size, no uses have
been recorded in Sabah and Sarawak.
1. Ehretia timorensis Decne
Fig. 3.
(of Timor island)

Nouv. Ann. Mus. Hist. Nat. 2 (1834) 395; Ng l.c. 64. Synonym: E. laevis Roxb. var. timorensis
(Decne) C.B. Clarke in Hooker f., Fl. Br. Ind. 4 (1883) 142; King l.c. 493.
Small tree to 10 m tall. Bark smooth, whitish grey. Twigs pubescent or glabrous. Leaves
thin-coriaceous, glabrous, drying dark brown to black, lower surface paler; ovate to elliptic,
714 x 35 cm; base slightly cuneate, or sometime rounded or subcordate, margin entire,
apex shortly acuminate; midrib slightly grooved on the upper leaf surface, lateral veins 59
pairs, rather straight, ascending, intercoastal veins laxly reticulate; petioles 1017 mm long.
Inflorescences much-branched, bearing many flowers, axillary and terminal, 511 cm long.
Flowers small, numerous, sessile or subsessile; calyx 12 mm long, with 5 small triangular
lobes; corolla white, subrotate, tube 715 mm long, lobes 23 mm
101
Fig. 3. Ehretia timorensis. A, flowering leafy twig; B, flower; C, flower in longitudinal section. (AC
from Meh FMS 10175.)
102
long, recurved; filaments attached at 0.51 mm above the base of corolla tube, anthers 0.5
0.7 mm long; style 23 mm long. Fruits globose, smooth, 34 mm diameter, ripening yellow
or orange; pericarp thin; endocarp breaking up at maturity into 4 separate single-seeded parts;
persistent calyx much shorter than the fruit.
Distribution. Myanmar to Australia. In Sabah, known from Ranau district. No record from
Sarawak.
Ecology. Mixed dipterocarp forest.
4. PTELEOCARPA Oliv.
(Greek, Ptelea = a rutaceous genus, karpos = fruit; having fruits resembling that of Ptelea)
Trans. Linn. Soc. London 28 (1873) 518; Merrill, EB (1921) 510; Masamune, EPB (1942) 637;
Anderson, CLTS (1980) 154; Cockburn, TS 2 (1980) 22; Whitmore & Tantra, CLS (1986) 154; Ng,
TFM 4 (1989) 64; Whitmore, Tantra & Sutisna, CLK 1 (1989) 33.
Trees. Leaves spirally arranged, stalked. Inflorescences terminal, many-flowered panicles.

Flowers: calyx tubular, deeply 5-lobed; corolla tube short, 5-lobed; stamens 5, exserted from
the corolla throat; ovary thick-stalked, 2-locular, style divided into 2 almost from its base,
ovules 1 per locule. Fruit a samara.
Distribution. A monotypic genus, distributed in Thailand, Peninsular Malaysia, Sumatra and
Borneo (Sabah, Sarawak, Brunei and Kalimatan).
Uses. In Sabah and Sarawak, no recorded use, but in Sumatra, the timber which is fairly hard
and durable is used for construction works inside the house (Burkill l.c. 1855).
Pteleocarpa lamponga (Miq.) Bakhuizen ex Heyne
Fig. 4.
(of Lampong, Sumatra)

Nut. Pl. Ned. Ind., 2nd ed. (1927) 1309; Anderson l.c.154; Whitmore & Tantra l.c. 154; Ng l.c. 64;
Whitmore, Tantra & Sutisna l.c. 33. Basionym: Dodonaea ?lamponga Miq., Fl. Ind. Bat., Suppl.
(1855) 511. Type: Teijsmann, s.n., Sumatra, Lampong Prov. (holotype L; isotype BO). Synonyms: P.
malaccensis Oliv. l.c. 518, Cockburn l.c. 22; P. longistyla Becc., Malesia 1 (1877) 130, Merrill l.c.
510, Masamune l.c. 637.
Tree to 37 m tall, 50 cm diameter; crown bushy; bole straight. Bark smooth to shallowly
fissured; inner bark pale brown or pale orange, laminated. Sapwood pale yellow-brown.
Leaves membranous to thin-coriaceous, glabrous, withering yellow before shedding;
obovate-elliptic, 314 x 16 cm; base attenuate, margin entire, apex acuminate; midrib
prominent on both sides of the leaf, lateral veins 47 pairs, fine, fading toward the margin,
intercostal veins finely reticulate. Inflorescences terminal, many-branched, many-flowered,
to 7 cm long. Flowers bright yellow; pedicels c. 5 mm long; sepals 5, c. 2 mm long,
imbricate; corolla tube c. 2 mm long, corolla lobes 5, 36 mm long; stamens 5, exserted from
the mouth of corolla
103
Fig. 4. Pteleocarpa lamponga. A, flowering leafy twig; BC, flowers; D, fruiting leafy twig. (AC
from living trees cultivated at the FRIM arboretum; D from Enggoh FDNB 10558.)
104
tube; ovary c. 3 mm long, stipe c. 1 mm, styles 2, c. 0.51 mm long; ovules 1 per locule.
Fruits flat, winged, broadly elliptic to orbicular, c. 5.5 x 5 cm, pale green or yellowish,
ripening pink to light red. Seed 1 per fruit, elongate.
Distribution. As for the genus. In Sabah, known from Beaufort, Ranau and Kudat (Banggi
Island) districts. In Sarawak, recorded from Balai Ringin and Matang FR. Also in Brunei and
Kalimantan.
Ecology. In mixed dipterocarp and kerangas forests, to 600 m. The winged fruits are
dispersed by wind.
105
106
__________________________________________________________________________________________
__________________________________________________________________________________________
CAPRIFOLIACEAE
Noorma Wati Haron
University of Malaya,
Kuala Lumpur, Malaysia
King & Gamble, J. As. Soc. Beng. 72, 2 (1903) 112; Merrill, EB (1921) 582; Ridley, FMP 2 (1923) 1; Masamune, EPB
(1942) 719; Backer & Bakhuizen f., FJ 2 (1965) 357; Kern & van Steenis, FM 1, 4 (1951) 175, l.c. 1, 6 (1972) 928;
Anderson, CLTS (1980) 158; Corner, WSTM 3rd ed. 1 (1988) 204; Wong & Saw, TFM 4 (1989) 66; Whitmore, Tantra &
Shrubs, small trees, sometimes lianas or perennial herbs; twigs terete. Leaves simple or rarely
compound, opposite or alternate, sometimes with stipules. Inflorescences terminal, corymbose
cymes, panicles or umbellate corymbs. Flowers bisexual or unisexual, radially or bilaterally
symmetrical, 45-merous; calyx (4)5-lobed or toothed, the lobes usually small, calyx tube fused to
the ovary; corolla (3)5-lobed or sometimes 2-limbed, lobes imbricate in bud, corolla tube variable
from very short to long and narrow; stamens inserted on the corolla tube, usually as many as corolla
lobes, anthers dorsifixed or basifixed, usually introrse, rarely extrorse, 2-locular, the locules parallel,
opening by longitudinal slits; ovary inferior, 25(8)-locular, style simple with capitate or lobed
stigma; ovules 1 to numerous, pendulous, anatropous, placentation axile. Fruit usually a fleshy berry
or a drupe, sometimes an achene or a capsule. Seeds one to several per fruit; testa bony, rarely
membranous.
Distribution. Fifteen genera with about 400 species, cosmopolitan, mostly in the temperate regions of
N America and E Asia and montane tropical areas. In Sabah and Sarawak, represented by 2 genera
with 8 species.
Ecology. Growing along streams and on hill slopes in secondary forest, and primary montane forest.
Flowers of most species are pollinated by insects such as bees, flies and beetles, although selfpollination may also occur.
Uses. Many genera are of horticultural value as ornamental flowering shrubs and most species treated
here are used for such purposes. The family produces no important commercial timbers.
Taxonomy. Kern and van Steenis (1951) are of the opinion that Caprifoliaceae is closely related to
Valerianaceae, an opinion shared by Thorne (Nord. J. Bot. 3 (1983) 85117) and Cronquist (Integr.
Syst. Class. Fl. Pl. 1981) who placed the two families in the order Dipsacales. Dahlgren (Nord. J. Bot.
3 (1983) 119149), on the other hand, excluded Caprifoliaceae from the order Dipsacales and split the
family into three smaller families, the Caprifoliaceae s.s., Sambucaceae and Viburnaceae, and placed
these families in the order Cornales. Takhtajan (Syst. Magnol. 1987) agreed with Dahlgren in splitting
the Caprifoliaceae into three smaller families but included them in the order Dipsacales. In the present
107
account, a broader concept of the family as proposed by Kern & van Steenis (l.c.), Cronquist (l.c.) and
Thorne (l.c.) is followed.
Key to genera
Leaves simple. Flowers in umbellate corymbs or few-flowered paniculate inflorescences. Fruit a
drupe.........................................................................................................................................Viburnum
Leaves pinnately or bipinnately compound. Flowers in compound, corymbose cymes or manyflowered paniculate inflorescences. Fruit a 35-seeded berry......................................................
Sambucus L.
Sp. Pl. (1753) 269; Kern & van Steenis l.c. (1951) 190; Backer & Bakhuizen f. l.c. 357; Corner l.c. 205; Wong &
Saw l.c. 66.
About 25 species, distributed in subtropical and temperate regions worldwide.

Small trees or shrubs, rarely perennial herbs. Leaves opposite or alternate; leaflet margins serrate or
narrowly lobed; stipules present or absent. Flowers radially symmetrical, bisexual or unisexual, with
flowers of both sexes usually in the same inflorescence; calyx tube ovoid or turbinate, limb 35toothed; corolla rotate, 35-lobed, lobes imbricate or rarely valvate in bud; stamens 5, inserted at the
base of corolla tube, filaments filiform, anthers oblong; ovary 35-locular, ovule 1 in each locule, style
short, stigmas 3 or 5.
In Sabah, represented by one introduced and naturalised species, S. javanicus, commonly found in
secondary hill and lower montane forest, at 6001500m.
VIBURNUM L.
(an old Latin plant-name)
Sp. Pl. (1753) 267; King & Gamble, J. As. Soc. Beng. 72, 2 (1903) 112; Merrill, EB (1921) 582; Ridley, FMP 2 (1923) 1;
Masamune, EPB (1942) 719; Kern, Reinwardtia 1 (1951) 111; Kern & van Steenis, FM 1, 4 (1951) 180; Backer &
Bakhuizen f., FJ 2 (1965) 358; Anderson, CLTS (1980) 158; Corner, WSTM 3rd ed. 1 (1988) 183; Wong & Saw, TFM 4
Trees, shrubs or treelets. Leaves simple, opposite, pinnately or palmately (non-Bornean taxa) veined,
without stipules. Inflorescence a terminal, umbellate corymb or a few-flowered panicle; bracts and
bracteoles usually small, caducous. Flowers bisexual, radially symmetrical; calyx tube turbinate or
cylindrical, limb 5-toothed; corolla white, pink or yellowish, rotate, bell-shaped or tubular, 5-lobed;
stamens 5, exserted, anthers basifixed or dorsifixed; ovary 1-locular, style short, conical, stigmas 3,
ovules anatropous, pendulous, solitary. Fruit a drupe, crowned by persistent calyx and style. Seeds
oblong; testa horny or stony; endosperm often hard and copiously folded.
Distribution. About 225 species; north temperate regions, South America, Madagascar and tropical
Asia. In Sabah and Sarawak, 7 species are known.
Ecology. In primary and secondary forests to 1800 m.
108
CAPRIFOLIACEAE (NOORMA WATI HARON)
Uses. None of the species found in Sabah and Sarawak is known to be of economic importance.
Key to Viburnum species
1. Leaf margin coarsely crenate-serrate to serrulate along the upper two-thirds, and entire or
faintly denticulate at the lower third..............................................................5. V. lutescens
Leaf margin entire throughout.............................................................................................2
2. Leaves narrowly elliptic-oblanceolate, apex acute to acuminate. Fruits not compressed
and more or less rounded in cross-section....................................................3. V. clemensae
Leaves obovate, elliptic-orbicular, elliptic-oblong, or broadly elliptic, apex cuspidate,
rounded, truncate or emarginate, rarely acute-acuminate. Fruit compressed, bilobate,
biconvex or somewhat unequally 4-angular in cross-section..............................................3
3. Leaves obovate or elliptic-orbicular, thick-coriaceous........................................................4
Leaves elliptic-oblong to broadly elliptic, thin-coriaceous.................................................5
4. Leaves obovate, upper surface shiny; apex rounded, truncate, emarginate, or very rarely
abruptly cuspidate; midrib angular on lower surface, distinct glands present at the base on
both sides. Fruits c. 5 4 mm....................................................................7. V. vernicosum
Leaves elliptic-orbicular, upper surface not shiny; apex cuspidate; midrib flat on lower
surface, both sides without distinct glands. Fruits 1011 810 mm..............2. V. beccarii
5. Leaves elliptic-oblong, both surface densely punctate, lower surface hispidulous; midrib
raised and angular on the lower surface. Fruits biconvex in cross-section............................
...................................................................................................................4. V. hispidulum
Leaves broadly elliptic, glabrous to minutely papillose; midrib raised and rounded on the
lower surface. Fruits bilobed or somewhat unequally 4-angular in cross-section...............6
6. Peduncles more than 12 cm long. Fruits c. 16 3 mm, bilobed in cross section ...................
................................................................................................................1. V. amplificatum
Peduncles to 3 cm long. Fruits c. 7 6 mm, somewhat unequally 4-angular in
cross-section..............................................................................................6. V. sambucinum
1. Viburnum amplificatum Kern
(Latin, amplificatio = enlargement; the large leaves and fruits)
l.c. 150; Kern & van Steenis l.c. 188, Sar. Mus. J. 9 (1960) 679, l.c. (1972) 929; Whitmore, Tantra & Sutisna l.c. 40. Type:
Elmer 21741, North Borneo, Tawao, Elphinstone Prov. (holotype G; isotypes L, S).
Small treelet to 6 m tall. Sapwood white. Bark smooth, greyish brown; inner bark pale cream. Young
parts sparsely hairy to almost glabrous. Leaves thin-coriaceous, glabrous; broadly elliptic to rarely
elliptic-obovate, 7.719.8 4.68.1 cm; base cuneate to acute, margins entire, apex cuspidate; midrib
flat and slightly sunken above, raised and rounded beneath; lateral veins 58 pairs, visible but not
109
prominent on both surfaces; intercostal veins visible but not prominent on both surfaces; petioles 1.3
2.9 cm long. Inflorescence a terminal corymb; peduncles to 12 cm; primary rays to 7. Flower buds
ovoid to subglobose; calyx tube c. 1.5 mm long, lobes 5, narrowly ellipsoid; corolla bell-shaped;
stamens c. 1 mm long, anthers basifixed; ovary slightly oblong, style less than 1 mm long, tubular,
stigmas fused. Fruits greenish, narrowly oblong to ellipsoid, grooved, c. 16 3 mm, bilobed in crosssection. Seeds oblong, almost black, with rough surface.
Distribution. Endemic to Borneo. Known from a few collections from Lahad Datu, Tawau and Ranau
districts in Sabah. Not yet reported from Sarawak; also known from Kalimantan.
Ecology. Along streams or on hillsides in primary forest to 360 m.
2. Viburnum beccarii Gamble
Fig. 1.
(Odoardo Beccari, 18431920, Italian botanist and explorer)

In King & Gamble l.c. 114; Ridley l.c. 2; Kern l.c. 120; Kern & van Steenis l.c. (1951) 183; Wong & Saw l.c. 67. Type:
Scortechini 375b, Peninsular Malaysia (n.v.).
Shrub or treelet to 5 m tall. Bark smooth; inner bark light brown. Twigs terete, glabrous. Young parts
densely covered with minute, short, stiff hairs. Leaves thick-coriaceous, densely punctate on both
surfaces, not shiny above; elliptic-orbicular, 7.39.8 5.67.5 cm; base rounded, margins entire,
apex cuspidate; midrib flat beneath, at base on both sides without distinct glands; lateral veins 56
pairs, obscure above, prominent beneath; intercostal veins visible beneath, obscure above; petioles,
1.72.3 cm long. Inflorescence a terminal corymb; peduncles 48 cm long. Flowers with clavate
calyx tube, to 0.3 cm long; petals white, tubular, lobes 5, acute; stamens about 4 mm long, attached to
the base of corolla tube; ovary glabrous, cylindrical, style less than 1 mm long, stigmas capitate.
Fruits ovoid to globose, grooved, green with reddish base when young, ripening blackish red, 1011
810 mm, somewhat unequally 4-angular in cross-section. Seeds ovoid, dark brown to almost
black, smooth.
Distribution. Sumatra, Peninsular Malaysia, Borneo. In Sarawak, uncommon, known only from Mt.
Murud, Lawas; not yet recorded in Sabah.
Ecology. Along river banks, on rocky areas or sandstone cliffs in lower montane forest at 14001800
m.
3. Viburnum clemensae Kern
Fig. 2.
(Mary Strong Clemens, who collected plants on Mt. Kinabalu in 1915, 1916, 1917 and 193133)
l.c. (1951) 157; Kern & van Steenis l.c. (1951) 189; Whitmore, Tantra & Sutisna l.c. 40. Type: Clemens 29978, North
Borneo, Mt. Kinabalu, Tenompok (A).
Small tree to 13 m tall. Bark brown; inner bark yellow to brown. Young parts sparsely hairy. Leaves
thin-coriaceous, densely papillose on both surfaces; elliptic-oblanceolate, 4.912.5 25.5 cm; base
cuneate, margins entire, apex acute to acuminate; midrib slightly sunken above, raised and rounded
110
Fig. 1. Viburnum beccarii. A, fruiting leafy twig; B, flower bud; C, flower in longitudinal section; D, stamen; E, fruits; F,
fruit in longitudinal section; G, seed; H, side view of seed. (AH from S. 32934.)
111
Fig. 2. Viburnum clemensae. A, flowering leafy twig; B, part of inflorescence; C, flower bud; D, flower bud in longitudinal
section; E, stamen; F, fruits; G, fruit in longitudinal section; H, seed. (AH from SAN 48000.)
112
beneath, glands distinct at base on both sides; lateral veins 56 pairs, visible but not prominent
beneath, obscure above; intercostal veins visible but not prominent on both surfaces; petioles 13 cm.
Inflorescences terminal panicles; peduncles to 8.5 cm long; bracts and bracteoles minute. Flower
buds subglobose; calyx tube 1.52 mm long, lobes 5, triangular; petals white; anthers basifixed; ovary
ovoid-oblong, style less than 1 mm long, stigma surface rounded. Fruits ovoid-ellipsoid to obovoid,
13 67 mm, subtended by remains of calyx lobes, not compressed and more-or-less rounded in
cross-section. Seeds black.
Distribution. Endemic to Borneo. In Sabah, known from Kota Belud, Tambunan, Tawau, Kundasang
and Ranau areas; in Sarawak, not yet recorded; also occurs in Kalimantan.
Ecology. In primary and secondary lower montane forests on ridge tops or hill slopes at 13001800
m.
4. Viburnum hispidulum Kern
Fig. 3.
(Latin, hispidulus = with minute, coarse, erect hairs; the young parts and lower leaf surface)
l.c. 1 (1951) 136; Kern & van Steenis l.c. (1951) 187; Anderson l.c. 158; Whitmore, Tantra & Sutisna l.c. 41 (cited as V.
hispidum Kern). Type: Clemens 31902, North Borneo, Mt. Kinabalu, Upper Kinataki River (holotype B; isotype L).
Tree to 30 m tall. Bark fissured; inner bark orange. Sapwood cream-yellow. Young parts sparsely
hispidulous. Leaves thin-coriaceous, densely punctate on both surfaces, lower surface hispidulous,
especially on midrib and lateral veins; elliptic-oblong, 6.617 2.87.8 cm; base cuneate to slightly
rounded, margins entire, apex cuspidate; midrib raised and angular beneath, lateral veins 69 pairs,
visible but not prominent beneath, obscure above, intercostal veins visible but not prominent beneath,
obscure above; petioles 1.36 cm long. Inflorescence a terminal, umbellate corymb; penduncles 6.7
9 cm long; primary rays to 6; bracteoles present. Flower buds globose, rotate-cupular when open;
calyx tube c. 2 mm long, lobes 5, triangular; petals white, with a strong sweet scent; stamens exserted,
anthers dorsifixed, yellowish, thick; ovary cylindrical, style c. 1 mm long, stigmas capitate. Fruits
ellipsoid to obovoid, c.10 56 mm, green, ripening black, biconvex in cross-section. Seeds oblongellipsoid, dark brown to black with rough surface.
Distribution. Endemic to Borneo. In Sabah, recorded from Ranau, Sg. Inarat, Mt. Kinabalu, Mt. Trus
Madi areas; and in Sarawak, from Bario, Limbang, Kapit, Kelabit Highlands, Lawas, and Mt. Serapi;
also occurs in Kalimantan.
Ecology. In primary lower montane forest at 16001800 m.
5. Viburnum lutescens Blume

(Latin, lutescens = becoming pale yellow; the lower surface of dried leaves)
Bijdr. Fl. Ned. Ind. 13 (1826) 655; King & Gamble l.c. 114; Merrill l.c. 582; Ridley l.c. 2; Masamune l.c. 719; Kern l.c. 142;
Kern & van Steenis l.c. (1951) 188; Backer & Bakhuizen f. l.c. 360; Corner l.c. 206; Whitmore, Tantra & Sutisna l.c. 41;
Wong & Saw l.c. 67. Type: Blume, s.n., Java (holotype L).
113
Fig. 3. Viburnum hispidulum. A, fruiting leafy twig; B, flowering leafy twig; C, flower buds; D, flower bud in longitudinal
section; E, young fruits; F, fruits in longitudinal sections; G, seed. (AG from S. 50970.)
114
Shrub or treelet to 9 m tall. Twigs terete. Young parts stellate hairy. Leaves thin-coriaceous, usually
glabrous; ovate-elliptic to oblong-elliptic, 9.511.6 4.25.5 cm; base cuneate to slightly rounded,
margin entire or faintly denticulate to about one-third from the base, upper two-thirds coarsely
crenate-serrate to serrulate, apex acuminate; midrib flattened to slightly rounded beneath, lateral
veins 68 pairs, prominent beneath, obscure above, intercostal veins prominent beneath; petioles 1.2
2 cm long. Inflorescence a terminal, sometimes axillary, panicle; peduncles 2.53.4 cm long; bracts
and bracteoles present, hairy. Flowers fragrant, cream-white; buds ovoid to subglobose; calyx tube c.
1 mm long, lobes 5, ovate-triangular; corolla rotate-campanulate, lobes 5, stamens shortly exserted, c.
1 mm long, anthers basifixed; ovary cylindrical, style less than 1 mm long, stigmas united. Fruits
oblong-ellipsoid, slightly flattened, grooved, 812 6 mm, ripening black. Seed dark brown.
Distribution. Peninsular Malaysia, Java, Borneo, and Lesser Sunda Islands. In Sabah, uncommon,
known only from a few collections from Tambunan district; not yet recorded from Sarawak; also
occurs in Kalimantan.
Ecology. Lowland to lower montane forest at 7501200 m.
6. Viburnum sambucinum Blume

(resembling Sambucus)
l.c. (1826) 656; Miquel, Fl. Ind. Bat. 2 (1856) 120; King & Gamble l.c. 113; Merrill l.c. 582; Ridley l.c. 1; Masamune l.c.
719; Kern l.c. 129; Kern & van Steenis l.c. (1951) 186; Backer & Bakhuizen f. l.c. 359; Corner l.c. 205; Wong & Saw l.c.
67; Whitmore, Tantra & Sutisna l.c. 41. Type: Blume, s.n., Java (L).
Shrub or treelet to 10 m tall. Young parts hairy. Bark greyish brown, smooth. Leaves thincoriaceous, glabrous except for the sparsely hairy veins beneath; broadly elliptic to oblonglanceolate, 10.714 57 cm; base cuneate, margins entire, apex cuspidate; midrib slightly rounded
beneath, glands indistinct, lateral veins visible but not prominent on both surfaces, 46 pairs,
intercostal veins visible but not prominent beneath; petioles 1.43 cm long. Inflorescence a terminal
umbellate corymb; peduncles to 3 cm long. Flowers cream-white, fragrant; bud globose; calyx tube
cylindrical, lobes 5, ovate-triangular; corolla rotate, lobes 5; stamens 5, 23 mm long, exserted; ovary
cylindrical, more or less hairy, style short, stigmas united. Fruits dark purple, ovoid to globose, c. 7
6 mm, somewhat unequally 4-angular in cross-section. Seeds dark brown to black.
Distribution. Sumatra, Peninsular Malaysia, Java, Borneo, Sulawesi, and Lesser Sunda Islands. In
Sabah and Sarawak, uncommon.
Ecology. Lowland to lower montane forest to 1200 m.
7. Viburnum vernicosum Gibbs
Fig. 4.
(Latin, vernicosum = varnished, shiny; the upper leaf surface)

J. Linn. Soc. Bot. 42 (1914) 86; Merrill l.c. 582; Masamune l.c. 720; Kern l.c. 139; Kern & van Steenis l.c. 187; Whitmore,
Tantra & Sutisna l.c. 41. Type: Gibbs 3990, North Borneo, Mt. Kinabalu, Kiau (BM).
115
Fig. 4. Viburnum vernicosum. A, flowering leafy twig; B, part of an inflorescence; C, flower in longitudinal section; D, fruit;
E, fruit in side view; F, fruit in longitudinal section; G, seed. (AG from RSNB 4410.)
116
Shrub or treelet to 5 m tall. Bark dark brown. Twigs glabrous, terete. Young parts glabrous. Leaves
thick-coriaceous, densely punctate on both surfaces, shiny above, glands distinct at base on both sides
of midrib; obovate to ovate-elliptic, 6.57.5 45.2 cm; base rounded to cuneate, margins entire, apex
rounded, truncate, emarginate, or rarely abruptly cuspidate; midrib angular beneath, lateral veins 45
pairs, visible on both surfaces but more prominent beneath, intercostal veins visible beneath, obscure
above; petioles 811 mm long. Inflorescence a terminal, umbellate corymb; peduncles to 5.4 cm
long; bracteoles present. Flowers ovoid to obovoid in bud; calyx green, narrowly triangular; petals
cream-white, ovate triangular; stamens exserted, filaments thick, attached to the base of corolla; ovary
cylindrical to narrowly ellipsoid, style less than 1 mm long, stigmatic surface flat. Fruits ovoid,
grooved, c. 5 4 mm, somewhat unequally 4-angular in cross-section. Seeds ovoid-globose, black,
surface rough.
Distribution. Endemic to Borneo. In Sabah, known from Mt. Kinabalu; in Sarawak, from Lawas.
Also occurs in Kalimantan.
Ecology. Lowland forest on ridges and lower montane forest, at 9601800 m.
117
118
__________________________________________________________________________________________
___________________________________________________________________________
CASUARINACEAE
Runi S. Pungga
Kuching, Malaysia
Ridley, FMP 3 (1924) 371; MitcheIl, MF 27 (1964) 118; Backer & Bakhuizen. f., FJ 2 (1965) 8; Smythies, CST (1965) 33;
Burgess, TBS (1966) 70; Johnson, Nuytsia 1 (1972) 261, Telopea 2 (1980) 83, l.c. 3, 2 (1988) 113, J. Adelaide Bot. Gard. 6,
1 (1982) 73; Keng, OFMSP (1978) 128; Ng, TFM 3 (1978) 45; Anderson, CLTS (1980) 159; Cockburn, TS 2 (1980) 23;
Corner, WSTM 3rd ed. 1 (1988) 207; Whitmore, Tantra & Sutisna, CLK 1 (1989) 41; Wilson & Johnson, Fl. Austral. 3
(1989) 100.
Dioecious or less commonly monoecious trees or shrubs; bole straight, cylindrical, sometimes fluted;
short buttresses sometimes present. Bark greyish to reddish brown, rough, fissured and shedding off
into narrow, elongate flakes. Twigs of two kinds, woody and persistent, and non-woody and
caducous; the latter modified into green needle-shaped structures, branched or unbranched,
superficially resembling those of pine trees, internodes rounded or quadrangular in cross-section,
usually with longitudinal grooves or furrows, nodes swollen. Leaves in whorls of 418, tiny, scalelike, apex rounded-acute to narrowly acute. Male inflorescences usually terminal, simple or
compound, erect, spike-like structures; each male flower of one or two scale-like perianth lobes
(falling at anthesis) and a single stamen with basifixed anther. Female inflorescence a globose cone,
terminal or subterminal on woody persistent twigs or on green needle-shaped twigs; each female
flower of one-carpellate ovary with 2-branched, divergent stigmas, subtended by a bract and two
bracteoles. Fruit cone (infructescence) a woody structure developed from a whole female
inflorescence, with several alternating woody bracts, each subtending a pair of woody bracteoles,
bracteole-pairs gaping open at maturity. Fruitlet a samara with membranous wing. Seed solitary in
each samara, often with more than one embryo, each with two large cotyledons.
Distribution. Four genera with about 100 species; from SE Asia to Australia and Polynesia. Three
genera with 4 species in Sabah and Sarawak.
Ecology. Often growing gregariously in sandy beach vegetation (Casuarina equisetifolia) and in
kerangas, hill, and montane forest on ultramafic soils and limestone outcrops (Ceuthostoma and
Gymnostoma spp.), from sea level to 2000 m.
Uses. In the past, timbers of Casuarina equisetifolia were used for shingles and other construction
work, but they are now of little commercial importance. The timber is hard and heavy, excellent for
firewood and for making charcoal and matches.
Taxonomy. Prior to Johnsons taxonomic work (l.c. 1980, 1982, 1988), the family was considered
monogeneric having Casuarina as the only genus. Johnson, however, reclassified Casuarina s.l. into
4 smaller genera, namely Allocasuarina, Casuarina s. str., Ceuthostoma, and Gymnostoma, based on
the number of scale-leaves per node, branching pattern, the cross-sectional outline and position and
119
arrangement of stomata in the grooves of the needle-shaped twigs, the structure of the infructescence,
the form of the bracts and bracteoles, and chromosome numbers. In the present account, we follow
Johnsons scheme but note that it is difficult to distinguish Ceuthostoma from Gymnostoma using
only sterile specimens and without the help of a scanning electron microscope.
Key to genera
1.
Needle-shaped twigs rounded in cross-section; scale-leaves 518 per node................................

.................................................................................................................................1. Casuarina
Needle-shaped twigs quadrangular in cross-section; scale-leaves 4 per node............................2
2. Furrows of internodes deep and narrow, concealing the stomata. Fruit cones terminal, c.
1 cm diameter, with 16 fertile fruitlets; bracts thin and inconspicuous; bracteoles inserted
at an oblique angle with the fruit cone axis, apex acute....................................2. Ceuthostoma
Furrows of internodes shallow and open, exposing the stomata. Fruit cones terminal
or subterminal, 2.25 cm diameter, with 2035 fertile fruitlets; bracts broad, welldeveloped; bracteoles inserted perpendicularly to the fruit cone axis, apex acute
or rounded......................................................................................................3. Gymnostoma
1. CASUARINA L.
(resembling the plumage of the Cassowary; the needle-twigs)

rhu laut (Malay)
Amoen. Acad. 4 (1759) 143; Ridley, FMP 3 (1924) 371; Mitchell, MF 27 (1964) 118; Backer & Bakhuizen f., FJ 2 (1965) 9;
Smythies, CTS (1965) 33; Hutchinson, Gen. Fl. Pl. 2 (1967) 143; Johnson, Nuytsia 1 (1972) 261, J. Adelaide Bot. Gard. 6, 1
(1982) 79; Keng, OFMSP (1978) 128; Ng, TFM 3 (1978) 45; Anderson, CLTS (1980) 159; Cockburn, TS 2 (1980) 23;
Corner, WSTM 3rd ed. 1 (1988) 209; Whitmore, Tantra & Sutisna, CLK 1 (1989) 41; Wilson & Johnson, Fl. Austral. 3
(1989) 100.
Dioecious or monoecious trees; bole cylindrical, straight; crown narrow, branches ascending, with a
rather untidy appearance. Bark greyish to reddish brown, ridged, fissured and peeling off into
oblong flakes. Needle-shaped twigs mostly unbranched; internodes 0.51.5 cm long, rounded in
cross-section, furrows of internodes deep and closed. Scale-leaves 518 per node, narrowly acute.
Male inflorescences simple, elongated spikes, at the end of long needle-shaped twigs. Female
inflorescences lateral on woody persistent twigs. Fruit cones globose or ovoid-cylindrical; bracts
thin at the exposed portions, distally not expanded; bracteoles protruding slightly from the cone
surface, not greatly thickened. Samaras glabrous, dull grey or pale yellow-brown.
Distribution. About 17 species; from the Bay of Bengal to Malesia, Melanesia, Polynesia, New
Caledonia and Australia. Only C. equisetifolia is found in Borneo (Sabah, Sarawak, Brunei and
Kalimantan).
120
CASUARINACEAE (RUNI PUNGGA)
Ecology. Found on infertile, sandy or acidic soils along the coasts (C. equisetifolia) and elsewhere
(other species) on mountains.
Casuarina equisetifolia L.
(Latin, folius = leaf; with foliage resembling that of horsetails, Equisetum)
Fig. 1.
Amoen. Acad. 4 (1759) 143; Ridley l.c. 372; Mitchell l.c. 119; Backer & Bakhuizen f. l.c. 10; Smythies l.c. 119; Ng l.c. 45;
Anderson l.c. 159; Cockburn l.c. 23; Johnson l.c. (1980) 84, l.c. (1982) 79; Corner l.c. 210; Whitmore, Tantra & Sutisna l.c.
41; Wilson & Johnson l.c. 100. Type: Rumphius, Herbarium Amboniense 3 (1743) t. 57.
Monoecious trees to 40 m tall, 100 cm diameter; bole straight, cylindrical, usually fluted towards the
base; crown conical when young, becoming untidy at maturity. Bark greyish brown, ridged, fissured,
and flaking into oblong pieces. Needle-shaped twigs to 30 cm long, mostly unbranched, greenish
yellow, green stripes distinct; internodes 0.51.2 cm long, 0.51 mm thick, furrows usually densely
hairy. Scale-leaves (7)8(9) per node, apex narrowly acute. Male inflorescences simple elongated
spikes, terminal on needle-shaped twigs; stamens with basifixed anthers. Female inflorescences on
lateral woody persistent twigs, c. 0.5 cm long; style red, 0.30.5 cm long. Fruit cones ovoidcylindrical, c. 2.5 x 2.2 cm, sparsely hairy; peduncles 0.31.3 cm long; bracts woody but rather thin
in the exposed portion; bracteoles acute, slightly protuding, more or less of equal size. Samara 0.6
0.8 cm long.
Distribution. As for the genus; in Sabah and Sarawak, the species is found only along sandy beaches.
Ecology. Naturally common in sandy soils near the sea but also grows rather well when planted far
inland.
Uses. The wood is suitable for light construction, piling and fuel. In Peninsular Malaysia, the Malays
use a decoction of the twigs as a lotion to soothe swelling. The powdered bark is used to treat
pimples. The species is protected in Sarawak due to its importance in controlling coastal erosion. The
tree is frequently planted as a wind-break along major roads.
2. CEUTHOSTOMA L.A.S. Johnson

(Greek, keuthos = hidden, stoma = minute opening;
the concealed stomata in the furrows of needle-twigs)
Telopea 3, 2 (1988) 133.
Dioecious trees. Needle-shaped twigs 4-ribbed, furrows deep and almost closed, concealing the rows
of stomata, mostly branched. Leaf-scales 4 per node. Female flowers in globose, terminal cone-like
structures. Fruit cones longer than broad; bracts somewhat expanded distally; bracteoles strongly
protruding and inserted at an oblique angle with the cone axis, not strongly woody, convex dorsally,
with prominent dorsal eccentric rib, neither ridged nor with protuberences. Samara dull pale brown.
121
Fig. 1. Casuarina equisetifolia. A, fruiting twig; B, detailed arrangement of bracts and bracteoles in fruit cone; C, samara or
fruitlet; D, female inflorescence; E, nodal portion of needle-shaped twig; FG, male inflorescences. (AB and DE from S.
13733, C from S. 42032, FG from SAN 87058.)
122
Distribution. Two species; Palawan Is., Borneo, Maluku, and New Guinea. Only C. terminale is
found in Borneo (Sabah).
Ecology. In hill and lower montane forests on ultramafic soils, to c. 1500 m.
Ceuthostoma terminale L.A.S. Johnson
Fig. 2.
(Latin, terminalis = apical; the position of fruit cones)
l.c. (1988) 135; Wilson & Johnson in Crane & Blackmore, Evol. Syst. & Foss. Hist. Ham. 2 (1989) 182. Type: Clemens
30757, North Borneo, Mt. Kinabalu, 12001500 m (holotype K; isotypes A, BO, L, NSW).
Tree to 30 m tall, 30 cm diameter. Bark rough, scaly or flaky, peeling off into long broad strips.
Needle-shaped twigs rounded-quadrangular in cross-section, little-branched, longest unbranched
ones exceeding 10 cm long; internodes 0.51 cm long, occasionally set with white hairs on the
furrows. Scale-leaves narrowly deltoid, 0.71 mm long. Male flowers in simple terminal spikes.
Female inflorescences cone-like, terminal on moderately elongated needle-shaped twigs. Fruit
cones c. 1 cm across, with 14 whorls of fertile fruitlets; bracts woody, 22.5 mm high and broad, not
markedly ridged; bracteoles strongly protruding, to 57 mm long, with a prominent dorsal eccentric
rib, tips sharply acute. Samaras c. 6 mm long, with transparent wing.
Distribution. Borneo, Philippines (Palawan Is.), Maluku (Halmahera), and Irian Jaya. In Borneo, so
far known from Mt. Kinabalu and Mt. Tawai in Sabah.
Ecology. In primary and secondary forests on ultramafic soils, at 1001500 m.
3. GYMNOSTOMA L.A.S. Johnson

(Greek, gymnos = exposed, stoma = minute opening;
the position of stomata in the grooves of needle-twigs)
Telopea 2, l (1980) 83, J. Adelaide Bot. Gard. 6, 1 (1982) 83; Wilson & Johnson, Fl. Austral. 3 (1989) 103.
Trees or tall shrubs, dioecious or monoecious. Needle-shaped twigs mostly branched, internodes 4ribbed, furrows shallow and open, exposing the stomata, quadrangular in cross-section, ridges
rounded to somewhat sharp-edged. Scale-leaves in whorls of 4, apex rounded-acute to narrowly
acute. Male flowers in simple or compound spikes at the ends of needle-shaped twigs. Female
flowers in terminal or subterminal globose cones-like structures on woody persistent twigs or on
elongated branches of needle-shaped twigs. Fruit cones globose or cylindrical; bracts woody, much
expanded distally, with the abaxial surface below the apex broader than high; bracteoles prominently
protruding and inserted perpendicularly to the fruit cone axis, convex, dorsally neither split nor
furnished with protuberances. Samaras grey to pale brown.
123
Fig. 2. Ceuthostoma terminale. A, fruiting twig; B, part of needle-shaped twig showing the arrangement of scale-leaves; C,
a portion of woody twig; DE, fruit cones; F, male inflorescence. (A and D from RSNB 4490, BC from RSNB 4963, E from
SFN 26398, F from SMHI 1573.)
124
Distribution. About 17 species; Malesia, Solomons Is., tropical NE Australia, Fiji, and New
Caledonia; 2 species in Sabah and Sarawak.
Ecology. In open and closed lowland to montane habitats on shallow soils, stream banks, and peaty or
sandy substrates. Often found in shrubby plant communities on ultramafic soils.
Uses. The timber is used mainly as firewood and for piling work.
Key to Gymnostoma species

Needle-shaped twigs 11.5 mm thick, internodes 0.51 cm long; lateral branches long, with less or no
secondary branches, mostly exceeding 10 cm. Fruit cones globose, to 2.2 cm
across......................................................................................................................................1. G. nobile
Needle-shaped twigs c. 0.5 mm thick, internodes 0.20.3 cm long; lateral branches short, with many
secondary branches, not exceeding 10 cm long. Fruit cones ellipsoid, c. 2.5 4.5 cm.
....................................................................................................................................2. G. sumatranum
1. Gymnostoma nobile (Whitmore) L.A.S. Johnson

(Latin, nobilis = noble, excellent; its graceful growth habit)
l.c. (1982) 84. Basionym: Casuarina nobilis Whitmore, Tropical Rain Forests of the Far East (1975) 134, Ng l.c. 46,
Anderson l.c. 159, Cockburn l.c. 25. Type: A. Cuadra SAN A3292, Sabah, Sipitang Distr., Seungau (holotype K; isotypes A,
KEP, L, NSW, US).
Tree to 40 m tall, 50 cm diameter; bole cylindrical, short buttresses often present; crown bushy
conical, becoming umbrella-shaped. Bark greyish to reddish brown, fissured and scaly. Wood hard,
inner part reddish brown, occasionally with black streaks and often with a wavy figure. Needleshaped twigs with a few lateral branches, the longest unbranched ones mostly exceeding 10 cm long,
11.5 mm thick; internodes 0.51 cm long. Scale-leaves with rounded-acute apex. Male flowers in
compound, cone-like structures. Female flowers in terminal or subterminal capitate inflorescences.
Fruit cones globose, c. 2.2 cm across, usually 1 or more cones terminal or subterminal on lateral,
woody persistent twigs or needle-shaped twigs; each cone with 2030 fertile fruitlets, and whorls of
sterile, undeveloped fruitlets at its upper and basal parts; bracteoles slightly projecting, rounded-acute
at the tips.
Vernacular name. Sarawakrhu ronang (Iban, Malay).
Distribution. Palawan Is., Borneo (Sabah, Sarawak, Brunei, and Kalimantan).
Ecology. Lowland and hill kerangas forest to 1000 m, on sandstone and ultramafic rocks. Occasional
in forest on impoverished sandy infertile soils in coastal areas and near the central zone of peat
swamp forests.
Uses. Commonly planted at roadsides and in gardens. Its wood is useful for construction and fuel.
125
Fig. 3. Gymnostoma sumatranum. A, fruiting twig; B, detail of nodes of a needle-shaped twig; C, detailed surface of woody
twig; DE, fruit cones; F, samara; GH, male inflorescences; I, male flower; J, female inflorescence. (AD and from Chew
CWL 1389, E from Anderson S. 8431, GI from SAN 37835, J from KEP 65596.)
126
2. Gymnostoma sumatranum (Jungh. ex de Vriese) L.A.S. Johnson

(of Sumatra)
Fig. 3.
l.c. (1982) 86. Basionym: Casuarina sumatrana Jungh. ex de Vriese, Tijdschr. Natuur. Gesch. Physiol. 11 (1844) 115,
Mitchell l.c. 119, Backer & Bakhuizen f. l.c. 9, Smythies l.c. 34, Anderson l.c. 159, Cockburn l.c. 24. Type: Junghuhn, s.n.,
Sumatra, Toba Plateau (holotype L; isotype U).
Tree to 20 m tall, 60 cm diameter; bole cylindrical, often twisted or fluted, sometimes with short
buttresses; crown spreading with its main branches ascending and topped by softly bushy needleshaped twigs. Bark greyish brown, sometimes marked with white patches, rough and fissured.
Needle-shaped twigs c. 0.5 mm thick, with many short lateral branches, bushy, the longest
unbranched ones seldom exceeding 10 cm long; internodes short, 0.20.3 cm long. Scale-leaves with
rounded-acute apex. Male flowers in simple, terminal spikes, anthers basifixed. Female flowers in
terminal capitate structures. Fruit cones ellipsoid, c. 2.5 4.5 cm, usually solitary on the apex of
woody persistent twigs, each with 2035 fruitlets; bracteoles strongly protruding, with acute tips.
Vernacular name. Sabah and Sarawakrhu bukit (Malay).
Distribution. Sumatra and Borneo. In Sabah, the species is known from the Tawau Hills, Batanga
near Sipitang, Mt. Tawai and Melinau River, and Mt. Silam, whereas in Sarawak it occurs in many
localities.
Ecology. Apparently confined to hill, ridge and lower montane forests on ultramafic soils and in
kerapah vegetation on sandstone, shale or acidic soils, at 6001800 m. Grows well when planted in
fertile soil.
Uses. Attractive ornamental tree suitable for road sides and gardens.
127
128
__________________________________________________________________________________________
CHLORANTHACEAE
John B. Sugau
Sandakan, Malaysia
Merrill, EB (1921) 209, PEB (1929) 41; Masamune, EPB (1942) 231; Anderson, CLTS (1980) 162; Verdcourt, FM 1, 10
Monoecious or dioecious herbs, shrubs or trees; tissues typically aromatic. Nodes of stem and branch
typically conspicuously swollen. Leaves simple, decussate or sometimes clustered together in
pseudowhorls; blade pinnately veined, serrate or dentate, the teeth often thickened at the tips; petioles
often connected by an interpetiolar ridge at the base or joined into a distinct sheath. Inflorescences
spicate, paniculate, or capitate, axillary or terminal, unisexual or bisexual. Flowers without sepals and
petals; male flowers with or without bract, stamens 13 or numerous (Hedyosmum) and organised in
a cone-like structure; female flowers exposed or enclosed by a cup-shaped bract, with a perianth-like
structure attached to the ovary, ovary usually inferior, 1-locular, ovules solitary, orthotropous,
pendulous, with two-layered integuments and thick nucellus, style short or absent, stigmas truncate, 2lipped, depressed subcapitate, or rarely linear or clavate; bisexual flowers with an anther-bearing
structure attached to one side of the ovary. Fruit a fleshy drupe, ovoid, obovoid or globose to
ellipsoid, small, free or in a cluster enclosed by the bract; endocarp hardened and crustaceous. Seeds
subglobose, without aril; embryo minute, cotyledons variable or scarcely developed; endosperm
copious, oily or not.
Distribution. Four genera with about 80 species, distributed mainly in the tropics but extending south
to New Zealand (Ascarina) and north to Japan, China, Korea and E Russia (Chloranthus and
Sarcandra). In Sabah and Sarawak, four genera with four species, of which only Ascarina
philippinensis reaches tree-size.
Ecology. Mostly found in moist habitats, some ascending to about 2650 m (in upper montane forest
on Mt. Kinabalu in Sabah). Flowers are pollinated by wind or insects; fruits and seeds are dispersed
by birds.
Taxonomy. The Chloranthaceae was placed in the Piperales by Lindley (1821) who first described
the family. Although this has been widely accepted and followed, Smith (1972) assigned the
Chloranthaceae to its own order, Chloranthales. It is considered one of the most primitive of extant
Angiosperm families, with a fossil pollen record going back as far as the early Cretaceous.
129
Key to genera
1. Petioles joined at the base to form a conspicuous membranous interpetiolar sheath. Inflorescence
with many conspicuous ovate to leaf-like bracts, 0.51 cm long. Flowers
unisexual...............................................................................................................................
Hedyosmum Swartz
Prod. Veg. Ind. Occ. (1788) 84; Swamy, J. Arn. Arb. 34 (1953) 402; Endress, Bot. Jahrb. 91
(1971) 39; Verdcourt, FM 1, 10 (1986) 143.
About 30 species, largely distributed in the New World from Mexico to Peru, Brazil, and the
W Indies.
Monoecious or dioecious herbs, shrubs or treelets; stems smelling of ginger-root when
crushed; stipules forming an interpetiolar sheath with comb-like margin. Leaves decussate,
chartaceous. Inflorescences unisexual or bisexual, axillary or terminal, sometimes united with
the stem near the base. Male flowers in cone-like, solitary or paniculate structures, filaments
very short or absent, anthers numerous, 2-locular, locules parallel, opening lengthwise.
Female flowers in capitate or paniculate clusters; perianth tube joined to the ovary, limb very
short, 3-toothed; style very short or absent, stigmatic at apex, stigmas truncate, rarely linear or
clavate, often caducous. Fruits free or in a cluster enclosed by the bracts, ellipsoid, globose or
ovoid, often 3-angular, sometimes crowned with persistent perianth-like lobes; pericarp juicy,
endocarp hard. Seeds 3-angular, pale yellow.
One species (H. orientale) occurs in E Asia extending from S China to Malesia (Sumatra,
Borneo, and Sulawesi). Uncommon in Sarawak, not yet recorded in Sabah. In Sarawak, in hill
and ridge forests in the Hose mountains, Bt. Kajang, Bt. Sarpandai, Bt. Kenawang, and the
Usun Apau plateau.
Petioles not joined to form any sheath (at most connected by an interpetiolar ridge or a low
coriaceous sheath). Inflorescence with tiny inconspicuous bracts, less than 0.5 cm long. Flowers
unisexual or bisexual.........................................................................................................................2
2. Leaf apex acute to shortly cuspidate; leaf margin recurved when dry; lateral veins close together
(not more than 10 mm apart). Inflorescences to 1.52 cm long. Flowers
unisexual...............................................................................................................................Ascarina
Leaf apex acuminate-caudate; leaf margin not recurved when dry; lateral veins well-spaced
(mostly more than 10 mm apart). Inflorescences longer than 2 cm. Flowers bisexual, with a male
part (the anther-bearing organ) attached to the ovary........................................................................3
3. Leaves finely pustulate all over the lower surface, margins typically finely serrate. Infructescence
branches very fine (less than 0.5 mm thick), distal parts not zig-zag. Male part with a 3-lobed
organ bearing 4 anthers...........................................................................
Chloranthus Swartz
Phil. Trans. Roy. Soc. 77 (1787) 359; Merrill, EB (1921) 209, PEB (1929) 41; Masamune
EPB (1942) 231; Swamy, J. Arn. Arb. 34 (1953) 375; Verdcourt, FM 1, 10 (1986) 129.
About 200 species, distributed from Japan, China, E Russia to India, Sri Lanka and Malesia.
Shrubs or treelets. Leaves decussate or sometimes clustered together in pseudowhorls;
stipules minute, subulate. Inflorescence a compound spike with opposite branches; bracts
tiny, boat-shaped. Flowers: male parts basally attached to the ovary or enveloping it,
130
CHLORANTHACEAE (SUGAU)
anthers introrse, without filaments, variously arranged; female parts exposed, stigma truncate.
Fruits fleshy, usually white; endocarp thin, fibrous. Seeds subglobose, narrowed at base,
minutely apiculate.
In Borneo, one species (C. erectus). In Sabah and Sarawak, scattered in the lowland, hill and
montane forests, to 2500 m. Also in Brunei and Kalimantan.
Leaves not pustulate on the lower surface, margins typically set with coarse teeth. Infructescence
branches coarse (11.5 mm thick), distal parts noticeably zig-zag. Male part a narrow clavate
structure bearing 2 anthers...................................................................
Sarcandra Gardner
Calc. J. Nat. Hist. 6 (1846) 348; Swamy & Bailey, J. Arn. Arb. 3 1(1950) 127; Verdcourt,
Kew Bull. 39 (1984) 66, FM 1, 10 (1986) 123.
Two species, distributed from India and Sri Lanka to Myanmar, Indo-China, China, Japan and
Malesia.
Monoecious shrubs or treelets; stems and branches without vessels. Leaves decussate or
sometimes clustered in pseudowhorls; stipules minute, subulate, attached to the inconspicuous
interpetiolar sheath. Inflorescence a spike-like panicle, terminal or axillary; bracts boatshaped, inconspicuous. Flowers: male parts attached to one side of the ovary at about a third
from the top, anthers introrse, opening lenghtwise; ovary ovoid, style absent, stigma
depressed subcapitate. Fruits with succulent pericarp. Seeds ellipsoid, testa membranous.
Only one species (S. glabra subsp. brachystachys var. brachystachys) in Borneo (Sabah). In
Sabah, mostly found in damp habitats, at 10001500 m.
ASCARINA J. R. Forst. & G. Forst.

(Greek, askaris = intestinal worm; the anther shape)
Char. Gen. (1775) 59, l.c. ed. 2 (1776) 117; Anderson, CLTS (1980) 162; Verdcourt, FM 1, 10 (1986) 137; Whitmore,
Tantra & Sutisna, CLK 1 (1989) 45.
Small to medium-sized trees, monoecious or dioecious; all parts glabrous. Stipules very small. Leaves
decussate, mostly elliptic to obovate; margins frequently serrate or crenate, teeth often gland-tipped;
lateral veins close together (not more than 10 mm apart) and subparallel for the most part.
Inflorescences compound spikes with several spreading branches, bracteate. Flowers: males
bracteate, reduced to 15 subsessile or sessile anthers, each with 2 parallel linear locules, opening
lengthwise; females consisting of an exposed sessile ovoid-globose ovary, without outer bracts or
with 2 often caducous inner bracts (and if the flowers are paired then also with bracteoles), stigmas
sessile, truncate or 2-lipped or horseshoe-shaped. Infructescence branches coarse (11.5 cm thick).
Fruits ovoid or obovoid, with thin succulent pericarp; epidermis without stomata. Seeds ovoid,
flattened, with smooth testa.
Distribution. About 12 species, distributed from Madagascar to the Pacific Islands, New Caledonia
and New Zealand. In Malesia (Borneo, Philippines, Sulawesi and New Guinea), 4 species. In Sabah
and Sarawak, one species.
Ecology. In lower montane or more often in upper montane rain forest at 15003300 m, sometimes in
lowland and hill forest.
131
Fig. 1. Ascarina philippinensis. A, flowering (male) leafy twig; B, a portion of male inflorescence; C, leaf margin showing
the thickened, gland-tipped teeth; D, fruiting leafy twig; E, infructescence; F, older female inflorescence; G, ovary in
longitudinal section. (AC from SAN 134198, DG from Kokawa & Hotta 5989.)
132
CHLORANTHACEAE (SUGAU)
Ascarina philippinensis C. B. Robinson
Fig. 1.
Philip. J. Sc. 4 (1909) Bot. 70; Merrill, En. Philip. 2 (1923) 22; Smith, J. Arn. Arb. 57 (1976) 409; Anderson l.c. 162;
Verdcourt l.c. (1986) 139; Whitmore, Tantra & Sutisna l.c. 45. Type: Williams 2541, Philippines, Mindanao, Mt. Apo
(holotype PNH, destroyed; isotypes K, NY). Synonym: A. reticulata Merr., Philip. J. Sc. 12 (1917) Bot. 263, l.c. (1923) 22.
Small to medium-sized tree, to 18 m tall, c. 35 cm diameter. Bark rough, flaky to slightly to distinctly
fissured, white to dark reddish brown. Twigs easily broken. Stipules linear to ovate-boat-shaped, 1
2.5 mm long, often caducous; leafless internodes often present, with sheaths to 6.5 mm long. Leaves
coriaceous, glossy on the upper side, somewhat paler on the lower side; 411 x 26 cm; base cuneate,
margin crenate with thickened teeth, apex acute to shortly cuspidate; midrib prominent, finely
longitudinally ridged, lateral veins 1214 pairs, subparallel for the most part, fine and slightly raised
on both sides; petioles 0.52 cm long. Inflorescences 1.52 cm long; bracts ovate, inconspicuous,
1.53 x 12 mm. Flowers: males whitish or pale greenish yellow, with 2 collateral stamens, anthers
2.54 x 11.5 mm, the projecting connective subacuminate; females congested, usually separated by
not more than 2 mm; ovary green. Fruits obovoid, 23 x 1.52 mm, with spicy aroma, turning purpleblack when ripe; endocarp pale yellow. Seeds 1.52 mm long.
Distribution. Borneo, Philippines, Sulawesi and New Guinea. In Sabah and Sarawak, principally in
the mountains; also in Kalimantan.
Ecology. Lower montane to upper montane rain forest, common at 15002650 m on Mt. Kinabalu.
Flowering in June, July and September; fruits ripen in JuneAugust.
133
134
__________________________________________________________________________________________
CRYPTERONIACEAE
J.T. Pereira
Sandakan, Malaysia
Merrill, EB (1921) 452; Masamune, EPB (1942) 545; Backer & Bakhuizen f., FJ 1 (1965) 257; van Beusekom-Osinga & van
Beusekom, Blumea 22 (1975) 255; Shea in Cockburn, TS 1 (1976) 72; van Beusekom-Osinga, FM 1, 8 (1977) 187;
Anderson, CLTS (1980) 164; Keating, Ann. Missouri Bot. Gard. 71, 3 (1984) 801; van Vliet & Baas, Ann. Missouri Bot.
Gard. 71, 3 (1984) 783; Patel et al., Ann. Missouri Bot. Gard. 71, 3 (1984) 858; Corner, WSTM 3rd ed. 1 (1988) 222; Ng,
TFM 4 ( 1989) 78; Whitmore, Tantra & Sutisna CLK 2, 1 (1990) 50; Mentink & Baas, Blumea 37 (1992) 189.
Trees. Twigs with swollen nodes marked by a transverse line, internodes finely ridged or winged.
Leaves simple, opposite, pinnately veined; stipules absent (except in saplings). Inflorescences
terminal and axillary panicles, few- to many-branched. Flowers bisexual or unisexual (plant
dioecious), radially symmetrical, 45-merous; sepals valvate, triangular, persistent or caducous;
petals present or absent (in Crypteronia), if present, valvate or rarely imbricate, alternate with the
sepals, in bud hood-shaped and enveloping the stamens, caducous; stamens inflexed in bud, persistent
or caducous, in one or two epipetalous whorls, filaments sometimes very short, connective broadly to
narrowly divergent, conduplicate or not, sometimes with a dorsal tubercle or a large outgrowth,
anthers adnate, linear to suborbicular, lengthwise dehiscent, introrse; ovary superior or inferior, 2
4(5)-carpellate, 16-locular, septa not or rarely only partly connate, style terminal, terete, mostly
persistent, stigmas capitate to punctate, ovules inserted horizontally or vertically on the septum, 1 to
many per locule, anatropous, placentation parietal, septal or basal. Fruit a chartaceous to woody
capsule, subglobose to ellipsoid, impressed at the septum, loculicidally dehiscent into 26 valves,
sometimes with persistent style and stigma at the apex (Crypteronia). Seeds 1 to many, compressedellipsoid, located basally, apically, centrally or laterally within its membranous wing; raphe running
freely within the wing; embryo straight.
Distribution. Three genera with about 12 species, distributed in tropical SE Asia and Malesia. In
Sabah and Sarawak, three genera with 10 species.
Ecology. Frequent in lowland to lower montane forest, up to about 1500 m. The monotypic
Dactylocladus is confined to peat swamp forest.
Taxonomy. The definition, delimitation and systematic position of Crypteroniaceae in the order
Myrtales is controversial. In the earlier system of classification (Bentham & Hooker f., Gen. Pl. 1
(1862) 782), the genus Crypteronia on which the family was founded, was originally placed under
Lythraceae. A. De Candolle (Prodr. 16 (1868) 677), assigned this genus to its own family, the
Crypteroniaceae, a decision accepted by Hutchinson (Fam. Fl. Pl. 1 (1959) 167; Gen. Fl. Pl. 2 (1967)
135
33), and also by Cronquist (Int. Syst. Class. Fl. Pl. (1981) 633). Niedenzu (in Engler & Prantl, Nat.
Pfl. Fam. 3, 7 (1898), however, placed Crypteronia in the tribe Crypteronioideae of the Blattiaceae (=
Sonneratiaceae).
Van Beusekom-Osinga & van Beusekom (l.c.), and van Beusekom-Osinga (l.c.), recognized 5 genera
in the family, namely Alzatea (confined to Peru and Bolivia), Axinandra, Crypteronia and
Dactylocladus (in tropical SE Asia and Malesia), and Rhynchocalyx (restricted to tropical S Africa).
Van Vliet (Blumea 27 (1981) 395) and Thorne (in Young & Seigler (eds.), Phytochemistry and
angiosperm phylogeny (1981) 227) proposed the inclusion of Crypteroniaceae as a subfamily in the
Melastomataceae, a suggestion not accepted by Mentink & Baas (Blumea 37 (1992) 216). Johnson &
Briggs (Ann. Missouri Bot. Gard. 71, 3 (1985) 732) excluded Rhynchocalyx from the Crypteroniaceae
and assigned it to its own family, the Rhynchocalycaceae. Concurrently, Graham (Ann. Missouri Bot.
Gard. 73, 3 (1985) 775) segregated Alzatea from Crypteroniaceae and placed it in its own family, the
Alzateaceae.
Brummitt (Vasc. Pl. Fam. & Gen. (1992) 549), in agreeing with the concepts of Graham and Johnson
& Briggs, recognized Crypteroniaceae as a well-defined family comprising Axinandra, Crypteronia,
and Dactylocladus. In the present account, Brummitts delimitation of the Crypteroniaceae is
followed.
Key to genera
1. Leaf apex typically obtuse, rarely acute; intercostal veins almost invisible on both surfaces.
Functional stamens very short, 0.51 mm long. Plants typically of peat swamp
forest.......................................................................................................................3. Dactylocladus
Leaf apex acute, acuminate to caudate or cuspidate; intercostal veins typically prominent on one
or both sides of the leaf. Functional stamens longer, 1.54 mm long. Plants of other
habitats.............................................................................................................................................2
2. Flower subtended by 1 bract; petals absent; stamens or staminodes as many as sepals, persistent;
ovary superior, 24-locular, ovules many per locule. Fruits small, chartaceous, typically with
persistent style and stigma. Seeds many.....................................................................2. Crypteronia
Flower subtended by 3 bracts, the two lateral ones often tiny; petals present; stamens twice as
many as sepals or petals, non persistent; ovary inferior, (4)6-locular, ovules 1 or 2 per locule.
Fruits large, woody, style and stigma caducous. Seeds few..........................................1. Axinandra
1. AXINANDRA Thwaite
(Greek, axine = axe, andros = male; the hatchet-shaped stamens)

In Hooker, J. Bot. 6 (1854) 66; Ridley, FMP 1 (1922) 821; van Beusekom-Osinga & van Beusekom, Blumea 22 (1975) 262;
van Beusekom-Osinga, FM 1, 8 (1977) 187; Ng, TFM 4 (1989) 78; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 50.
Synonyms: Axinandra sect. Naxiandra Baill., Adansonia 12 (1876) 85; Naxinandra (Baill.) Krasser in Engler & Prantl,
Nat. Pfl. Fam. 3, 7 (1893) 197.
Trees. Leaves: apex acuminate to caudate or cuspidate; midrib flattened above, prominent beneath;
lateral veins ascending and joining to form looped marginal veins, intercostal veins
136
CRYPTERONIACEAE (PEREIRA)
prominent on one or both sides, glabrous. Inflorescences terminal or axillary, erect, few-branched;
axes more or less angular; bracts triangular; bracteoles 3 per flower, the two lateral ones often tiny.
Flowers bisexual, receptacle pubescent in flower, glabrous in fruit; sepals (4)5, caducous in fruit;
petals, (4)5, flimsy, valvate-connate or imbricate-conduplicate or imbricate-contorted, more or less
connate or coherent and dropping simultaneously as a unit; stamens (8)10, 1.53.5 mm long,
caducous, filaments terete to more or less flattened, connective divergent, with distinct dorsal
tubercle, anthers broadly linear at apex; ovary inferior, (2)3-carpellate, (4)6-locular, glabrous,
ovules 1 or 2 per locule, inserted basally to the septum, style (subulate-)terete, sometimes protruding
from mature buds, glabrous, caducous in fruit, stigmas punctate. Fruits large, woody, globose to
ellipsoid, splitting into 26 valves in the apical part that protrudes from the enlarged receptacle;
receptacle with distinct encircling rim; valves triangular, coarse, glabrous. Seeds few, depressedellipsoid, located basally and obliquely in the narrowly oblong wing; raphe running along the embryo.
Distribution. Four species, distributed from Sri Lanka to Peninsular Malaysia and Borneo. In Sabah,
only A. coriacea is recorded, whereas in Sarawak, A. alata and A. beccariana are known.
Ecology. Lowland to lower montane forest to 1200 m.
Key to Axinandra species

1. Internodes winged. Leaves oblong; lateral veins 1214 pairs. Bracteoles of median flowers 11.5
mm long..............................................................................................................................1. A. alata
Internodes not winged. Leaves elliptic, ovate-elliptic to ovate; lateral veins 612 pairs. Bracteoles
of median flowers, 0.251 mm long..................................................................................................2
2. Leaves coriaceous to subcoriaceous; acumen obtuse; intercostal veins prominent and distinctly
reticulate on both sides. Inflorescences 2.55 cm long; ultimate branches 1.54 cm long; main
axis 12 mm thick; bracts 0.251 mm long..................................................................3. A. coriacea
Leaves chartaceous; acumen acute; intercostal veins obscure and faintly reticulate on both sides.
Inflorescences 520 cm long; ultimate branches 4.57 cm long; main axis 0.30.5 mm thick;
bracts 14 mm long..................................................................................................2. A. beccariana
1. Axinandra alata Baill.
(Latin, alatus = furnished with wing-like extensions; the internodes)

Adansonia 12 (1876) 86; Merrill, EB (1921) 452; Masamune, EPB (1942) 545; van Beusekom-Osinga & van Beusekom l.c.
263; van Beusekom-Osinga l.c. 187. Type: Beccari PB 3651, Borneo, Sarawak (holotype FI; isotypes K, P).
Tree. Twigs glabrous; internodes terete at base, becoming acutely quadrangular upwards with 4
gradually widening wings towards the nodes; wings 13 mm wide, ending in acute, often upwardcurved processes. Leaves drying brown-green above, brown below, chartaceous; oblong, 5-10 x 35
cm; base rounded, apex cuspidate, acumen 0.30.7 cm long; midrib flattened on the upper leaf
surface, raised on the lower surface, lateral veins 1214 pairs, distinctly looping towards the margin,
137
flattened above, raised below, intercostal veins prominent on both surfaces, distinctly reticulate,
glabrous; petioles 35 mm long, 12 mm thick, glabrous. Inflorescences axillary, 815 cm long, main
axes 11.2 mm thick, ultimate branches 57 cm long, rachis pubescent; bracts c. 1 mm long, sparsely
pubescent, caducous; bracteoles of median flowers 11.5 mm long, narrowly triangular, densely hairy,
persistent; pedicels 12 mm long, c. 0.5 mm thick, pubescent. Flowers: sepals 0.50.8 x 11.5 mm,
triangular, glabrous inside, pubescent outside; petals 3.54 x 11.5 mm; filaments 1.52 mm long;
styles 35 mm long, protruding from mature buds. Fruits not seen.
Distribution. Endemic to Borneo (Sarawak only), very uncommon and only known from the type.
2. Axinandra beccariana Baill.

l.c. 85; Merrill l.c. 452; Ridley l.c. 826; Masamune l.c. 545; van Beusekom-Osinga & van Beusekom l.c. 263; van
Beusekom-Osinga l.c. 187; Ng l.c. 78. Type: Beccari PB 3458, Borneo, Sarawak (holotype FI; isotypes K, P). Synonyms:
A. maingayi C.B. Clarke in Hooker f., Fl. Br. Ind. 2 (1879) 581; A. borneensis Bakhuizen f., Rec. Trav. Bot. Neerl. 40 (1943)
preprint 332.
Tree. Twigs glabrous to minutely pubescent; internodes not winged. Leaves glabrous, drying greenbrown on both surfaces, chartaceous; elliptic to ovate, 510 x 25 cm; base cuneate, apex acuminate
to acute, acumen 0.30.5 cm long, tip acute; midrib flattened on the upper leaf surface, raised on the
lower leaf surface, lateral veins 812 pairs, looping towards the margin, flattened above, slightly
raised below, intercostal veins obscure and faintly reticulate on both sides, glabrous; petioles 58 mm
long, 12 mm thick. Inflorescences axillary to terminal, 520 cm long, main axes 0.30.5 mm thick,
ultimate branches 4.57 cm long, rachis sparsely pubescent to glabrous; bracts 14 mm long,
subglabrous, persistent, bracteoles of median flowers 0.51 mm long, narrowly triangular to
triangular; pedicels 12.5 mm long, c. 0.5 mm thick. Flowers: sepals c. 0.5 x 1 mm, triangular,
glabrous inside, subglabrous to glabrous outside; petals c. 3 x 1 mm; filaments 11.5 mm long; styles
c. 2 mm long, hardly or not protruding from mature buds. Fruits 1.52 x 11.5 cm; valves c. 0.8 cm
long. Seeds not seen.
Distribution. Peninsular Malaysia (one collection, Maingay 65412, from Melaka) and Borneo
(known from Sarawak only).
3. Axinandra coriacea Baill.
Fig. 1.
l.c. 85; Merrill l.c. 452; van Beusekom-Osinga & van Beusekom l.c. 264; van Beusekom-Osinga l.c. 187. Lectotype (van
Beusekom-Osinga & van Beusekom, 1975): Beccari PB 2622 Borneo, Sarawak (FI; isolectotypes K, P).
Tree to 55 m tall, 90 cm diameter; buttresses to 2 m tall and 0.5 m wide. Bark grey-brown, smooth to
flaky; inner bark pale brown to red-brown. Sapwood whitish to brown. Twigs glabrous to sparsely
138
Fig. 1. Axinandra coriacea. A, flowering leafy twig; B, old flowers with petals and stamens fallen off; C, mature flower bud
with style protruding; D, mature open flower with petals fallen off, showing style and stamens; E, fruit. (AB from S. 39235,
CD after FM 1, 8 (1977) 187, fig. 11, E from S. 37302.)
139
hairy; internodes not winged. Leaves drying greenish to dark brown above, brown below, coriaceous
to subcoriaceous, glabrous above, sparsely hairy to glabrous below; elliptic to ovate-elliptic, 5
15 x 1.57 cm; base cuneate to rounded, apex acuminate to caudate, acumen 0.51 cm long, tip obtuse;
midrib flattened on the upper leaf surface, raised on the lower leaf surface, lateral veins 612 pairs,
looping towards the margin, flattened above, slightly raised below, intercostal veins prominent and
distinctly reticulate on both sides; petioles 37 mm long, 12 mm thick, glabrous or sparsely hairy.
Inflorescences axillary to terminal, 2.55 cm long, main axes 12 mm thick, angular, ultimate
branches 1.54 cm long, rachis densely pubescent; bracts 0.251 mm long, subglabrous, bracteoles of
median flowers 0.251 mm long, triangular; pedicels 12 mm long, c. 1 mm thick. Flowers: sepals
0.51 x 11.5 mm, triangular, glabrous inside, densely to sparsely pubescent outside; petals c. 6 x 1.5
mm; filaments 1.52 mm long; styles 35 mm long, protruding from mature buds. Fruits 1.24 x 13
cm; valves 56, 0.51.7 x 0.30.7 cm. Seeds: wings broadly obtuse at apex.
Distribution. Endemic to Borneo (Sabah, Sarawak and Kalimantan).
Ecology. In lowland to lower montane forest up to 1200 m. In Sabah, it is recorded mainly in forest
on ultramafic soil (Mt. Silam, Bidu-Bidu Hills, Bukit Kiabau, etc.), whereas in Sarawak it is found in
mixed dipterocarp and kerangas forests, up to 1200 m.
2. CRYPTERONIA Blume
(Greek, kruptos = hidden, eros = love; probably alluding to the small flowers)
Bijdr. Fl. Ned. Ind. (1826) 1151; Ridley, FMP 1 (1922) 821; Keng, OFMSP (1969) 151; van Beusekom-Osinga & van
Beusekom, Blumea 22 (1975) 258; Shea in Cockburn, TS 1 (1976) 73; van Beusekom-Osinga, FM 1, 8 (1977) 191;
Anderson, CLTS (1980) 164; Corner, WSTM 3rd ed. 1 (1988) 222; Whitmore, Tantra & Sutisna, CLK 1 (1989) 51; Ng,
TFM 4 (1989) 79; Pereira & Wong, Sandakania 6 (1995) 41. Synonyms: Henslowia (non Blume) Wall., Pl. As. Rar. 3
(1831) 13, t. 221; Quilamum Blanco, Fl. Filip. 1 (1837) 851.
Medium-sized to large trees; buttresses sometimes present. Twigs with nodes somewhat swollen,
glabrous to pubescent. Stipules absent or very small. Leaves coriaceous, subcoriaceous to
chartaceous, apex acute, acuminate to caudate; midrib flattened or sunken above, lateral veins
distinctly to vaguely looping, intercostal veins prominent on one or both sides, glabrous to pubescent.
Inflorescences terminal and/or axillary panicles, branching 13 orders, many-flowered; bracts 1 per
flower. Flowers bisexual or unisexual (only in C. paniculata); sepals persistent even in fruit; petals
absent; stamens or staminodes 5, as many as sepals, persistent, functional stamens 1.54 mm long,
glabrous, filaments flattened, connective tissue divergent, anthers apically or laterally attached to the
connective; ovary superior, 24-locular, ovules many per locule, inserted horizontally or vertically
(basally) to the septum, style terete, glabrous to hairy, stigmas capitate to punctate. Fruits small,
hardly larger than the flowers, chartaceous, typically with persistent styles and stigmas, subglobose to
ovoid-ellipsoid, impressed at the septa, splitting loculicidally into 24 valves, styles and stigmas
adhering at apex. Seeds many, tiny, ovoid-ellipsoid, inserted on the septum or at the base of the
septum; wing membranous, acute to obtuse at base or apex; raphe running closely along the embryo.
140
Distribution. Seven species with 2 varieties, distributed in tropical and subtropical SE Asia from
Assam to Bengal, Myanmar, Thailand, Indo-China and Malesia. In Sabah and Sarawak, six species
are recorded.
Ecology. Lowland to lower montane forest, up to about 1500 m.
Key to Crypteronia species

1. Leaf intercostal veins obscure on both sides. Flowers unisexual (plants dioecious); ovary 2locular. Seeds inserted on the septum, the membranous wing with a broadly rectangular base and
a tapered or acute apex................................................................6. C. paniculata (var. paniculata)
Leaf intercostal veins prominent on one or both sides. Flowers bisexual; ovary 34-locular. Seeds
inserted at the base of the septum; the membranous seed wing with a tapered or acute to narrowly
rectangular
or
obtuse
base
and
a
broadly
obtuse
to
acute
apex...................................................................................................................................................2
2. Midrib distinctly sunken on the upper leaf surface. Inflorescence bracts 2.59 mm long..................
...................................................................................................................................1. C. borneensis
Midrib flat on the upper leaf surface. Inflorescence bracts 0.51.5 mm long...................................3
3. Leaves oblong to ovate-oblong; lateral veins 1020 pairs. Inflorescences branching to the 3rd
order, rarely to only the 2nd order; main axis 35 mm thick................................5. C. macrophylla
Leaves narrowly elliptic, elliptic to ovate-elliptic; lateral veins typically not more than 10 pairs.
Inflorescences branching to the 1st or 2nd order (typically to the 1st order); main axis 13 mm
thick...................................................................................................................................................4
4. Leaf apex acuminate to caudate, acumen 0.51.8 cm long; leaf base distinctly cuneate; intercostal
veins distinctly reticulate to tessellate. Petioles slender, only 11.5 mm thick. Ovary glabrous.
Fruits ellipsoid...............................................................................................................2. C. elegans
Leaf apex shortly acute, acumen if present shorter, 0.10.4 cm long; leaf base cordate to rounded;
intercostal veins sparsely reticulate. Petioles thicker, 27 mm thick. Ovary sparsely to densely
minutely papillose to pubescent. Fruits subglobose to ovoid...........................................................5
5. Inflorescences with ultimate branches (15)2040 cm long; main axis 23 mm thick; pedicels
0.10.5 mm long, densely pubescent. Sepals broadly spread open in the mature flower, sparsely to
densely pubescent outside. Styles sparsely pubescent at base, glabrous at upper part. Seed wing
tapered at apex.............................................................................................................4. C. griffithii
Inflorescences with ultimate branches not more than 20 cm long; main axis slender, 12 mm
thick; pedicels 11.5 mm long, glabrous. Sepals connate, forming a dome-shaped structure even
in the mature flower, glabrous outside. Styles glabrous throughout. Seed wing broadly obtuse at
apex..........................................................................................................................3. C. glabriflora
1. Crypteronia borneensis J.T. Pereira & Wong

(of Borneo)
Sandakania 6 (1995) 45. Type: Tarmiji & Talib SAN 90941, Borneo, Sabah, Beaufort (holotype SAN; isotype K).
141
Medium-sized tree to 25 m tall, 45 cm diameter. Bark greyish brown, smooth; inner bark brown to
greyish. Sapwood yellowish white to brown. Twigs glabrous. Leaves drying dark brown to greenbrown on both sides, coriaceous to subcoriaceous; broadly ovate, ovate-elliptic to elliptic, 1026 x 4
20 cm; base cordate, rounded to cuneate, apex acute to acuminate, acumen 0.51.0 cm long; midrib
distincly sunken on the upper surface, distinctly raised on the lower leaf surface, lateral veins 815
pairs, distinctly looping towards the margin, sunken to flattened above, distinctly raised below,
intercostal veins obscure above, prominent below, laxly reticulate, glabrous; petioles 0.52 cm long,
25 mm thick, glabrous. Inflorescences terminal or in the axils of fallen leaves on older branches,
1425 cm long, branching to the 2nd or 3rd order, main axis 24 mm thick, ultimate branches 515
cm long, rachis pubescent; bracts 2.59 mm long, triangular, subglabrous, persistent. Flowers
bisexual, bracteoles (0.5) 12 mm long, narrowly triangular to subulate, sparsely hairy, persistent;
pedicels 0.52 mm long, c. 0.5 mm thick, minutely papillose or pubescent; sepals triangular, densely
tomentose inside, minutely papillose or tomentose outside, 0.51.5 x 0.51.5 mm; stamens 1.52.5
mm long, filaments 12 mm long, anthers 0.250.5 x 0.20.8 mm; ovary 3(4)-locular, 12 mm
across, styles 13.5 mm long, minutely densely papillose or tomentose, stigmas capitate. Fruits
subglobose, papillose to densely tomentose. Seeds inserted at the base of the septum; wing broadly
obtuse at apex, tapered or acute at base.
Vernacular name. Sabahtampasu (Dusun).
Distribution. Endemic to Borneo (Sabah, Sarawak and Brunei).
Ecology. In lowland primary mixed dipterocarp forest, on clay-rich, well-drained alluvium and sandy
soils, to c. 600 m. Frequently near river banks.
2. Crypteronia elegans J.T. Pereira & Wong

(Latin, elegans = elegant; the inflorescences)
Sandakania 6 (1995) 48. Type: Paie S. 36393, Borneo Sarawak , 7th Division, Kapit, Bukit Bakar (holotype SAN; isotypes
K, KEP, L, MO, SAR).
Medium-sized tree to 27 m tall, 75 cm diameter; buttresses to 1.5 m tall and 1.5 m wide. Bark reddish
brown to blackish, flaky; inner bark red with deep yellow. Sapwood orange- straw-coloured. Twigs
glabrous. Leaves drying dark brown to brown on both sides, subcoriaceous; narrowly elliptic, 7
12 x 25 cm; base cuneate, apex acuminate to caudate, acumen 0.51.8 cm long; midrib flattened on
the upper leaf surface, raised on the lower leaf surface, lateral veins 810 pairs, looping towards the
margin, flattened at both sides, intercostal veins prominent and distinctly reticulate or tessellate on
both sides, glabrous; petioles 12 cm long, 11.5 mm thick. Inflorescences terminal and axillary, up
to 24 cm long, branching to the 1st or 2nd order, main axis 12 mm thick, ultimate branches 718 cm
long, rachis puberulose; bracts 0.51 mm long, early caducous. Flowers bisexual; bracteoles 0.51.5
mm, narrowly triangular, usually caducous; pedicels 0.51.5 mm long, c. 0.5 mm thick, puberulose;
sepals triangular, sparsely puberulose outside, densely puberulose inside, 0.51 x 0.751 mm; stamens
1.52.5 mm long, filaments 12 mm long, anthers c. 0.5 mm across; ovary 3-locular, 1.52 mm
across, styles 13 mm long, glabrous, stigmas punctate. Fruits ellipsoid, glabrous to subglabrous.
Seeds inserted at the base of the septum; wing tapered at base and apex.
142
Distribution. Endemic to Borneo (Sarawak and Brunei); not yet recorded for Sabah. In Sarawak,
recorded at Kelabit Highlands and Bukit Bakar, Kapit, while in Brunei, it is found in the Temburong
District.
Ecology. In lowland mixed dipterocarp forest and secondary forest, to 1100 m, mainly on low hills
and ridges.
3. Crypteronia glabriflora J.T. Pereira & Wong

(Latin, glaber = without hair, florus = flowers)
Fig. 2.
Sandakania 6 (1995) 50. Type: Dransfield JD 7225, Brunei, Temburong, Bukit Belalong (holotype SAN; isotypes BRUN,
K, KEP).
Tree to 20 m tall, 40 cm diameter. Bark reddish brown, cracking; inner bark reddish. Sapwood pale
yellowish brown. Twigs glabrous. Leaves drying dark brown to brown above, pale brown below,
coriaceous; ovate-elliptic to elliptic, 720 x 38 cm; base cordate to rounded, apex shortly acute,
acumen 0.10.4 cm long; midrib flattened on the upper leaf surface, slightly raised on the lower leaf
surface, lateral veins 810 pairs, looping towards the margin, flattened above, flattened to slightly
raised below, intercostal veins prominent and laxly reticulate on both sides, glabrous; petioles 0.41
cm long, 23 mm thick, glabrous. Inflorescences terminal and in the axils of fallen leaves on older
branches, 1220 cm long, branching to the 1st or 2nd order, main axis 12 mm thick, ultimate
branches 520 cm long, rachis puberulose; bracts 0.51 mm long, triangular, glabrous, caducous.
Flowers bisexual; bracteoles 0.51 mm long, narrowly triangular, pubescent, persistent; pedicels 1
1.5 mm long, c. 0.5 mm thick, glabrous; sepals triangular, connate, forming a dome-like structure
even in the mature flower, glabrous on both sides, slightly hairy at margin below, 11.5 x 11.5 mm;
stamens 34 mm long, filaments 2.53.5 mm long, anthers c. 0.5 mm across; ovary 3-locular, 11.5
mm across, densely minutely papillose, styles 24.5 mm long, glabrous, stigmas punctate. Fruits
subglobose, minutely papillose. Seeds inserted at the base of the septum; wing broadly obtuse at
apex, narrowly obtuse at base.
Distribution. Endemic to Borneo (Sarawak and Brunei only); not yet recorded in Sabah. In Sarawak,
it occurs on Mt. Pagon, Limbang and in Brunei in the Temburong District.
Ecology. Mixed dipterocarp forest to 820 m, on clay soil overlying sandstone.
4. Crypteronia griffithii C.B. Clarke
(W. Griffith, 18101845, physician and botanist in India and Malaya)

In Hooker f., Fl. Br. Ind. 2 (1879) 574; Ridley l.c. 822; van Beusekom-Osinga & van Beusekom l.c. 260; van BeusekomOsinga l.c. 196; Corner l.c. 222; Ng l.c. 80; Whitmore, Tantra & Sutisna l.c. 51. Type: Griffith 2513, Malacca (holotype K;
isotype L).
Medium-sized to large tree to 40 m tall, 70 cm diameter; buttresses to 0.6 m tall and 0.6 m wide. Bark
brownish to grey, smooth to fissured or flaky; inner bark brown to red. Sapwood white to yellow or
pale brown. Twigs glabrous. Leaves drying green-brown to brown or dark brown on both sides,
coriaceous; elliptic to ovate-elliptic, 940 x 415 cm; base cordate to rounded, apex shortly acute,
143
Fig. 2. Crypteronia glabriflora. A, flowering leafy twig; B, mature open flower; C, young fruit with the stamens still
persistent. (All from Dranfield JD 7225.)
144
acumen 0.10.4 cm long; midrib flattened on the upper leaf surface, raised on the lower leaf surface,
lateral veins 810(14) pairs, flattened above, raised below, looping towards the margin, intercostal
veins prominent and laxly reticulate beneath, glabrous; petioles 0.51.5 cm long, 37 mm thick,
glabrous. Inflorescences in the axils of fallen leaves on older branches, 2045 cm long, branching to
the 1st or 2nd order, main axis 23 mm thick, ultimate branches (15)2040 cm long, rachis densely
pubescent; bracts 0.51 mm long, triangular, hairy, persistent to caducous. Flowers bisexual;
bracteoles 0.51 mm long, triangular with slender tip; pedicels 0.10.5 mm long, 0.51 mm thick,
densely pubescent; sepals triangular, broadly spread open in the mature flower, densely to sparsely
pubescent on both sides, 12 x 11.5 mm; stamens 23.5 mm long, filaments 1.53 mm long, anthers
c. 0.5 mm across; ovary 3-locular, 1.52 mm across, sparsely pubescent, styles (2)34 mm long,
sparsely pubescent at base, glabrous at upper part, stigmas punctate. Fruits subglobose, sparsely
pubescent. Seeds inserted at the base of the septum; wing tapered or acute at apex, narrowly
rectangular at base.
Vernacular names. Sabahrambai-rambai (Bajau). Sarawakubah semut (Iban).
Distribution. Myanmar, C Sumatra, Peninsular Malaysia and Borneo. In Sabah and Sarawak,
common in most districts; also in Kalimantan.
Ecology. Scattered in primary forest, up to 600 m; often on yellow to brown-blackish sandy soil,
sometimes on clayey soil. In Peninsular Malaysia and Borneo, the species often forms an association
with Cladomyrma maschwitzii ants.
Uses. The wood has been reported to be durable and used in house-building.
5. Crypteronia macrophylla Beus.-Osinga

(Greek, makros = large, phullon = leaves)
Blumea 22 (1975) 261; van Beusekom-Osinga l.c. 198; Whitmore, Tantra & Sutisna l.c. 51. Type: Ashton S. 19372, Borneo,
Sarawak, Anap, Ulu Muput Kanan, Bt. Naoung (holotype L; isotypes K, SAN, SAR).
Medium-sized tree to 25 m tall, 60 cm diameter. Bark greyish to pale yellowish brown, smooth to
flaky; inner bark orange. Twigs glabrous. Leaves drying greenish brown above, brown below,
coriaceous; oblong to ovate-oblong, 1845 x815 cm; base cordate to slightly rounded, apex acute,
acumen c. 0.5 mm long; midrib flattened on the upper leaf surface, distinctly raised on the lower leaf
surface, lateral veins 1020 pairs, flattened above, raised below, distinctly to vaguely looping towards
the margin, intercostal veins prominent and distinctly reticulate to tessellate on both sides, glabrous;
petioles 11.5 cm long, 25 mm thick, glabrous. Inflorescences in the axils of leaves on older nodes,
3040 cm long, branching to the 2nd or 3rd order, main axis 35 mm thick, rachis pubescent; bracts
0.51 mm long, triangular, short-hairy, caducous. Flowers bisexual; bracteoles 0.51 mm long,
narrowly triangular; pedicels 1.52 mm long, c. 0.5 mm thick, densely thinly pubescent; sepals
triangular, 0.51 mm across, pubescent on both surfaces; stamens 1.53 mm long, filaments 12.5
mm long, anthers 0.51 x 0.51 mm; ovary (3)4-locular, 12.5 mm across, densely pubescent to
thinly tomentose, styles 0.23 mm long, densely pubescent, stigmas punctate. Fruits subglobose,
pubescent. Seeds inserted at the base of the septum; wing acute at base and apex.
145
TEE FLORA OF SABAH AND SARAWAK VOL. 2 (1996)
Vernacular names. Sarawakbedulang semut, ubah semut, ubah samak (Iban); teletang (Kelabit).
Distribution. Endemic to Borneo (Sarawak and Kalimamtan only). In Sarawak, known from Anap,
Rejang, Kapit, Ulu Melinau, Balleh, Baram and Kuching areas; in Kalimantan, recorded from West
Kutei, Pontianak and Sg. Kahayan.
Ecology. Primary lowland to hill forest, on clay, igneous, sandstone to basalt soils, to 1200 m,
sometimes near river banks. Young branches are often inhabited by ants.
6. Crypteronia paniculata Blume
(Latin, paniculatus = panicle; the inflorescence)

l.c. 1151; Ridley l.c. 821; van Beusekom-Osinga & van Beusekom l.c. 259; van Beusekom-Osinga l.c. 194; Anderson l.c.
165; Corner l.c. 224; Whitmore, Tantra & Sutisna l.c. 51; Ng l.c. 80. Type: Blume, s.n., Java (holotype L).
var. paniculata
Tree to 36 m tall, 105 m diameter; buttresses to 1.5 m tall, 1.2 m wide. Bark brown to grey-brown,
finely fissured to scaly; inner bark brown. Sapwood light brown to yellowish white. Twigs glabrous.
Leaves drying deep brown to brown-green above and greenish-brown below, chartaceous; ovateelliptic to elliptic, 621 x 310(12) cm; base cuneate to sometimes rounded, apex shortly to narrowly
acuminate, acumen 0.21.5 cm long; midrib flattened on the upper leaf surface, raised on the lower
leaf surface, lateral veins 710 pairs, flattened, vaguely looping towards the apex, intercostal veins
obscure on both sides, laxly reticulate, glabrous; petioles 0.20.8 cm long, 12 mm thick, glabrous.
Inflorescences axillary and/or terminal, 930 cm long, branches to the 1st or 2nd order, main axis 1
1.5 mm thick, ultimate branches 1020 cm long, rachis puberulose; bracts c. 0.5 mm long, triangular,
glabrous, caducous. Flowers unisexual (plants dioecious); bracteoles (0.5)11.5 mm long, narrowly
triangular to subulate, glabrous; pedicels 11.5 mm long, c. 0.5 mm thick, glabrous; sepals triangular,
densely to sparsely puberulose inside, papillose to subglabrous outside, 0.51.5 x 0.51 mm; males:
stamens 24 mm long, filaments 1.53.5 mm long, anthers c. 0.5 x 0.51.5 mm; pistillodes 0.51 mm
long; females: staminodes 0.51 mm long; ovary 2-locular, 0.51.5 mm across, papillose to
puberulose, styles (0.8)11.5 mm long, papillose, stigmas capitate. Fruits subglobose, slightly
impressed at the septum, minutely papillose to puberulose. Seeds inserted on the septum; wing acute
or tapered at apex, broadly rectangular at base.
Vernacular names. Sabahkinkidon mantok (Dusun Kadazan). Sarawakubah semut (Iban).
Distribution. Continental SE Asia; from E Bengal to Assam, Andaman Islands, Myanmar, Peninsular
Malaysia, Borneo (Sabah, Sarawak, Kalimantan), Sumatra, Java, Lesser Sunda Islands, and the
Philippines. In Sabah, more common in the west coast; in Sarawak, recorded in the 1st, 2nd and 7th
Divisions. In Kalimantan, documented for W Kutei.
Ecology. Frequent in primary and secondary hill forests, on sandy clay soil, to c. 1500 m.
146
Uses. The timber is claimed to be hard and durable and is utilized in house-building in W Java and S
Sumatra. It is also used in making cart-wheels and also for fuel. In the Philippines, the bark has been
used to treat blisters. In Java, the young shoots are eaten as salad.
3. DACTYLOCLADUS Oliv.
(Greek, daktulos = finger, klados = twigs; the branching mode of the twigs)
jongkong (Malay)
In Hooker, Ic Pl. (1895) t. 2351; Merrill, EB (1921) 452; Masamune, EPB (1942) 547; Brown, FTSB (1955) 261; Smythies,
CST (1965) 89; Burgess, TBS (1966) 391; van Beusekom-Osinga & van Beusekom, Blumea 22 (1975) 261; van BeusekomOsinga, FM 1, 8 (1977) 199; Anderson, CLTS (1980) 165; Whitmore, Tantra & Sutisna, CLK 1 (1989) 51.
Large trees; without buttresses. Twigs quadrangular, swollen at the nodes. Leaves coriaceous, apex
typically obtuse, rarely acute; midrib thick, flattened on the upper leaf surface, raised on the lower
leaf surface, lateral veins vague and flattened on both sides, looping towards the margin, intercostal
veins almost invisible on both surfaces, glabrous. Inflorescences terminal and/or axillary, erect,
poorly branched; bracts and bracteoles caducous, minute; pedicels short. Flowers bisexual; sepals
triangular, persistent; petals unguiculate, blades irregularly lobed, suborbicular, pubescent on the
margin, enclosing stamens in bud, caducous; functional stamens 0.51 mm long, persistent, glabrous,
anthers oblong to broad-linear, filaments terete; ovary half-inferior, lower part adnate to the
receptacle, puberulose, (3)45-locular, ovules 3 per locule, basally inserted in between the septa,
septa not connate, styles pubescent, persistent, stigmas capitate. Fruits broad-ellipsoid, small,
pubescent, (3)45-valved. Seeds narrow-ellipsoid, flat, situated centrally to the membranous wing,
apex obtuse to irregularly shaped, broader than the base; raphe running close to the embryo.
Distribution. Monotypic, so far known with certainty from Borneo only. A doubtful record has been
reported from Irian Jaya (cf. van Beusekom-Osinga l.c. 201).
Ecology. Common in lowland peat swamp forest.
Dactylocladus stenostachys Oliv.
(Greek, stenos = narrow, small, stachys = spike; the inflorescence)
Fig. 3.
In Hooker l.c. t. 2351; Merrill l.c. 452; Masamune l.c. 457; Brown l.c. 261; Smythies l.c. 89; Burgess l.c. 391; van
Beusekom-Osinga & van Beusekom l.c. 262; van Beusekom-Osinga l.c. 200; Anderson l.c. 165; Whitmore, Tantra &
Sutisna l.c. 51. Lectotype (van Beusekom-Osinga & Beusekom, 1975): Haviland 2916, Borneo, Sarawak (K; isolectotype
L).
Large tree to 40 m tall, 150 cm diameter; trunk base of big trees developing masses of small
adventitious roots; pneumatophores (sharp-pointed aerial roots) yellowish, appearing at the base or
around the tree. Bark brown to grey, smooth in immature tree, reddish brown, fissured to scaly in
mature tree; inner bark yellowish to brown. Sapwood yellow to white, with small canal-like
147
perforations. Twigs glabrous. Leaves drying greenish above, brown to greenish below; elliptic, ovate,
obovate to spathulate, 3.510(15) x 27 cm; base cuneate to slightly rounded; lateral veins 1015
pairs; petioles 0.10.7 cm long, 0.10.3 mm thick, glabrous. Inflorescences 616 cm long, branching
to the 1st order, rachis pubescent; bracteoles persistent, triangular, c. 0.5 mm long; pedicels 0.251
mm long, densely puberulose. Flowers: sepals puberulose on both surfaces, c. 0.5 x 0.51 mm;
stamens 0.51 mm long, filaments 0.250.5 mm long, anthers 0.50.8 x 0.5 mm; ovary c. 1 mm
across, styles 0.52 mm long, stigmas capitate. Fruits 34 mm across; valves 12 x 11.5 mm. Seeds
23 mm long, inserted at the base of the septum.
Vernacular names. Sabahjongkong (Malay), medang miang (Kedayan), medang tabak (Brunei
Malay), tanjiong awan (Malay). Sarawakgatal (Murut), jongkong (Malay), medang jongkong
(Murut), merebong (Iban).
Distribution. Endemic to Borneo (Sabah, Sarawak, Brunei and Kalimantan).
Ecology. Common in all types of peat swamp and mixed swamp forests. Occasionally in kerangas
forest. In the peat swamp forest, it is common and dominant constituent of the padang keruntum
phasic communities. Flowering and fruiting throughout the year, particularly from May to August.
Uses. The timber is moderately hard and moderately heavy, with density ranging from 495 to 610
kg/m3 air dry. The heartwood is not differentiated from the sapwood and is light brown when fresh
and darkening to pink-brown or red-brown. Texture is fine and even, with straight or slightly
interlocking grains. The timber is easy to work and produces a smooth and lustrous surface. It is not
durable when exposed to weather or in contact with the ground, but suitable for weather-boarding,
flooring, partitioning, utility furniture, concrete shuttering, and for veneers in plywood manufacture
(Burgess l.c.; Wong, DMT (1982) 82).
149
150
__________________________________________________________________________________________
CTENOLOPHONACEAE
Runi S. Pungga
Kuching, Malaysia
Exell & Mendona, Consp. Fl. Angol. 1, 2 (1951) 248; Hutchinson, Fam. Fl. Pl. (1959) 265; van Hooren & Nooteboom,
Blumea 29 (1984) 559, FM 1, 10 (1988) 629; Brummitt, Vasc. Pl. Fam. & Gen. (1992) 550.
A monogeneric family closely allied to the Ixonanthaceae and Linaceae. Ctenolophonaceae differ
from the Ixonanthaceae in having indumentum of stellately tufted hairs, opposite leaves, interpetiolar
stipules, hypogynous flowers with twisted, caducous, extrastaminal disc, a 2-forked style with each
fork topped by a capitate stigma, and a soft-hairy arillode. From the Linaceae, Ctenolophonaceae
differ in their indumentum, interpetiolar stipules, extrastaminal disc, free filaments, 2-forked style and
well-developed soft-hairy arillode.
CTENOLOPHON Oliv.
(Greek, ctenos = comb, lophorus = crested; the crested arillode)

Trans. Linn. Soc. 28 (1873) 516; Beccari, Malesiana 1 (1877) 119; King, J. As. Soc. Beng. 64, 2 (1895) 105; Merrill, EB
(1921) 313; Ridley, FMP 1 (1922) 423; Masamune, EPB (1942) 356; Cockburn, TFM 1 (1972) 306, TS 1 (1976) 204;
Anderson, CLTS (1980) 239; van Hooren & Nooteboom, Blumea 29 (1984) 560, FM 1, 10 (1988) 629; Whitmore, Tantra &
Sutisna, CLK 2, 1 (1990) 208; Brummitt, Vasc. Pl. Fam. & Gen. (1992) 550.
Trees. Indumentum of stellately tufted and simple hairs. Stipules interpetiolar, boat-shaped, cadacous.
Leaves simple, opposite, entire, coriaceous. Inflorescences terminal panicles or lateral racemes.
Flowers bisexual, hypogynous; sepals 5, thick, persistent; petals 5, thick, linear, twisted in bud,
caducous; disc extrastaminal; stamens 10, filaments free, short and long, inserted halfway on the inner
side of the disc; ovary 2-locular, with 2 ovules in each locule; style apically 2-forked, each fork
topped by a capitate stigma. Fruit a woody capsule, splitting irregularly down one side. Seeds
solitary, pendulous, with soft-hairy arillode.
Distribution. Two species, distributed from W Africa to Malesia (Peninsular Malaysia, Sumatra,
Borneo, Philippines and New Guinea). One species in Sabah and Sarawak.
Ctenolophon parvifolius Oliv.
Fig. 1.
(Latin, parvus = small, folium = leaf)
151
CTENOLOPHONACEAE (RUNI PUNGGA)
l.c. 516, t. 43, f. 17; Beccari l.c. 120; King l.c. 106; Merrill l.c. 313; Ridley l.c. 423; Masamune l.c. 356; Cockburn, Gard.
Bull. Sing. 24 (1969) 4, l.c. (1972) 306, l.c. (1976) 205; Anderson l.c. 239; van Hooren & Nooteboom l.c. (1988) 631;
Whitmore, Tantra & Sutisna l.c. 208. Type: Maingay KD 382, Malaya (holotype K; isotype L). Synonym: C. grandifolius
Oliv. l.c. 517, t. 43, f. 810, Beccari l.c. 119, King l.c. 106, Merrill l.c. 313, Ridley l.c. 403, Masamune l.c. 356.
Tree to 35 m tall, 40 cm diameter; buttresses steep and sharp. Bark grey to grey-brown, finely
cracked or scaly, hard and lenticellate. Leaves thick-coriaceous, drying grey-brown; elliptic to
elliptic-ovate, 59 x 26 cm; base rounded to broadly cuneate, apex broadly acute to obtusely
acuminate; midrib slightly sunken on the upper leaf surface, raised on the lower leaf surface, lateral
veins looping and joining at the margin without forming a distinct marginal vein, intercostal veins
reticulate and slightly raised on the lower leaf surface. Inflorescences dense panicles, terminal or in
the axils of upper leaves. Flowers: sepals ovate, whitish hairy; petals linear-oblong, 59 mm long;
filaments 3.56 mm long; ovary ovoid, hairy, style 5.56.5 mm long. Fruits ellipsoid, 1525 x 510
mm, hairy, apex whitish hairy, sharply acute, with persistent clavate swollen stalk and sepals. Seeds
ellipsoid, to 1.8 cm long, brown, shiny, surrounded by crest-like arillode.
Vernacular names. Sabahbesi-besi, obah (Dusun). Sarawakkrian, litoh (Iban).
Distribution. Sumatra, Peninsular Malaysia, Borneo (Sabah, Sarawak), the Philippines and New
Guinea.
Ecology. Found in swamp forest and on hills, on rather poor soils over sandstone and ultramafic
rocks. Common in peat swamp forest.
Uses. The timber is hard and very durable, and good for house construction and general purpose
timber.
153
154
__________________________________________________________________________________________
DAPHNIPHYLLACEAE
Balu Perumal
World Wide Fund for Nature (WWF),
Backer & Bakhuizen f., FJ 1 (1963) 505; Huang, Taiwania 11 (1965) 57, l.c. 12 (1966) 137; Hsuan Keng, OFMSP (1969)
415; Whitmore, TFM 1 (1972) 181; Anderson, CLTS (1980) 165.
A monogeneric family comprising about 9 species, widely distributed in E Asia and Malesia
extending from India to C China, Japan, and Malesia. In Sabah and Sarawak, represented by 2
species.
Taxonomy. The genus Daphniphyllum was formerly considered to be a member of the
Euphorbiaceae, but now it is placed in a family (Daphniphyllaceae) and an order (Daphniphyllales) of
its own. The relationship of this small family is thought to be with the Hamamelidaceae or perhaps
with the Magnoliaceae. Embryological, palynological and wood anatomical studies confirmed its
exclusion from the Euphorbiaceae and its alliance with the Hamamelidaceae. Chemically, the family
is known to produce iridoid glycosides and the daphniphylline series of alkaloids. So far this alkaloid
seems to be restricted to the family.
DAPHNIPHYLLUM Blume
(Greek, daphne = laurel, phullon = leaf;
having leaves resembling those of Daphne, Thymelaeaceae)
Bijdr. Fl. Ned. Ind. 17 (1826) 1153; Merrill, EB (1921) 336; Ridley, FMP 3 (1924) 234; Masamune, EPB (1942) 395;
Backer & Bakhuizen f. l.c. 505; Huang l.c. (1965) 57, l.c. (1966) 137; Whitmore l.c. 181; Anderson l.c. 165.
Dioecious trees or shrubs. Leaves spirally arranged, simple, entire, pinnately veined; without
stipules. Inflorescences axillary racemes; bracts basal, caducous. Flowers unisexual; calyx 26lobed, lobes imbricate in bud, basally united; petals absent. Flowers: males with 512 stamens,
radially arranged, anthers introrse or extrorse, dehiscing through longitudinal slits; female: ovary
superior, incompletely 2(34) locular, styles short, united basally, stigmas thick, recurved, ovules 2
per locule, subapical, pendulous, anatropous; staminodes sometimes present. Fruit a drupe. Seed 1;
endosperm thick, oily; embryo tiny, apical.
Ecology. In a wide variety of habitats, from the lowlands to mountain summits; in kerangas,
limestone and montane forests, in both secondary and primary forests.
155
Uses. Some species, e.g., Daphniphyllum laurinum, are known to be poisonous due to the presence of
the alkaloid daphniphylline, found in the bark and seeds. In Banka (Sumatra, Indonesia), a decoction
of the root has been reported as a medicine for diarrhoea and thrush (Heyne, Nutt. Pl. Ned. Ind. 3
(1917) 80).
Key to Daphniphyllum species

Tree to 15 m tall, restricted to montane forest. Leaves usually coriaceous, margin recurved. Calyx in
female flower not persistent. Anthers usually oblong or ellipsoid, extrorse, filaments usually long.
Fruit with prominent recurved styles...............................................................................D. glaucescens
Shrub or treelet to 3 m tall, restricted to lowland forest. Leaves usually chartaceous, margin slightly
undulating. Calyx in female flower persistent. Anthers crescent-shaped, introrse, filaments short. Fruit
with minute styles.......................................................................................
D. laurinum (Benth.) Baill.
tud. Gen. Euph. (1858) 565; Merrill l.c. 336; Ridley l.c. 234; Masamune l.c. 395; Huang l.c. (1966) 139;
Whitmore l.c. 181; Anderson l.c. 165. Basionym: Gouphia laurina Benth. in Hooker, Kew Jour. 6 (1854) 9. Type:
Wallich 1836, Singapore (holotype K; isotypes BM, E). Synonym: Daphniphyllum bancanum Kurz, Tijdschr. Ned.
Ind. 27 (1864) 188, Ridley l.c. 234, Masamune l.c. 395.
Leaves alternate or often spirally clustered towards the ends of twigs; blades 1128 3.512 cm;
intercostal veins faintly scalariform. Flowers: males with the calyx 34-lobed, pistillodes rarely
present; females with a bell-shaped, 4-lobed calyx, style very short, spreading, staminodes rarely
present. Fruits 712 57 mm, surface finely warty.
W Malesia; widely distributed in Sumatra, Peninsular Malaysia, and Borneo; also cultivated in Java. In
Sabah and Sarawak, common in forest on sandy soils or on sandstone rocks by the seashore, as well as
inland on limestone at 500900 m.
Daphniphyllum glaucescens Blume
Fig. 1.
(Greek, glaukos = covered with whitish or pale bloom; the lower leaf surface)
l.c. 1153; Merrill l.c. 336; Masamune l.c. 395; Huang l.c. (1966) 161; Whitmore l.c. 181; Anderson l.c. 165. Type: Blume
1908, Java, Mt. Salak (holotype L; isotypes NY, US).
Tree to 15 m tall, 30 cm diameter. Twigs terete, brown, marked with many triangular leaf scars.
Leaves occasionally clustered towards the ends of twigs; blades usually coriaceous, rarely
chartaceous, dark green above, greyish brown, glaucous and papillose below; narrowly elliptic,
elliptic, elliptic-oblong, or rarely lanceolate, 816 35 cm; base acute, cuneate or rounded, margin
recurved; midrib prominent below, somewhat impressed above, lateral veins 712 pairs, intercostal
veins reticulate; petioles narrowly triangular in cross-section, 14.5 cm long, c. 0.7 mm thick.
Inflorescences: males slender racemes, 38 cm long; females racemose, 36 cm long, pedicels terete,
616 mm long, c. 0.2 mm thick. Flowers: males purple, calyx bell-shaped, 45-lobed, filaments 0.2
0.6 mm long, anthers usually oblong or ellipsoid, 0.61.5 0.20.5 mm, extrorse; females: calyx not
persistent, styles spread out upwards or coiled, as long as the ovary, staminodes present. Fruits
ellipsoid, obovoid or ovoid, 614 510 mm, ripening blackish purple, surface uneven with a few
tiny knobs; persistent styles prominent and recurved.
156
DAPHNIPHYLLACEAE (BALU PERUMAL)
Fig. 1. Daphniphyllum glaucescens subsp. borneense. A, fruiting leafy twig; B, fruit; C, longitudinal section of fruit; D,
leafy twig with a male inflorescence; E, male flower; F, stamen; G, leafy twig with female inflorescences; H, ovary; I, ovary
in longitudinal section. (AC from SAN 56346, DF from SAN 54275, GI from SAN 21965.)
157
Key to subspecies
Leaves narrowly elliptic, length 2.63.8 times the width; apex acute-acuminate. Fruits obovoid or
ovoid, 68 57 mm................................................................................................................
subsp. dichotomum Huang
l.c. (1966) 190. Lectotype (Huang, 1966): Clemens, s.n., Borneo, Sabah, Ranau, Mt. Kinabalu
(UC; isolectotypes A, BO, L, NY).
Endemic to Borneo. In Sabah, known from a few collections from Mt. Kinabalu (Clemens
10946, 30912, 32385; SAN 79578); and in Sarawak, from Mt. Api, Ulu Melinau, Baram
Distr. (S. 30884). Also known in Kalimantan. In lower montane and limestone hill forest, at
10001500.
Leaves elliptic to elliptic-oblong, length 1.52.7 times the width; apex rounded, cuspidate or abruptly
acuminate. Fruits ellipsoid 1012 810 mm................................................................
subsp. borneense (Stapf) Huang
Fig.1.
l.c. (1966) 192. Basionym: Daphniphyllum borneense Stapf, FMK (1894) 224, Merrill l.c.
336, Anderson l.c. 165. Type: Haviland 1070, Borneo, Sabah, Mt. Kinabalu (holotype K;
isotypes BM, L, UC).
Endemic to Borneo. In Sabah, common in the lower and upper montane forest on Mt.
Kinabalu, at 1,5004000 m; in Sarawak, known only from Mt. Murud, the Hose mountains
and Usun Apau plateau, at 23002600 m. Also reported from Kalimantan.
Vernacular name. Sarawakteradam bukit (Iban).
Distribution. E Asia extending from Sri Lanka to Japan and Malesia.
Ecology. In Sabah and Sarawak, common on mountain ridges or slopes of lower and upper montane
forest (usually associated with Magnolia and Symplocos) and subalpine vegetation, at 10004000 m.
Flowering from February to July.
158
__________________________________________________________________________________________
EPACRIDACEAE
S.P. Lim
Sandakan, Malaysia
Hooker f., Fl. Br. Ind. 3 (1882) 477; Merrill, EB (1921) 467; Masamune, EPB (1942) 577; Sleumer, FM 1, 6 (1964) 422;
Whitmore, TFM 2 (1972) 32.
Small trees or shrubs. Leaves spirally arranged, simple, stiff and coriaceous, with several longitudinal
main veins; stipules none. Inflorescence an axillary and/or terminal raceme, with very short-stalked
flowers, each flower subtended by small bracts. Flowers bisexual or unisexual (plants dioecious),
radially symmetrical, 45-merous, often fragrant; sepals free, coriaceous, imbricate and persistent;
corolla tube present, lobes valvate or imbricate; stamens 45, rarely 2, free, inserted in the corolla
tube, alternate with the corolla lobes, included or exserted, anthers 1-locular, dehiscing by a
longitudinal slit, becoming inverted during development; disc entire or 5-lobed; ovary superior, 110locular, ovules one per locule, placentation axile. Fruit with a single hard stone (pyrene) containing
several locules, or with several 1-locular stones. Seeds small; embryo straight; endosperm fleshy.
Distribution. About 31 genera and 400 species, extending from Myanmar and Thailand, Indo-China,
Malesia to the Pacific islands, Australia, New Zealand, Hawaii and the SW part of South America;
Australia and Tasmania have the most species. In Malesia, 3 genera with 18 species, 4 of which are
known also from outside Malesia. In Sabah and Sarawak, 2 genera with 4 species are known.
Ecology. In open coastal vegetation on white sands, kerangas forest and forest on ultramafic soils; in
the lowlands and on mountains up to 4000 m. In Australia, the Epacridaceae assume the importance
of the Ericaceae in other continents, although in Malesia, they can be found growing with Ericaceae.
Taxonomy. Epacridaceae is distinguishable from Ericaceae in having free sepals (those in Ericaceae
generally connate), 45 stamens rather than 10, with unilocular anthers splitting longitudinally rather
than bilocular anthers opening by pores.
Key to genera
Leaves without distinct stalk or with a short stalk up to 1 mm long; veins rather
inconspicuous on the lower surface, either very faint and numerous, or 13 unbranched veins
parallel for the most part and running to the apex with other branched veins diverging toward
the margins. Fruit containing a single hard stone, with (2)5(10) locules (chambers) each
with a seed.................................................................................................................1. Styphelia
159
Leaves with distinct stalk longer than 1 mm; veins very conspicuous on the lower leaf
surface, with 13 unbranched veins running to the apex, only the central one straight, the
other veins curved and almost parallel to the leaf margins. Fruit containing 810 separable
stones, each equivalent to an ovary locule with one seed.................................2. Trochocarpa
1. STYPHELIA Sm.
(Greek, stuphelos = set closely together; the leaves)
Sp. Bot. New Holl. (1793) 45; Hooker f., Fl. Br. Ind. 3 (1882) 477; Merrill, EB (1921) 467; Masamune, EPB (1942) 577;
Sleumer, Blumea 12 (1963) 145, FM 1, 6 (1964) 424; Whitmore, TFM 2 (1972) 32.
Shrubs or small trees, monoecious (with bisexual flowers) or dioecious; young branches minutely
puberulent, becoming glabrous and fissured in older parts. Leaves without a distinct stalk or only a
short stalk not more than 1 mm long; venation inconspicuous on the lower side, the veins either very
faint and numerous or 3 parallel veins running straight to the apex, the other veins branched and
diverging toward the margins. Inflorescence a short raceme or a solitary flower terminating a short
peduncle; bracteoles 2, opposite, ovate, keeled dorsally, margin ciliate. Flowers 5-merous; sepals with
ciliate margin; corolla lobes valvate, recurved in open flower, inner side hairy on the upper half,
outside glabrous; stamens 5, attached at or near the top of the corolla tube, filaments short, filiform;
ovary (2)5(10)-locular, style attenuate, stigma a slight swelling. Fruit globose to ellipsoid with a
hard stone, containing(2)5(10) locules, base covered with persistent sepals (in Bornean taxa). Seed
one in each locule.
Distribution. About 130 species. Eight species in Malesia (one species widespread, also in IndoChina, Myanmar and Thailand), one species in Marianas, six species throughout the Pacific area, 13
species in New Caledonia, eight species in New Zealand and the majority in Australia. Three species
are found in Sabah, while only a single species has been documented for Sarawak.
Ecology. On seashores, and acidic and peaty soil in montane forest up to alpine elevation. Often
gregarious.
Uses. Some fruits are edible, while leaves and roots of some species are used as medicine.
Key to Styphelia species

1. Leaves generally large, (15)2555(85) mm long, (2)510(11) mm wide. Veins very
fine and hardly visible.................................................................................. 2. S. malayana
Leaves much smaller, typically up to 14 mm long, 3 mm wide. Veins visible, with 13
main veins running straight to the apex, the other veins branched and diverging toward
the margins...........................................................................................................................2
2. Leaves with a short distinct stalk not more than 1 mm long, glaucous on the lower
surface; leaf apex acute without a stiff point. Racemes 36 mm long, with 49(13)
flowers each.
Flowers unisexual.......................................................................................3. S. suaveolens
160
EPACRIDACEAE (LIM)
Leaves sessile, not or very slightly glaucous on the lower surface; leaf apex acuminate
with a stiff sharp slender point. Racemes 13 mm long, with only 12 flowers each.
Flowers bisexual...........................................................................................1. S. abnormis
1. Styphelia abnormis (Sond.) J. J. Sm.
Fig. 1.
(Latin, abnormis = abnormal or unusual)

Ic. Bog. 4 (1910) 82; Merrill l.c. 467; Masamune l.c. 577; Holthuis & Lam, Blumea 5 (1942) 224; Sleumer l.c. (1963) 151,
l.c. (1964) 433. Basionym: Leucopogon abnormis Sond. in Lehm., Pl. Preiss. 1 (1845) 325. Type: DUrville & Lesson, s.n.
1825, New Guinea (holotype P). Synonyms: Leucopogon acuminatus (non R. Br.) Duperrey, Voy. Coquille Bot. Atlas
(1826) t. 53; Leucopogon lancifolius Hook. f., Ic. Pl. (1852) t. 898; Leucopogon moluccanus Scheff., Nat. Tijd. Ned. Ind. 32
(1873) 419; Leucopogon malayanus Jack var. moluccanus (Scheff.) Kurz, J. As. Soc. Beng. 46, 2 (1877) 217; Styphelia
lancifolia (Hook. f.) J.J. Sm. l.c. (1910) 82; Styphelia moluccana (Scheff.) J.J. Sm. l.c. 82.
Shrub or small tree. Leaves sessile, upper surface waxy and glabrous, lower surface not or only very
slightly glaucous, glabrous; narrowly elliptic, lanceolate or oblanceolate, (6)1014(16) x (1)23(
4) mm; base narrow and truncate, subglabrous, margin finely serrulate-ciliate, apex acuminate with a
conspicuous stiff sharp slender point; 13 unbranched veins running to the apex, the other (outer)
veins on each side branched and diverging towards the margins. Inflorescences axillary, minutely
puberulent, 13(4) mm long, with 12, rarely 3 flowers; bracteoles 2, opposite, ovate, 1.32 x 11.5
mm, keeled dorsally ending in a short mucro, margin ciliate, glabrous on surface. Flowers bisexual;
sepals ovate, 23 x 1.21.5 mm, glabrous on both surfaces; corolla tube c. 2 mm long, tube hidden by
sepals, corolla lobes 1.52 x 0.60.8 mm, inner surface densely covered with pale long soft hairs;
stamens with filament 0.5 mm long, anthers oblong, 0.60.7 mm long, slightly exserted; ovary
globose, c. 1 mm across, glabrous, styles 1.21.5 mm long, glabrous. Fruits broadly ellipsoid with
truncate apex, to 3 x 1.52 mm, tipped with a persistent style, glabrous; stone with 45 locules.
Distribution. Confined to Malesia: Borneo, SE Sulawesi, Maluku and New Guinea. In Borneo, it
occurs only in Sabah, recorded for Balambangan Island, Gaya Island, Menggatal and Bukit Padang in
Kota Kinabalu.
Ecology. On sandy beaches (Balambangan Island), among rocks (Gaya Island), and on sand and
degraded sandstone (Menggatal and Bukit Padang, Kota Kinabalu). Flowering and fruiting throughout
the year.
2. Styphelia malayana (Jack) Spreng.

(of Malaya)
Syst. Veg. 4 (1827) Cur. Post. 67 (as malaica); Hooker f. l.c. 477; Merrill l.c. 467; Masamune l.c. 577; Lam, Blumea 5
(1945) 572; Sleumer l.c. (1963) 147, l.c. (1964) 426; Whitmore l.c. 32. Basionym: Leucopogon malayanus Jack, Mal. Misc.
1, 5 (1820) 20. Type: Jack, s.n., Singapore (not preserved).
var. malayana
Sleumer l.c. (1963) 147, l.c. (1964) 426.
161
Fig. 1. Styphelia abnormis. A, flowering leafy twigs; B, detail of leaf venation; C, flower; D, corolla slit open to show
stamens; E, ovary with style and stigma; F, infructescence with two fruits. (AB from Mohd. Shah & Kadim MS 946, CF
from SAN 85553.)
162
EPACRIDACEAE (LIM)
Shrub or small tree. Leaves sessile, upper surface waxy and glabrous or with minute appressed stiff
hairs along the middle, lower surface glaucous, glabrous to puberulent; narrowly elliptic to lanceolate,
(15)2555(85) x (2)510(11) mm; base narrow and truncate, covered in tiny hairs, margin
glabrous to ciliate especially towards the base and apex, apex acute or acuminate with a conspicuous
stiff sharp slender point, glabrous or minutely puberulent; veins numerous, very fine and without
distinct midrib, hardly visible. Inflorescences axillary, densely long-hairy, up to 15 mm long with up
to 10 flowers; bracteoles 2, opposite, ovate, 1.52 x 22.5 mm, keeled dorsally, apex mucronulate,
margin ciliate, glabrous or subglabrous on the surface. Flowers bisexual; sepals ovate to lanceolate,
34.5 x 1.52.5 mm, glabrous or with appressed fine hairs along the middle on the inner surface,
glabrous or subglabrous on the outer surface; corolla tube c. 3 mm long, hidden by or slightly exserted
from the sepals, corolla lobes 2 x 1.5 mm, inner surface covered with rusty tomentose indumentum;
stamens with filament 0.61 mm long, anthers oblong, 0.20.8 mm long, slightly exserted; ovary
globose to ellipsoid, c. 1mm across, glabrous, styles 12.5 mm long, glabrous or covered with dense
flexuous hairs especially at the base. Fruits globose to ellipsoid, 35 x 24 mm, tipped with a
persistent hairy or glabrous style, glabrous; stone with 5 locules.
Vernacular names. Sarawakchuchur atap (Malay), maki china (Malay), melamut (Iban), rusak
(Bidayuh). Bruneiambok gobang (Brunei Dusun), ludang-ludang (Tutong), terindak ati (Brunei
Malay, Kedayan).
Distribution. Myanmar, Thailand, Indo-China, Peninsular Malaysia, Sumatra (including Banka,
Billiton, and Riouw), Anambas Is. and Borneo. In Sabah, it is confined to the northwest parts (Mt.
Kinabalu, Mt. Tambuyukon and Labuan). In Sarawak, recorded only from the 1st Division (Mt.
Santubung, Bako National Park and Bungoh Hill) and 4th Division (Lambir Hill, Murud Hill and Mt
Dulit) but generally present throughout. In Brunei, it has been documented from the Tutong and Belait
districts and the Berakas Forest Reserve in Muara district. Also in Kalimantan.
Ecology. On nutrient-poor dry sandy soils (0150 m), in montane kerangas forest on sandstone (400
2400 m) and ultramafic soils (16002750 m). Flowering and fruiting throughout the year but mainly
in July and August.
Uses. Leaves and roots are used to prepare a concoction drunk for stomach ache and for
administration after childbirth. Fruits are edible.
3. Styphelia suaveolens (Hook. f.) Warb.

(Latin, suaveolens = sweet-smelling; the flower)
In Sarasin, Reisen 2 (1905) 329; Merrill l.c. 468; Masamune l.c. 577; Lam l.c. 571; Sleumer l.c. (1963) 148, l.c. (1964) 428.
Basionym: Leucopogon suaveolens Hook. f., Ic. Pl. (1852) t. 898. Type: Low, s.n., Sabah, Mt Kinabalu (holotype K).
Synonyms: Styphelia philippinensis Merr., Philip. J. Sc. 20 (1922) 419; Leucopogon philippinensis (Merr.) Hosokawa,
Trans. Nat. Hist. Soc. Formosa 30 (1940) 336.
Shrub or small tree. Leaves glabrous and waxy on the upper surface, glaucous, glabrous to slightly
hairy on the lower surface; narrowly elliptic to oblanceolate, (4)814(16) x (1)23 mm; base
cuneate, margin glabrous and sometimes ciliate toward the apex, apex acute without a stiff sharp
slender point; 13 unbranched veins parallel for the most part and running to the
163
apex, and 1 or 2 outer veins on each side branched towards the margin, obscure on the upper leaf
surface; petioles shorter than 1 mm, subglabrous or minutely puberulent up to the leaf base.
Inflorescences terminal, minutely puberulent, 13 in a leaf axil, 36(9) mm long, each with 49(
13) flowers; bracteoles 2, opposite, ovate to triangular, 11.5 x 1.21.8 mm, keeled dorsally, margin
ciliate, surface minutely puberulent or with tiny glistening hairs. Flowers unisexual (plant dioecious);
sepals triangular 22.5 x 21.5 mm, both surfaces glabrous or with tiny glistening hairs all over the
outer surface; corolla tube c. 2 mm long, hidden by sepals, corolla lobes 1.52 x 1 mm, inner surface
densely covered with pale long curly hairs; in male flowers, filaments of stamens c. 1mm long,
anthers oblong, 11.5 mm, exserted, reduced pistil columnar, c. 2 mm long; in female flower,
filaments of stamens c. 0.5 mm long, anthers reduced, c. 0.3 mm long, less exserted compared to
those in the male flower, ovary globose, c. 0.8 mm across, glabrous, styles 0.51 mm long, glabrous.
Fruits subglobose, 2.5 x 2 mm, tipped with a persistent style, glabrous; stone with 25 locules.
Distribution. Borneo, Philippines, Sulawesi, Timor, New Guinea, Solomon Islands, Australia and
New Zealand. In Borneo, it is restricted to Mt. Kinabalu, Sabah.
Ecology. In montane forest at 18004000 m, on ultramafic rock, granite and sandstone. Flowering
throughout the year.
2. TROCHOCARPA R. Br.
(Greek, trocho = wheel-like, karpos = fruit)
Prod. (1810) 548; Sleumer l.c. (1963) 163, l.c. (1964) 436.
Shrubs or small trees. Leaves distinctly stalked; veins conspicuous on the lower surface, with 13
unbranched veins running to the apex, only the central one straight, the others curved and almost
parallel to the leaf margins. Inflorescence a raceme, terminal or axillary or both. Flower bisexual, 5merous; bracts below the flowers 2 (opposite) or 710, margin ciliate; corolla lobes valvate, plane to
recurved, inner suface hairy, outer surface glabrous; stamens 5, attached at or near the top of the
corolla tube, filaments short, filiform; ovary (8)10(11)-locular, styles columnar, stigmas obtuse or
capitate. Fruits globose, with 810 separable stones, each developed from an ovary locule. Seed one
in each locule.
Distribution. About 12 species. A single species in Borneo, one species in Sulawesi, 6 species in
New Guinea and 6 species in Australia with one in both Australia and New Guinea.
Ecology. In open places, up to 600 m in Australia, and NW New Guinea and between 30004000 m
in Sabah, Central Sulawesi and the main range of New Guinea.
Trochocarpa celebica (J. J. Sm.) Steen.
Fig. 2.
(of Celebes = Sulawesi)

In Lam l.c. 573; Merrill l.c. 467; Masamune l.c. 577; Lam l.c. 573; Sleumer l.c. (1963) 166, l.c. (1964)
439. Basionym: Styphelia celebica J. J. Sm., Ic. Bog. 4 (1910) 81. Type: Abendanon, s.n., 1909, Central
164
EPACRIDACEAE (LIM)
Fig. 2. Trochocarpa celebica. A, flowering and fruiting leafy twigs; B, detail of leaf venation; C, inflorescence; D, corolla
slit open to show stamens; E, ovary with style and stigma; F, fruit. (AB from SAN 54715, CF from SAN 82977.)
165
Celebes (holotype BO). Synonyms: Styphelia learmonthiana Gibbs, J. Linn. Soc. Bot. 42 (1914); Trochocarpa
learmonthiana (Gibbs) H.J. Lam, l.c. 574.
Low shrub or tree; young branches minutely puberulent, becoming glabrous and fissured in older
parts. Leaves waxy and glabrous to slightly hairy on the upper surface, not glaucous, glabrous on the
lower surface; elliptic to ovate, (3)57(9) x (1)23(4) mm; base cuneate, glabrous or subglabrous
on the upper surface, margin glabrous to ciliate at the acute leaf apex; venation conspicuous, with 13
unbranched veins running to the apex, only the central one straight, the other (outer) veins on each
side curved and almost parallel to the leaf margins, branched toward the margin; petioles 12 mm
long, glabrous or subglabrous. Inflorescences axillary and terminal, densely puberulent, up to 8 mm
long, with up to 12 flowers; bracteoles 2, opposite, c. 1.5 x 2 mm, ovate, dorsally keeled, ending in
short mucro, imbricate, margin ciliate, surface glabrous. Flowers: sepals ovate to triangular, 22.5
x 1.31.8 mm, glabrous, margin ciliate; corolla tube c. 3 mm long, exserted from the sepals; corolla
lobes 2 x 11.5 mm, inner surface with dense long coarse hairs at the base; stamens with filament c. 2
mm long, anthers linear, 0.81 mm long, very exserted; disc 5-lobed; ovary globose, c. 1 mm across,
glabrous, styles 1.31.7 mm long, columnar, ridged, glabrous. Fruits globose to ovoid, 24 x 35
mm, tipped with persistent style, glabrous, containing 10 separable stones; base covered by persistent
sepals.
Distribution. Borneo and Sulawesi. In Borneo, it is confined to Mt. Kinabalu, Sabah.
Ecology. In montane kerangas or open summit vegetation and in rock crevices. On Mt. Kinabalu, it
occurs between 3000 m and 4000 m. Flowering throughout the year.
166
ERYTHROXYLACEAE
R.C.K. Chung
Kepong, Malaysia
King, J. As. Soc. Beng. 62, 4 (1893) 190 (as Order Lineae, in part); Koorders, Exk. Fl. Java (1912) 415; Merrill, EB (1921)
313; Ridley, FMP 1 (1922) 324 (as Order Lineae, in part); Masamune, EPB (1942) 357; Payens, FM 1, 5 (1958) 543; Backer
& Bakhuizen f., FJ 1 (1963) 440; Hutchinson, Fam. Fl. Pl. 1 (1959) 263, Gen. Fl. Pl. 2 (1967) 605; Cockburn, TFM 1 (1972)
194, TS 1 (1976) 82; Keng, OFMSP (1978) 175; Anderson, CLTS (1980) 171; Ashton, MNDTS 2 (1988) 238; Corner,
WSTM 3rd ed. 1 (1988) 254; Whitmore, Tantra & Sutisna, CLK 1 (1989) 108.
Shrubs or trees. Stipules intrapetiolar, rarely extrapetiolar, completely fused or rarely bifid, often
caducous. Leaves alternate (distichous) or rarely opposite (Aneulophus), simple, entire. Flowers
axillary, solitary or in clusters, bisexual, heterostylous (with short- and long-styled forms), 5-merous,
radially symmetrical; calyx campanulate, lobes 5, imbricate or valvate, persistent; petals 5, free,
caducous, imbricate or involute in bud, usually with a ligular appendage near the base on the inner
side; stamens 10, in two whorls of 5, filaments connate at least at the base, anthers basifixed, 2locular, dehiscing longitudinally; disc absent; ovary superior, mostly 3-carpellate, 3-locular with 2 of
the locules usually sterile, fertile locule 1- or 2-ovulate; styles 3, erect, free or partly connate, stigmas
capitate (often oblique) or clavate; ovules pendulous, anatropous. Fruit a drupe; endocarp hard.
Seeds with or seldom without endosperm; embryo straight.
Distribution. Four genera with about 270 species; Aneulophus (2 species), Nectaropetalum (6
species) and Pinacopodium (2 species) are restricted to Africa, while Erythroxylum (c. 250 species) is
a genus centered mainly in the Andes and the Amazon basin in tropical and subtropical S America. In
Sabah and Sarawak, 4 species of Erythroxylum are known.
Ecology. In primary and secondary forests, to c. 1600 m, especially on slopes of low hills.
Taxonomy. In the past the genus Erythroxylum has been incorporated in different families, viz. the
Malpighiaceae (de Jussieu, Gen. Pl. (1789) 281), Linaceae (Bentham & Hooker, Gen. Pl. 1 (1862)
244, and Ridley l.c. 323), and Erythroxylaceae (Payens l.c., Hutchinson l.c., Keng l.c., and others). In
recent classifications, it is widely accepted that the Erythroxylaceae differ from the Malpighiaceae in
their fruits which are not winged and at maturity do not split into 3 nut-like parts, and from the
Linaceae in their ligulate petals and 3-locular ovary with only one locule fully developing.
167
ERYTHROXYLUM P. Browne
(Greek, erythros = red, xylon = wood)
Civ. Nat. Hist. Jam. 1 (1756) 278; Linnaeus, Syst. Nat. ed. 10, 2 (1759) 1035 (Erythroxylon); King, J. As. Soc. Beng. 62, 4
(1893) 190; O. E. Schulz, Pflanzenr. 29 (1907); Koorders, Exk. Fl. Jav. (1912) 416; Merrill, EB (1921) 313; Ridley, FMP 1
(1922) 324; Masamune, EPB (1942) 357; Payens, FM 1, 5 (1958) 543; Backer & Bakhuizen f., FJ 1 (1963) 441; Burkill,
EPMP (1966) 966; Plowman, Taxon 25 (1976) 141; Ashton, MNDTS 2 (1988) 238; Corner, WSTM 3rd ed. 1 (1988) 254;
Whitmore, Tantra & Sutisna, CLK 1 (1989) 108; Ng, MFFSS 1 (1991) 66; Chung, Sandakania 7 (1996) 67.
Shrubs or trees. Young twigs flattened, usually becoming terete when older; small distichous bracts
(ramenta) often found at the base of lateral twigs and between the leaves. Stipules small, entire, rarely
two-lobed, often 2-keeled below, sometimes emarginate or 2-dentate at apex, caducous and leaving a
distinct scar. Leaves alternate (distichous), involute in bud, pinnately veined; midrib often sunken
above, prominent below, lateral veins numerous, fine, joining and forming marginal loops and
anastomosing irregularly with the intermediate lateral, intercostal, and intramarginal veins to form a
distintive reticulate venation; petioles articulate at the base. Flowers appearing as the leaves mature,
subtended by bracts; pedicels thickened under the calyx, mostly angular, with 2 bracteoles at the base;
calyx lobes imbricate in bud, persistent; petals alternating with the calyx lobes, with a 3-lobed, ligulelike appendage inserted on the apex of the claw; stamens 10, arranged in 2 whorls of 5, persistent,
filaments connate into a persistent staminal tube at base, outermost alternate with the petals, arising
directly from the staminal tube, innermost sometimes ventral to the obtuse or dentate rim of the
staminal tube, anthers ellipsoid, basifixed, cordate at the base; ovary (1)3-locular, only one locule
fertile, with 1 ovule; styles 3, erect, free or partly connate, stigmas capitate (often oblique) or clavate,
rounded or rarely acute. Fruit a drupe, 3-locular, with one containing a seed, the other two empty but
enlarged; mesocarp pulpy; endocarp hard and connate into a 3-locular pyrene. Seeds flattened; with or
without endosperm; testa thin-coriaceous; embryo oblong, straight, erect, cotyledons flat to planoconvex, radicle distinct, pointing towards the fruit apex.
Distribution. A pantropical genus with about 250 species, with the centre of distribution in S
America; 6 species are known in the Malesian region, of which 4 are documented in Sabah and
Sarawak.
Ecology. Primary and secondary forests, from sea level to 500 m, especially on slopes of low hills.
Uses. The family includes the important cocaine-producing plants, Erythroxylum coca (Bolivian coca)
and E. novogranatense (Peruvian coca), whose leaves yield the important alkaloid cocaine, a narcotic
widely used in modern medicine. In the past, E. novogranatense was cultivated in Sri Lanka, Java and
Thailand to produce cocaine, used as a local anaesthetic and in the prescription of some medicines.
The timber of some species is hard, strong, and durable and used locally in tropical America and
Africa for construction purposes. In SE Asia, however, due to its small size, the timber is of minor
importance.
168
ERYTHROXYLACEAE (CHUNG)
Key to Erythroxylum species

1. Shrub or treelet to
long..............................
1.5
tall.
Flowers
solitary;
pedicels
1619
mm
Fig. 2HJ.
E. iwahigense forma calcicola R.C.K. Chung
l.c. 74. Type: Paul Chai S. 39898, Borneo, Sarawak, 4th Div., Mt. Mulu National
Park, Mt. Buda (holotype SAR; isotypes K, KEP, L, MO, SAN).
Leaves thick-coriaceous; elliptic or obovate, (1.6)2.44(4.3) x (1.1)1.31.8(2)
cm; base cuneate, apex acute with rounded or emarginate tip, rarely obtuse with
rounded tip; lateral veins (12)1420 pairs, 12(2.5) mm apart, intercostal veins
distinct, prominent to faint on both surfaces. Flowers solitary, pedicels 1619 mm
long.
Endemic to Sarawak. Very uncommon, known only from the type collection.
Limestone hills to 1000 m. The typical form, E. iwahigense forma iwahigense from
Palawan Is., differs from the present form by its thin-coriaceous leaves and broader
spacing between lateral veins, (3)48(10) mm.
Treelet or tree, 540 m tall. Flowers in clusters of 28, or rarely solitary but never
exclusively so; pedicels 310 mm long...............................................................................2
2. Staminal tube longer than calyx lobes..........................................................1. E. cuneatum
Staminal tube shorter than or at most as long as calyx lobes...............................................3
3. Leaves thin-coriaceous or chartaceous, apex acute with rounded or pointed tip, or
cuspidate, or rarely obtuse with rounded or emarginate tip. Calyx lobes 11.2 mm long.
Fruits broadly ellipsoid, 4.58 mm diameter................................................2. E. latifolium
Leaves thick-coriaceous, apex obtuse-emarginate. Calyx lobes 1.41.8 mm long. Fruits
ellipsoid, to 4 mm diameter...................................................................3. E. sarawakanum
1. Erythroxylum cuneatum (Miq.) Kurz
Fig. 1AG.
(Latin, cuneatus = wedge-shaped; the leaf base)

J. As. Soc. Beng. 43, 2 (1847) 135; O.E. Schulz l.c. 146; Payens l.c. 548; Merrill, Philip. J. Sci. Bot. (1908) 232, EPP (1923)
325; Cockburn l.c. (1976) 84; Anderson l.c.171; Ashton l.c. 238; Corner l.c. 255; Whitmore, Tantra & Sutisna l.c. 108;
Chung l.c. 69. Basionyms: Ficus cuneata Wall., Cat. (1828) no. 4534, nom. nud.; Urostigma ?cuneatum Miq. in Hooker,
Lond. J. Bot. 6 (1847) 585. Type: Wallich, Cat. no. 4534, India Orientalis (holotype K).
Shrub or small to large tree to 40 m tall, 65 cm diameter. Bark grey to reddish brown, smooth to
finely fissured and sometimes scaly, lenticellate, brittle; inner bark orange-brown to reddish brown.
Sapwood pale yellow; heartwood dark reddish brown. Twigs drying brown to black. Stipules
triangular to lanceolate, as long as petiole. Leaves chartaceous to thin-coriaceous, glabrous; elliptic
to elliptic-lanceolate or obovate, 1.912 x 0.95.8 cm; base cuneate, acute to obtuse, apex obtuse to
acute, with emarginate, rounded or rarely pointed tip; midrib sunken above, lateral veins 815 pairs,
213 mm apart, intercostal veins slightly raised to faint on both surfaces; petioles 29 mm long, 0.51
mm thick, glabrous. Flowers in clusters of 26(8), or rarely solitary but never exclusively so;
pedicels 310 mm long, 0.20.7 mm thick, glabrous; calyx lobes 0.51.2 mm high, triangular to
lanceolate; petals white, greenish white to light
169
green, 34 x 1.52 mm, claw distinctly narrowing towards the base, about a third as long as the petal;
filaments in short-styled form c. 23.5 mm long (for equal stamens) and c. 1.54.5 mm long (for
unequal stamens), in long-styled form c. 24 mm long (for equal stamens) and 0.51.5 mm long (for
unequal stamens), staminal tube longer than calyx lobes, 1.52 mm high, anthers 0.30.5 mm across;
ovary ellipsoid or ovoid, longer than staminal tube, 3-locular, styles in short-styled form 12 mm
long, shortly connate at base, in long-styled form 25 mm long, connate to a third to a half of their
length, stigma in short-styled form oblong-ovoid or clavate, in long-styled form capitate, in both
forms broader than the style. Fruits ripening bright red, ellipsoid, 812 x 2.54.5 mm, curved,
glabrous, apex acute, triangular with curved sides in cross section; locules arranged in a triangular
position, with the fertile one nearly of the same size as the 2 sterile locules. Seeds 610 x 1.53 mm;
endosperm present.
Vernacular name. Sabahperepat burong (Malay).
Key to forms
Leaf apex obtuse with rounded or emarginated tip......................................................................
forma cuneatum
Fig. 1AF.
Synonyms: E. burmanicum Griff., Posth. Papers, Not. Pl. As. 4 (1854) 468, t. 581, fig. 3; E.
longistipulatum Burck, Ann. Jard. Bot. Btzg. 11 (1893) 193; E. bancanum Burck l.c. 192, t.
16; E. cuneatum (Miq.) Kurz var. bancanum (Burck) O.E. Schulz. l.c. 148; E. platyphyllum
Merr. l.c. (1908) 232.
Shrub or small tree to 5 m tall, 5 cm diameter. Stipules lanceolate, (2.5)46(7.5) mm long.
Leaves obovate or elliptic to elliptic-lanceolate, (1.9)3.59.5(12) x (0.9)1.94.5(5.8) cm;
base cuneate, apex obtuse with emarginate or rounded tip; lateral veins 812 pairs, (2)39(
13) mm apart; petioles (2)37(9) mm long. Calyx lobes 0.51(1.2) mm high, triangular.
Fruits 810 x 2.54.5 mm. Seeds 68 x 1.53 mm.
Myanmar, Thailand, Indo-China, Sumatra (Banka and Riouw Islands), Peninsular Malaysia,
Singapore, Anambas and Natuna Islands, Java (western part, Kangean), Borneo (Sabah only),
Philippines, Lesser Sunda Islands (Sumbawa) and Maluku. Seasonal monsoon forest, lowland
mixed dipterocarp forest, beaches; also (rarely) on mountain peaks and limestone. In Sabah,
uncommon, found along beaches and on islands. No record from Sarawak, Brunei and
Kalimantan.
Leaf apex acute with rounded or emarginate or rarely pointed tip.............................................
forma sumatranum (Miq.) R.C.K. Chung
Fig. 1G.
l.c. 71. Basionym: E. sumatranum Miq., Fl. Ned. Ind., Suppl. (1862) 200, 512. Type:
Teijsmann, s.n. (= L. sheet no. 908.1252056), Sumatra, Palembang, Ogan Ulu (holotype L;
isotype BO). Synonyms: E. densinerve O.E. Schulz l.c. 142, Merrill l.c. (1921) 313,
Masamune l.c. 357; E. borneense Merr., PEB (1929) 112, Masamune l.c. 357.
Trees to 40 m tall, 65 cm diameter. Stipules triangular, (2)2.54(5) mm long. Leaves
elliptic to narrowly elliptic, lanceolate or rarely obovate, (2.8)4.27.7(10.2) x (0.9)1.7
2.8(3.2) cm; base acute to obtuse, apex acute with rounded or emarginate or rarely acute
tip; lateral veins 815 pairs, (2)37(9) mm apart; petioles (2)35(7) mm long. Calyx
lobes 0.51.2 mm high, triangular to lanceolate. Fruits 912 x 2.54 mm. Seeds 810 x 1.52.5
mm.
Sumatra, Borneo and the Philippines. In Borneo, found in mixed dipterocarp forest to
170
150 m; rather common in Sabah, but uncommon in Sarawak; also known in Kalimantan.
2. Erythroxylum latifolium Burck
Fig. 1H.
(Latin, latus = broad, folium = leaf)

l.c. 192; Merrill l.c. (1921) 313; Masamune l.c. 357; Chung l.c. 77. Type: Teijsmann, s.n., Sumatra, Lingga (holotype BO).
Synonyms: E. latifolium Burck var. angustatum O.E. Schulz l.c. 144; E. latifolium Burck var. longipetiolatum Boerl. &
Koord., Ic. Bog. 1 (1897) t. 6; E. cuneatum (non (Miq.) Kurz) Payens l.c. 549, p.p., quoad syn. E. latifolium, incl. var.
angustatum O.E. Schulz et var. longipetiolatum Boerl. & Koord.
Shrub or small tree to 6 m tall. Twigs brown to grey-brown. Stipules triangular, (2.5)33.5(4) mm
long. Leaves chartaceous or thin-coriaceous; elliptic or rarely obovate, (5.5)6.213(17) x (2.5
)2.85.7(6.4) cm; base cuneate to acute, apex acute with rounded or pointed tip, sometimes cuspidate
or rarely obtuse with rounded or emarginate tip; midrib sunken above, lateral veins 914 pairs, (3
)513(16) mm apart, intercostal veins distinct, slightly prominent below, faint above; petioles (3)4
8(10) mm long, 0.81.5 mm thick, glabrous. Flowers in clusters of 24; pedicels 56 mm long, 0.8
1 mm thick, glabrous; calyx lobes 11.2 mm high, triangular; petals 5; stamens 10, staminal tube
shorter than or at most as long as the calyx lobes. Fruits broadly obovoid, 912 x 4.58 mm, not
curved, glabrous, apex acute, triangular with curved sides in cross-section; locules arranged in a
triangle, with the fertile one nearly of the same size as the 2 sterile locules. Seeds 79 x 35 mm;
endosperm present.
Distribution. Sumatra (including Banka, Billiton, and Lingga), Anambas and Natuna Islands, and
Borneo. In Sarawak, known from Baram, Marudi FR, Lambir Hills NP and Sg. Dalam FR in the 4th
Div. and from the Lassa Protected Forest in the 6th Div. Not yet recorded from Sabah; also found in
Brunei.
Ecology. Lowland mixed dipterocarp, kerangas and peat swamp forests, to 20 m.
3. Erythroxylum sarawakanum R.C.K. Chung
Fig. 2AG.
(of Sarawak)
l.c. 78. Type: Ilias S. 17894, Borneo, Sarawak, Bako National Park (holotype SAR; isotypes A, BO, K, L, SAN, SING).
Shrub or treelet to c. 5 m tall, 4 cm diameter. Twigs grey-brown. Stipules triangular, 22.5 mm long.
Leaves thick-coriaceous, drying brown and glabrous above, pruinose below; obovate or elliptic, (4.3
)5.59(10.5) x (2.2)2.85(5.5) cm; base broadly cuneate to acute, apex obtuse with emarginate tip;
midrib sunken above, lateral veins (6)812 pairs, (4)712(13) mm apart, intercostal veins
distinctly prominent above, faint below; petioles (6)710(12) mm long, 12 mm thick, glabrous.
Flowers in clusters of 24; pedicels 45 mm long, c. 0.5 mm thick, glabrous; calyx lobes 1.41.8 mm
high, triangular; petals 5; stamens 10, staminal tube shorter than or at most as long as the calyx lobes.
Fruits ellipsoid, 9 x 3.54 mm, glabrous, with a rounded top, triangular with curved sides in crosssection; locules arranged in a triangle,
171
Fig. 1. Erythroxylum cuneatum forma cuneatum (AC, F), forma sumatranum (DE, G); E. latifolium (H). A, fruting leafy
twig; B, flower bud; C, long-styled flower with equal stamens; D, short-styled flower with equal stamens, with petals
removed; E, longitudinal section through flower, with petals removed; F, posterior clawed petal with 3-lobed ligule-like
appendage; H, fruiting leafy twig. (A from SAN 107458, B from S. 12753, C and F from KKSS 417, DE and G from SAN
30377, H from S. 25076.)
172
Fig. 2. Erythroxylum sarawakanum (AG); E. iwahigense forma calcicola (HJ). A, fruiting leafy twig; B, detail of lower
leaf surface; C, fruit; D, fruit in cross-section; E, mature flower bud with petals removed; F, flower bud in longitudinal
section; G, posterior clawed petal with 3-lobed ligule-like appendage; H, flowering leafy twig; I, fruit; J, fruit in crosssection. (AD from S. 17894, EG from S. 73603, HJ from S. 39898.)
173
with 2 sterile ones as large as the fertile one, seldom curved. Seeds 68 x2.53 mm; endosperm
present.
Distribution. Endemic to Sarawak. So far known only from a few collections from Bako National
Park (S. 17894, S. 10324, S. 73601), Mt. Matang (Clemens 22354) and Mt. Santubong (S. 37668 and
S. 73603).
Ecology. Lowland to hill forest, to c. 700 m.
Notes. E. sarawakanum differs from E. latifolium in its thick-coriaceous leaves, which are obovate or
elliptic, obtuse at the apex and with a emarginate tip; longer calyx lobes (1.41.8 mm long), and
ellipsoid fruits (up to 4 mm wide).
174
IXONANTHACEAE
Runi S. Pungga
Kuching, Malaysia
Exell & Mendona, Bol. Soc. Brot. 25 (1951) 105; Forman, Kew Bull. 19 (1965) 521; Hutchinson, Gen. Fl. Pl. 2 (1967) 592;
Anderson, CLTS (1980) 219; van Hooren & Nooteboom, Blumea 29 (1984) 550; Kool, FM 1, 10 (1988) 621; Brummitt,
Vasc. Pl. Fam. & Gen. (1992) 589.
Trees or shrubs. Indumentum of simple hairs. Stipules lateral, minute, scale-like, caducous. Leaves
simple, pinnately veined, alternate or spirally arranged. Inflorescences axillary, dichasially
corymbose cymes, lax or fascicled racemes, or panicles of racemes; bracts small, caducous; pedicles
articulate at the base or to the middle. Flower perigynous, bisexual, 5-merous, radially symmetrical;
sepals 5, shortly connate at base, lobes imbricate in bud; petals 5, free, imbricate or contorted in bud,
persistent in fruit; stamens 10 or (15)25, filaments inserted outside and against the disc, irregularly
coiled in bud, anthers basifixed or dorsifixed, 2-locular; ovary 25-locular, ovules 2 or 1 in each
locule; style 1, simple. Fruit a septicidal coriaceous capsule, 5-locular, valves often spuriously
septate, either with or mostly without a central axis, occasionally apically bifid. Seeds 1 or 2 per
locule, with a wing or with a suprahilar arillode; endosperm fleshy or spongy.
Distribution. Three (or possibly five) genera, distributed in the tropical and subtropical regions of the
Old World, Australia and the Pacific Islands. In Sabah and Sarawak, one genus (Ixonanthes).
Taxonomy. The family is closely allied to the Ctenolophonaceae and the Linaceae. It differs from the
Ctenolophonaceae in its indumentum consisting of simple hairs, lateral stipules, spirally arranged
leaves, perigynous flowers, persistent petals in fruits, intrastaminal disc, simple style, and seeds that
have a prominent basal wing or suprahilar arillode. From the Linaceae, it differs in its perigynous
flowers, persistent petals in fruits, free filaments inserted outside the disc, single style, capsular fruit,
and seeds with a basal wing or suprahilar arillode. Forman (l.c.) described Allantospermum from
Borneo and assigned it to the Ixonanthaceae. Nooteboom (Adansonia 2, 7 (1967) 161; FM 1, 6 (1972)
970) and Kool (l.c.), however, excluded the genus from the Ixonanthaceae and included it in the
Simaroubaceae, and recognised three genera, viz. Cyrillopsis (from S America), Ochthocosmus (from
tropical America and Africa) and Ixonanthes (from Indo-Malesia), in the family. Their decision is
followed in the present account.
175
IXONANTHES Jack
(Greek, ixos = birdlime, anthos = flower; the sticky flowers)
Mal. Misc. 2, 7 (1822) 51; Merrill, EB (1921) 313; Ridley, FMP 1 (1922) 324; Hallier, Beih. Bot. Centralbl. 39 (1923) 6;
Masamune, EPB (1942) 356; Burgess, TBS (1966) 247; Cockburn, TFM 1 (1972) 307, TS 1 (1972) 205; Anderson, CLST
(1980) 219; Kool, Blumea 26 (1980) 191, FM 1, 10 (1988) 622; Corner, WSTM 3rd ed. 1 (1988) 255; Whitmore, Tantra &
Sutisna, CLK 2, 1 (1990).
Trees or treelets growing monopodially in flushes, often with steep and sharp buttresses; branches
ascending. Bark grey-brown, smooth or finely fissured and lenticellate; inner bark dark reddish
brown, fibrous. Sapwood cream to pale yellow; heartwood dark purplish brown. Leaves spirally
arranged, thin- to thick-coriaceous; base cuneate, margin entire or remotely serrate, apex obtuse or
often emarginate, drying dark-brown; midrib sunken on the upper surface, raised on the lower surface;
lateral veins faint or distinctly raised on both surfaces; intercostal veins finely and irregularly
reticulate; petioles swollen at the base, grooved on the adaxial side, drying blackish. Flowers in longstalked, dichasially corymbose cymes; young buds often sticky or resinous; stamens 10 or (15)20 in
one whorl, subpersistent in fruit; disc annular or cup-shaped, intrastaminal; ovary 5-locular; ovules 2
in each locule, placentation axile; stigmas discoid. Fruits ellipsoid, apiculate, thinly woody, smooth.
Seeds with a firm testa; endosperm spongy, oily; embryo straight.
Vernacular names. Sarawakinggir burung (Malay, Iban), kayo mupa (Berawan), kayu mup (Penan
Tutoh), meridian (Bidayuh), pagar anak (Malay).
Distribution. Three species in continental SE Asia and Malesia (except Java and Lesser Sunda
Islands); 2 species in Borneo.
Ecology. In primary and secondary forests on hillsides, ridges and slopes, also in kerangas forest,
from sea level to 800 m.
Taxonomy. The genus has been divided (Hallier l.c., Kool l.c.) into two sections, viz. section
Brewstera represented by I. icosandra which does not occur in Borneo, and section Ixonanthes
represented by I. petiolaris and I. reticulata. Section Brewstera differs from section Ixonanthes in its
shorter petioles (23 mm), slightly glandular, serrate leaf margins, sub-4-whorled branching of the
peduncles, each flower with (15)20 stamens, and seeds with suprahilar arillode.
Key to Ixonanthes species

Leaves thin-coriaceous; lateral and intercostal veins faint on the lower leaf surface.
Inflorescences dense. Styles 1.52.5 mm long. Fruits 1.32 cm long; valves ultimately
apically bifid. Seeds 1013 x 34 mm...................................................................1. I. petiolaris
Leaves thick-coriaceous; lateral and intercostal veins prominent on the lower leaf surface.
Inflorescences lax. Styles 1520 mm long. Fruits 24.5 cm long; valves never apically bifid.
Seeds 1820 x 49 mm.........................................................................................2. I. reticulata
176
IXONANTHACEAE (RUNI PUNGGA)
1. Ixonanthes petiolaris Blume
Fig. 1.
(Latin, petiolaris = with a well-developed stalk; the leaves)

Mus. Bot. Lugd. Bat. 1 (1852) 396; Cockburn l.c. (1976) 207; Kool l.c. (1980) 199, l.c. (1988) 626; Whitmore, Tantra &
Sutisna l.c. 209. Type: Praetorius, s.n. (= L. sheet no. 908. 1261336) (L). Synonyms: I. multiflora Stapf ex Ridl., Kew
Bull. (1930) 75, Masamune l.c. 356, Anderson l.c. 220; I. philippinensis Elmer, Leafl. Philip. Bot. 10 (1930) 3758.
Tree to 45 m tall, 50 cm diameter; bole deeply fluted. Bark smooth, with numerous lenticels, pale
brown to yellowish brown or black, flaky and minutely ridged. Leaves thin-coriaceous, elliptic to
obovate, 615 x 37.5 cm; base acute, apex slightly obtuse; lateral and intercostal veins rather faint
on lower leaf surface; petioles 1420 mm long. Inflorescences dense; peduncles 3.57 cm long;
pedicels c. 5 mm long. Flowers at anthesis 1.53 x 12.5 mm; sepals elliptic to orbicular, 11.5 x 1
1.5 mm (in fruit fleshy and enlarging to 1.52 mm), thickened at base, margins with a c. 1.5-mm-wide
transparent band, apex rounded; petals elliptic-orbicular, 22.5 x 11.5 mm (in fruit chartaceous and
enlarging to 2.53 x 1.52 mm), thickened at base, margins with a narrow transparent band, apex
rounded; stamens 10, filaments to 15 mm long; ovary depressed globose, style 1.52.5 mm long.
Fruits ellipsoid, 1.3(2) x 0.8(1.2) cm; valves apically bifid. Seeds 1013 x 34 mm, with basal wing.
Vernacular name. Sabahpagar anak (Malay).
Distribution. Thailand, Sumatra, Peninsular Malaysia, Borneo, Philippines and Sulawesi. In Sabah
and Sarawak, fairly common; also in Kalimantan.
Ecology. In primary and secondary forests on slopes and ridges; from sea level to 800 m.
Uses. The timber is used in minor construction.
2. Ixonanthes reticulata Jack
Fig. 2.
(Latin, reticulatus = net-like; the leaf intercostal veins)

l.c. 51; Ridley l.c. (1922) 326; Hallier l.c. 9; Masamune l.c. 357; Browne, FTSB (1955) 175; Cockburn l.c. (1972) 307, l.c.
(1976) 206; Kool l.c. (1980) 200, l.c. (1988) 626; Corner l.c. 256; Whitmore, Tantra & Sutisna l.c. 209. Type: Jack, s.n.,
Sumatra, Tapanuli (L). Synonyms: I. grandiflora Hochr., Pl. Bog. Exsicc. (1904) 39, Merrill l.c. 313, Masamune l.c. 356; I.
beccarii Hallier l.c. 10, Anderson l.c. 220; I. crassifolia Hallier l.c. 10, Cockburn l.c. (1976) 206; I. grandifolia Ridl. l.c.
(1930) 74, Masamune l.c. 356.
Tree to 35 m tall, 30 cm diameter; bole straight and fluted; buttresses narrow, small. Bark smooth,
slightly fissured, scaly or cracked, pink, yellowish brown, fawn or grey to blackish. Wood white,
hard. Leaves thick-coriaceous, elliptic to obovate, 5.514 x 310 cm; base cuneate, apex obtuse or
emarginate; lateral and intercostal veins raised and prominent on both surfaces, intercostal veins
distinctly reticulate; petioles 1020 mm long. Inflorescences lax; peduncles 2.514 cm long; pedicels
0.51.5 cm long; sepals elliptic to linear-elliptic, 48 x 34 mm; petals elliptic-orbicular, 410 x 36
mm; stamens 10, filaments to 20 mm long; ovary globose, style 2025 mm long. Fruits ellipsoid, 2
4.5 cm long, valves not apically bifid, septa after fruit-dehiscence persistently connate with the
adjacent one. Seeds 1820 x 49 mm, with basal wing.
177
Fig. 1. Ixonanthes petiolaris. A, fruiting leafy twig; B, fruit; C, open fruit; D, fruit valve; E, flower; F, longitudinal section of
flower; H and I, stamens. (AB from SAN 50411, CD from SAN 44751, EI from SAN 40370.)
178
IXONANTHACEAE (RUNI PUNGGA)
Fig. 2. Ixonanthes reticulata. A, fruiting leafy twig; B, fruit; C, open fruit, D, flower bud; E, flower bud in longitudinal
section; FG, stamens. (A from S. 34367, B from SAN 73585, C from SAN 39762, DG from S. 13482.)
179
Vernacular name. Sabah and Sarawakinggir burung (Malay, Iban).

Distribution. India, Myanmar, Vietnam, S China, Hong Kong, Sumatra, Peninsular Malaysia,
Borneo, Philippines, Sulawesi, and New Guinea. Common in Sabah and Sarawak; also in Brunei and
Kalimantan.
Ecology. In primary and secondary forests on hillsides and ridges; also in kerangas and swamp
forests, from sea level to 2400 m.
Uses. The timber is utilised for minor construction and firewood.
180
LEEACEAE
A. Latiff
Universiti Kebangsaan Malaysia,
Bangi, Malaysia
Dumort., Anal. Fam. Pl. (1829) 27; Ridsdale, Blumea 22 (1974) 57, FM 1, 7 (1976) 755.
A monogeneric family closely allied to the Vitaceae in the order Rhamnales, separated because of the
development of a complex staminodial tube joined to the corolla, the ovary locules that each have a single
ovule, and pollen morphology (Ridsdale l.c. 1974).
LEEA van Royen ex L., nom. cons.

(James Lee, 17151795, a Scottish gardener)
mali-mali (Malay)
Syst. Nat. ed. 12, 2 (1767) 627; Merrill, EB (1921) 368; Ridley, FMP 1 (1922) 483; Masamune, EPB (1942) 439;
Suessenguth in Engler & Prantl, Nat. Pfl. Fam. 2, 20d (1953) 382; Backer & Bakhuizen f., FJ 2 (1965) 93; Ridsdale l.c.
(1974) 74, l.c. (1976) 755; Anderson, CLTS (1980) 134; Corner, WSTM 3rd ed. 2 (1988) 757; Latiff, TFM 4 (1989) 480.
Trees, erect or creeping shrubs, scramblers, or herbaceous plants with woody base; stems with spines
or unarmed. Leaves compound, 13-foliolate or 14-pinnate, usually imperfectly pinnate with
terminal leaflets; petioles expanded to form a stipular structure surrounding the stem apex; stipules
narrowly sheathing and subpersistent or large and caducous; leaflets glabrous or pubescent, lower
surface often with pearl-glands, margin serrate or crenate. Inflorescence a cyme, lax or condensed,
axillary or seemingly terminal, distinctly stalked. Flowers bisexual, radially symmetrical, 45merous; calyx bell-shaped, lobes triangular; corolla lobes valvate in bud, apical part of each lobe
joined into a keel, reflexed at maturity, basal part joined to each other and to the androecium;
staminodial tube consisting of thickened parts representing the filaments and thin parts or sinuses
between the filaments, the tube joined to the corolla above the base, but free above and below this
point, anthers introrse; ovary discoid, 410-locular, each locule with one ovule, style short, entire,
stigma slightly thickened. Fruit a berry, depressed subglobose. Seeds triangular-ovoid in crosssection; endosperm simply ruminate; embryo linear.
Distribution. About 32 species; from Africa and Madagascar to SE Asia, Malesia, Micronesia,
Melanesia and Australia. In Sabah and Sarawak, represented by 7 species, of which only 5 reach treesize.
181
Ecology. Quite common in a variety of habitats from beach vegetation to montane forest, but mostly
found in lowland secondary vegetation. The flowers are pollinated by flies, while the fruits are
dispersed by birds.
Uses. Members of the genus do not reach timber size. Locally, some species are used in traditional
medicine. Some Dusun groups in Sabah utilise leaves of Leea for scaring off evil spirits from entering
the house.
Key to Leea species

1. Stems and main branches spiny.........................................................................................................2
Stems and main branches not spiny...................................................................................................3
2. Plant single-stemmed. Leaves 1-pinnate. Inflorescences glabrous. Fruits 1015 mm in
diameter.........................................................................................................................1. L. aculeata
Plant many-stemmed. Leaves 2 or 34-pinnate. Inflorescences pubescent. Fruits 710 mm in
diameter........................................................................................................................3. L. angulata
3. Leaves 1-pinnate...................................................................................................................
Leea amabilis Veitch ex Masters
Gard. Chron. 27 (1882) 492; Masamune l.c. 439; Suessenguth l.c. 385; Ridsdale l.c. (1976)
765.
Treelet to 2 m tall. Flowers 5-merous, white. Fruits deeply grooved between segments, 1820
mm in diameter.
Uncommon and endemic to Sarawak.
Leaves 2- to 4-pinnate......................................................................................................................4
4. Stipules resembling narrow wings, subpersistent, scars long and thin. Leaflets without pearlglands. Flowers red......................................................................................................
Leea rubra Blume ex Spreng.
Syst. Veg. 1 (1824) 670; Merrill l.c. (1921) 396; Ridley l.c. 485; Burkill, EPMP (1935) 1327;
Masamune l.c. 439; Suessenguth l.c. 383; Ridsdale l.c. (1976) 776; Latiff l.c. 482; Corner l.c.
757.
Shrub to 3 m tall. Leaves 24-pinnate. Inflorescences rusty pubescent. Flowers 5-merous,
bright red. Fruits dark red, c. 8 mm diameter.
India, Bangladesh, Myanmar, Thailand, Cambodia, Laos, Vietnam, throughout Malesia and
Australia. In Borneo, known only from Sabah.
Stipules obovate, caducous, leaving broad-triangular scars. Leaflets with pearl-glands. Flowers
reddish orange or greenish white.......................................................................................................5
5. Young twigs and stipules densely pubescent. Inflorescence and leaflets with large, conspicuous,
light brown pearl-glands...............................................................................................2. L. aequata
Young twigs and stipules glabrous or only sparsely pubescent. Inflorescences and leaflets with
small, inconspicuous, caducous pearl-glands....................................................................................6
6. Leaves 2-pinnate; leaflet margin wavy to finely serrate. Flowers reddish orange.................
...................................................................................................................................4. L. guineensis
182
LEEACEAE (LATIFF)
Leaves 3-pinnate; leaflet margin crenate or coarsely serrate to shallowly dentate. Flowers greenish
white.................................................................................................................................5. L. indica
1. Leea aculeata Blume ex Spreng.
Fig. 1.
(Latin, aculeatus = prickly; the stems and branches)

l.c. 670; Merrill l.c. (1921) 368, PEB (1929) 183; Masamune l.c. 439; Suessenguth l.c. 383; Backer & Bakhuizen f. l.c. 93;
Ridsdale l.c. (1976) 773; Anderson l.c. 134. Type: Blume, s.n., Java, Mt. Salak (L). Synonym: L. sandakanensis Ridl., Kew
Bull. (1931) 499, Masamune l.c. 439, Suessenguth l.c. 386.
Single-stemmed shrub or small tree to 10 m tall; trunk and main branches with spines. Stipules
forming very narrow wings, c. 2 x 0.5 cm, scars of similar length. Leaves 1-pinnate; rachis 815 cm
long; petioles 26 cm long. Leaflets 59, glabrous, thin-coriaceous, pearl-glands globose, black;
elliptic to elliptic-oblong, occasionally ovate to ovate-oblong, 1018 x 47 cm; base rounded to
cuneate, margin infrequently serrulate, apex long-acuminate; lateral veins 612 pairs; petiolules to 2
cm long. Inflorescences 816 cm long, broad and many-branched, glabrous; bracts triangular, to c.
3 x 2 mm; peduncles 28 cm long. Flowers 5-merous, greenish white; calyx enclosing the corolla, c.
3 x 3 mm, glabrous, lobes c. 2 x 1 mm; corolla tube and staminodial lobes c. 4 mm long; corolla lobes
c. 3 x 2 mm; staminodial tube c. 3.5 mm long, upper free part c. 1.5 mm long, lobes slightly cleft,
sinuses shallow, lower free part 1.52 mm long, extending downwards to the ovary; free part of
filaments c. 1 mm long, anthers c. 1 mm across; ovary 46-locular, style c. 2 mm long. Fruits 1015
mm diameter, shallowly grooved, blue-black. Seeds usually 6, c. 8 x 4 mm.
Vernacular names. Sabahtogunumali (Dusun Kinabatangan). Sarawakkamali, kemali (Iban),
mali-mali (Malay), teloyang (Berawan).
Distribution. Sumatra, W Java, Borneo, Philippines, Sulawesi, Maluku and Irian Jaya. Common
throughout Sabah, less common in Sarawak.
Ecology. Widespread in secondary vegetation, particularly in riverine areas.
Notes. In Sabah and Sarawak, there are two species with spines. Unlike L. angulata, the spines in this
species are found only on the trunk and main branches.
2. Leea aequata L.
(Latin, aequatus = watery; the soft fruits)
Syst. Nat. ed. 12, 2 (1767) 627; King, J. As. Soc. Beng. 65, 2 (1896) 419; Winkler, Bot. Jahrb. 44 (1910)
537; Merrill l.c. (1921) 368; Ridley l.c. (1922) 486; Masamune l.c. 429; Corner l.c. 757; Suessenguth l.c. 388; Backer &
Bakhuizen f. l.c. 94; Ridsdale l.c. (1976) 775; Anderson l.c. 134; Latiff l.c. 481. Type: Herb. Linn. 1118: 1 (LINN).
Shrub, treelet or less frequently small tree to 8 m; stems and main branches not spiny; young twigs
usually densely pubescent. Stipules obovate, c. 4 x 3 cm, sparsely to densely pubescent, caducous,
scars broad-triangular, 23 cm long. Leaves 23-pinnate; rachis 720 cm long, petioles 814 cm
long, usually pubescent. Leaflets 5 to numerous, chartaceous, glabrous to
183
Fig. 1. Leea aculeata. A, flowering leafy twig; B, cut flower bud showing the staminodial tube; C, longitudinal section of
flower; D, young fruit; E, infructescence; F, fruit. (AC from S. 45409, DF from SAN 113617.)
184
LEEACEAE (LATIFF)
pubescent on the veins above, sparsely to densely pubescent beneath, pearl-glands rounded to discshaped, light brown, large and conspicuous; ovate to ovate-lanceolate or elliptic to elliptic-lanceolate,
1020 x 46 cm; base cuneate to truncate, sometimes subcordate or asymmetric, margins serrate, apex
acuminate to long-acuminate; lateral veins 812 pairs, usually densely pubescent; petiolules 515 mm
long, hairy. Inflorescences 615 cm long, rusty pubescent, covered with large, conspicuous, light
brown pearl-glands; bracts ovate, c. 6 x 5 mm, conspicuous; peduncles 24 cm long, lateral and
ultimate branches rather short, sometimes condensed. Flowers 5-merous, greenish white; calyx c.
4 x 4 mm, glabrous to densely pubescent, usually with pearl-glands, lobes c. 1 x 2 mm; corolla tube and
staminodial lobes c. 2 mm long; corolla lobes c. 2 x 1 mm; staminodial tube 1.52 mm long, upper free
part c. 1.5 mm long, lobes deeply emarginate, sinuses shallow, to 0.5 mm deep, lower free part 0.2
0.4 mm long; free part of filaments c. 1 mm long, anthers c. 1 mm across; ovary 47-locular, style 1,
c. 0.5 mm long. Fruits 912 mm diameter, red. Seeds 6, 46 mm across.
Distribution. India, Bhutan, Nepal, Bangladesh, Andaman Is., Myanmar, Thailand, Cambodia, Laos,
Vietnam, throughout Malesia except New Guinea; uncommon in Sabah and Sarawak.
Ecology. In secondary vegetation.
3. Leea angulata Korth. ex Miq.

(Latin, angulatus = angular; the branches and twigs)
Ann. Mus. Bot. Lugd. Bat. 1 (1863) 97; King l.c. 414; Ridley l.c. (1922) 485; Corner l.c. 757; Backer & Bakhuizen f. l.c. 94;
Ridsdale l.c. (1976) 772; Latiff l.c. 481. Type: Korthals, s.n., Java, Kerawang (L).
Weak straggler, bushy shrub, or tree to 10 m tall, frequently many-stemmed and suckering; trunk,
main and ultimate branches with whitish triangular spines. Bark smooth, inner bark pale. Stipules
forming narrow wings, c. 2 x 2 cm, usually extending the whole length of the petioles, scars narrow.
Leaves 2- or 34-pinnate; rachis 1418 cm long; petioles c. 46 cm long. Leaflets numerous,
glabrous, pearl-glands rounded, caducous; elliptic to elliptic-lanceolate or ovate to ovate-lanceolate, c.
10 x 4 cm; base rounded to cuneate, margin crenate, less frequently shallowly serrate, apex acuminate;
lateral veins 48 pairs, often with pubescent domatia, rarely sparsely pubescent along the whole
length; petiolules to 8 mm long. Inflorescences to 20 cm long, broad, many-branched, pubescent;
bracts triangular to narrowly triangular, to c. 3 x 2 mm; peduncles 47 cm long, main branches long,
ultimate branches lax. Flowers 5-merous, greenish white; calyx c. 2 x 2 mm, pubescent, lobes c. 1 mm
long; corolla tube and staminodial lobes c. 4 mm long; corolla lobes c. 2 x 1 mm; staminodial tube c. 2
mm long, upper free part 1.21.5 mm long, lobes obtuse, sinuses shallow, lower free part c. 0.5 mm
long; free part of filaments c 1.5 mm long, anthers c. 1.5 mm across; ovary 6-locular, style c. 2 mm
long. Fruits 710 mm diameter, greyish blue. Seeds usually 6, c. 5 x 3 mm.
Vernacular name. Sabahmali-mali (Malay).
Distribution. Nicobar Is., Thailand and Malesia. In Sabah, it has been recorded from the Tawau,
Semporna and Sandakan districts.
185
Ecology. Uncommon in secondary vegetation in the east coast of Sabah, especially in logged-over
forest.
4. Leea guineensis G. Don

(of Guinea)
Gen. Hist. 1 (1831) 712; Ridsdale l.c. (1974) 92, l.c. (1976) 777; Latiff l.c. 481. Type: G. Don, s.n. (BM). Synonym: L.
javanica (non Blume) Miq., Ann. Mus. Bot. Lugd. Bat. 1 (1863) 100, Merrill l.c. (1921) 369, Masamune l.c. 439; L. robusta
Ridl. l.c. (1922) 486.
Shrub with soft branches, to 5 m tall; stems and main branches not spiny. Young twigs glabrous,
rarely sparsely pubescent. Stipules obovate, c. 12 mm broad, caducous, glabrous, scars broadtriangular. Leaves 2-pinnate; rachis to 50 cm long; petioles 1014 cm long. Leaflets numerous,
glabrous above, scabrous beneath, pearl-glands small, inconspicuous; elliptic-lanceolate to broadly
elliptic, 8.319 x 4.510.8 cm; base obtuse, margin wavy to finely serrate, apex caudate; lateral veins
1016 pairs. Inflorescences 3-forked cymes, c. 15 cm long, very dense, rusty pubescent, glabrescent,
covered with small, inconspicuous, caducous pearl-glands; peduncles c. 3.5 cm long. Flowers 5merous, reddish orange; calyx c. 3 x 2 mm, glabrous, lobes c. 1 x1 mm; corolla tube and staminodial
lobes 35 mm long; corolla lobes c. 3 x 2 mm; staminodial tube 23 mm long, upper free part 12.5
mm long, lobes shallowly retuse, notched or cleft, sinuses shallow; free part of filaments c. 1 mm
long, anthers c. 1.5 mm across; ovary (4)6(8)-locular, style 12.5 mm long. Fruits 6-seeded, c. 10
mm diameter. Seeds c. 6 x5 mm.
Distribution. Tropical Africa, India, Andaman Is., Myanmar, Thailand, Cambodia, Laos, Taiwan,
Peninsular Malaysia, Sumatra, Java, Lesser Sunda Isl., Borneo, Sulawesi, New Guinea and
Micronesia. In Borneo, known from Kalimantan and Sabah. In Sabah, it is confined to the Kota
Marudu area.
Ecology. In secondary vegetation.
5. Leea indica (Burm. f.) Merr.

(of India)
Philip. J. Sci. 14 (1919) 245, l.c. (1921) 368; Craib, Fl. Siam En. 1 (1926) 318; Masamune l.c. 439; Corner l.c. 758; Merrill
& Perry, J. Arn. Arb. 22 (1941) 380; Ridsdale l.c. (1976) 779; Anderson l.c. 134; Latiff l.c. 482. Basionym: Staphylea
indica Burm. f., Fl. Ind. (1786) 75, t. 23, f. 2. Type: Burmann f. (1986) t. 23, fig. 2, description. Synonym: L. sundaica Miq.,
Fl. Ind. Bat. 1, 2 (1859) 610, Merrill l.c. (1921) 369, Ridley l.c. (1922) 485, Masamune l.c. 439, Backer & Bakhuizen f. l.c.
94
Shrub or small tree, to 10 m tall, many- or single-stemmed, frequently stilt-rooted; stems and main
branches not spiny; young twigs glabrous, rarely woolly or rough-hairy or pustulate or papillose.
Stipules obovate, to 6 x 4 cm, caducous, usually glabrous or sparsely pubescent, rarely densely soft or
bristly pubescent, scars broadly triangular, c. 4 cm long. Leaves 3-pinnate; rachis 1530 cm long,
glabrous to pubescent, rarely soft or bristly pubescent, or papillose; petioles 820 cm long. Leaflets
glabrous to pubescent, rarely densely so or woolly, chartaceous to thin-coriaceous, pearl-glands
small, inconspicuous, caducous, angular to
186
LEEACEAE (LATIFF)
subglobose; broadly ovate, ovate-oblong to ovate-lanceolate, 8-30 x 3-16;base cuneate, rounded to

subcordate, margin crenate, serrate to shallowly dentate, apex acute to acuminate; lateral veins 816
pairs; petiolules to 25 mm long, glabrous to pubescent. Inflorescences 1040 cm long, usually broad
and lax, rarely condensed, glabrous to pubescent, rarely soft or bristly pubescent or papillose, pearlglands small, inconspicuous, caducous; bracts triangular to linear, to 8 mm long; peduncles to 15 cm
long, lateral and ultimate branches numerous and spreading, rarely condensed. Flowers 5-merous,
greenish white; calyx c.3 x 2 mm, glabrous to pubescent, lobes c.2 x 1 mm; corolla tube and
staminodial lobes c. 3 mm long; corolla lobes c. 3 x 2 mm, staminodial tube 1.52.5 mm long, upper
free part 12 mm long, lobes shallowly retuse, notched or cleft, sinuses shallow to 0.4 mm deep,
rarely to 1 mm deep, lower free part c. 0.5 mm long; free part of filaments c 1.5 mm long, anthers c. 1
mm across; ovary 48-locular, style c. 2 mm long. Fruits 610 mm across, purple-black. Seeds 6, c.
5 x 4 mm.
Vernacular names. Sabahmali-mali (Malay), pantan pantan (Dusun Tuaran), togimamalid (Dusun
Kiau). Sarawakkayu kenupan (Penan), kemali (Iban).
Distribution. India, Sri Lanka, Nepal, Bangladesh, Andaman and Nicobar Is., Myanmar, Thailand,
Cambodia, Laos, Vietnam, Hainan, China, throughout Malesia, Australia to Pacific islands.
Widespread in Sabah and Sarawak; also in Brunei.
Ecology. Common in primary and secondary forests, from sea level to mountains, up to 1700 m.
187
188
LOGANIACEAE
K.M. Wong & John B. Sugau
Sandakan, Malaysia
Merrill, EB (1921) 490; Masamune, EPB (1942) 613; Leenhouts, Bull. Jard. Bot. Brux. 32 (1962) 417, FM 1, 6 (1962) 293,
Blumea 14 (1966) 230, Blumea 29 (1984) 423; Burgess, TBS (1966) 385; Cockburn, TS 1 (1976) 207; Anderson, CLTS
(1980) 239; Ashton, MNDTS 2 (1988) 307.
Trees, shrubs, woody climbers, stranglers, epiphytes or herbs. Leaves opposite, simple, entire;
stipules absent but petiole bases expanded on the margins and joining between leaves to form an
ochrea around the node (Mitrasacme, Mitreola and Norrisia), or developing transverse scale-like
appendages just above the leaf axil (axillary scales) that resemble truncate rims which are separate
between leaves (Fagraea) or joined between leaves to form a cup-shaped ochrea tightly encircling the
node (Fagraea, Geniostoma), or these appendages absent and the nodes with a transverse line or
thickening between petiole bases (Gelsemium, Norrisia, Strychnos). Inflorescence terminal or
axillary, a cyme, raceme, thyrse or a solitary flower. Flower nearly always bisexual, radially
symmetrical, (4)5-merous; sepals united or free, valvate or imbricate; petals fused and forming a
tube, lobes (4)5, valvate, imbricate or contorted in bud; stamens inserted on the corolla tube,
alternate with corolla lobes, anthers basifixed or rarely dorsifixed, dehiscing lengthwise; disc
sometimes present, ovary superior, usually 2-locular, rarely 1- or 4-locular, ovule 1many per locule,
amphitropous or anatropous, placentation axile; style usually one, stigma knob-like or club-shaped or
faintly or deeply bifid. Fruit a berry, capsule or drupe. Seeds 1many, with copious endosperm;
embryo minute, straight.
Distribution. About 28 genera with some 600 species have been placed in this family, taken in its
wider taxonomic sense. Chiefly pantropical, a few genera extending to warm-temperate regions,
mainly towards the south. In Malesia, 10 genera with c. 100 species; 7 genera in Sabah and Sarawak
of which Fagraea is the main genus.
Uses. Strychnine is obtained from Strychnos and other poisonous alkaloids are also known from
Mostuea and Gelsemium spp. Only a few species of Fagraea are taken for timber. Fagraea and
Gelsemium yield various species of ornamental importance or potential.
Taxonomy. Buddleja, commonly included in the Loganiaceae in the past is now generally accepted as
representing a separate family, Buddlejaceae (in the Scrophulariales), chiefly on account of the
occurrence of intraxylary phloem in its wood, pollen characters that resemble those of the
Scrophulariaceae and a number of characters (e.g., possession of stellate hairs and glandular-capitate
hairs) that are absent in most of the Loganiaceae.
189
Recent work appears to indicate that the Loganiaceae as here circumscribed, mainly following the
concept of Solereder (in Engler & Prantl, Nat. Pfl. Fam. 4, 2 (189295) 19), is still a heterogenous
assemblage of distinct families belonging to the Gentianales. It has been suggested that the
Loganiaceae sensu stricto should comprise just Logania, Mitrasacme and Mitreola; Gelsemium and
Geniostoma should be the bases of distinct families; Strychnos and Norrisia might be placed together
within the Strychnaceae; and Fagraea belongs properly to the Gentianaceae (Struwe et al., Cladistics
10 (1994) 175).
The concept for grouping the genera used here is therefore only one of convenience, as we await more
detailed studies that corroborate the newly proposed classification. Of importance in our flora are the
genera Fagraea, Geniostoma and Norrisia.
Key to genera
1. Small herbs.....................................................................................................................................2
Shrubs, trees, climbers, stranglers or epiphytes................................................................................3
2. Flowers 4-merous. Leaves small (less than 2 cm long), petioles inconspicuous...................
Mitrasacme Labill.
Nov. Holl. Pl. Spec. 1 (1804) 35; Merrill, EB (1921) 490; Masamune, EPB (1942) 615;
Leenhouts, Bull. Jard. Bot. Brux. 32 (1962) 440, FM 1, 6 (1962) 378.
About 40 species, mainly Australian, also in Tasmania, New Zealand, New Caledonia,
Malesia, the Carolines, and in Asia from the Deccan and Sri Lanka to Central Japan and
Korea.
Wiry, low, usually clump-forming herbs. Leaves often with only the midrib distinct.
In Sabah, one species (M. neglecta) recorded from Balambangan Is., and in Sarawak, one
species (M. pygmaea) recorded from Mt. Santubong.
Flowers 5-merous. Leaves larger (at least several cm long), with distinct petioles...............
Mitreola L.
Gen. Pl. ed. 1 (1737) 377. Synonym: Cynoctonum Gmel., Syst. (1791) 443; Leenhouts, FM 1,
6 (1962) 375.
Seven species; Madagascar, SE Asia, Malesia, and N and W Australia. In Sabah and E
Kalimantan, one species.
Low herb, less than 0.5 m high. Inflorescence axillary, dichasial with long cincinnate
branches.
3. Leaves with 35 main veins from the leaf base. Tendrils often present in leaf axils..............
Strychnos L.
Sp. Pl. 1 (1753) 189; Gen. Pl. ed. 5 (1754) 86; A. W. Hill, Kew Bull. (1917) 121; Merrill, EB
(1921) 490; Masamune, EPB (1942) 616; Leenhouts, FM 1, 6 (1962) 374.
About 150200 species, in tropics and subtropics. In Sabah and Sarawak, c. 11 species.
Woody climbers or rarely shrubs or treelets; usually with axillary, simple or double tendrils.
Petioles often inserted on swellings (leaf-cushions) at the nodes.
Leaves pinnately nerved. Tendrils absent.........................................................................................4
190
LOGANIACEAE (WONG & SUGAU)
4. Petiole bases developing conspicuous wings or axillary scales or forming a distinct cup-like
ochrea around the node......................................................................................................................5
Petiole bases not forming any special structures, the nodes at most marked by a transverse line or
thickening, or the ochrea very low and not more than 1 mm high, inconspicuous.............................................................................................................................................6
5. Fruit a dehiscent capsule. Inflorescence strictly axillary. Corolla campanulate to rotate Leaves
chartaceous to thin-coriaceous, with strongly recurved margin when dry.................
....................................................................................................................................2. Geniostoma
Fruit a berry, not dehiscent. Inflorescence terminal or axillary. Corolla salverform, infundibular
(funnel-shaped) or tubular. Leaves thin- to thick-coriaceous, the margins recurved only in some
thick-coriaceous leaves......................................................................................................1. Fagraea
6. Leaf apex rounded to acute or slightly acuminate. Style knob-like or slightly 2lobed.................................................................................................................................3. Norrisia
Leaf apex long-caudate. Style twice forked...........................................................................
Gelsemium Juss.
Gen. Pl. (1789) 150; Merrill, EB (1921) 490; Masamune, EPB (1942) 615; Leenhouts, FM 1,
6 (1962) 343.
Three species; SE Asia, SE North America, Mexico and Guatemala. In Sabah and Sarawak,
one species (G. elegans).
Liana or straggling shrub. Leaves pinnately nerved. Flower 5-merous; corolla lobes imbricate.
Fruit a dry capsule, 2-valved. Seeds winged.
1. FAGRAEA Thunb.
(J.T. Fagraeus, 172947, Swedish naturalist)
Merrill, EB (1921) 491; Masamune, EPB (1942) 613; Leenhouts, Bull. Jard. Bot. Brux. 32 (1962) 417, FM 1, 6 (1962) 293,
Blumea 14 (1966) 230, 29 (1984) 423; Burgess l.c. 385; Cockburn l.c. 207; Anderson l.c. 239; Ashton l.c. 307; Wong &
Sugau, Sandakania 8 (1996) 1.
Trees, shrubs, woody climbers, stranglers or epiphytes; buttresses absent or very small. Leaves strapshaped, elliptic, lanceolate, oblanceolate, obovate to broadly ovate, pinnately nerved, thin- to thickcoriaceous, margin mostly entire or crenulate (F. crenulata), mostly glabrous; petioles indistinct to
distinct, developing axillary scales above the base that loosely clasp the node (section Fagraea) or at
the very base and tightly clasping the node (sections Cyrtophyllum and Racemosae), the petiole base
with or without auricles developed. Inflorescences terminal or rarely axillary, either without any
branches (with only a solitary flower) or cyme-like, with well-developed primary branches that
rebranch 16 times (sections Fagraea and Cyrtophyllum), or branches typically very condensed and
grouped as distinct tiers along the main axis (section Racemosae); bracts small and scale-like,
bracteoles subtending flowers 23 pairs, small or in some species very large and forming an involucre
around the flower. Flowers small to very large, 5-merous; calyx with a firm cup-like basal part, lobes
imbricate; corolla mostly creamy-white, either tubular, salverform or narrowly to widely
infundibular, lobes contorted, shorter than to as long as the tube; stamens included or exsert; ovary
one-locular, ovules many, in 2 parietal placentas; style as long as corolla tube or
191
distinctly longer, stigma capitate, slightly 2-lobed. Fruit a berry, apex with persistent remains of
style. Seeds many, irregularly angular (sections Cyrtophyllum and Racemosae) or ellipsoid-rounded
(section Fagraea).
Distribution. About 6070 species; Sri Lanka, India, Indo-China, China, Hainan, Taiwan, throughout
Malesia, to northern Australia and the Pacific. 42 species in Borneo; in Sabah and Sarawak 40
species, with F. crenulata and F. kalimantanensis as yet known only from Kalimantan.
Ecology. In primary or secondary forest, in open places, riverbanks and forest edges; sea-level to
about 3000 m.
Uses. Some species of the genus have timber value but many have potential in ornamental
horticulture.
Taxonomy. Three sections of the genus are recognised (Wong & Sugau l.c.). Section Fagraea has
axillary scales that develop above the petiole base and loosely clasp the stem; inflorescences of either
a solitary flower or that are cyme-like with the primary branches rebranching 13 times; fruits that are
typically large at maturity (always more than 25 mm across), with the epidermis detaching from the
pericarp on drying; and ellipsoid-rounded seeds. Section Cyrtophyllum has axillary scales that develop
at the very base of the petiole and tightly clasp the stem; cyme-like inflorescences with primary
branches that rebranch 36 times; smaller fruits (less than 20 mm across) with the epidermis not
detaching from the pericarp on drying and angular seeds. Section Racemosae has axillary scales, fruits
and seeds similar to that in section Cyrtophyllum but the inflorescence typically has all its branches
very condensed and grouped in distinct tiers along the main axis; in addition it appears to be
characterised by stamens and styles that are not or only slightly exsert in the open flower in
comparison to the conspicuously exsert stamens and styles in flowers of section Cyrtophyllum.
Recent studies (Struwe, pers. comm.) support the isolation of a few species of Fagraea s.l. from New
Guinea, Maluku, Australia, Polynesia and Malesia as a distinct genus, but the present review of
Fagraea in Borneo (Wong & Sugau l.c.) reveals insufficient evidence to treat the sections as separate
genera.
Leenhouts delimitation of the species has not been adhered to in most cases, notably in his concept of
F. auriculata, F. blumei, F. elliptica, F. fragrans, F. gardenioides and F. racemosa. F. ceilanica is
here considered absent in Malesia and is an artificial assemblage of a large number of species (see
Wong & Sugau l.c.).
Key to Fagraea species

1. Fruits typically large at maturity (always more than 25 mm across), the epidermis detaching from
the pericarp on drying. Inflorescence either without any branching (with only a solitary flower) or
with well-developed primary branches (which resemble the lower internodes of the main axis in
length) that rebranch typically only once but exceptionally (in F. floribunda) to 3 orders. Axillary
scales
developing
above
the
petiole
base
and
loosely
clasping
the
node...................................................................................................................................................2
192
Fruits always smaller (less than 20 mm across), the epidermis not detaching from the pericarp on
drying (sometimes the pericarp wrinkled). Inflorescence either pendulous or with all branches
very condensed and grouped as distinct tiers along the main axis, or with well-developed primary
branches that rebranch 36 orders. Axillary scales developing at the very base of the leaf stalk and
slightly clasping the node................................................................................................................28
2. Petiole bases typically developing distinct, rim-like auricles or large lobe-like auricles that are
usually reflexed..................................................................................................................................3
Petiole bases without auricles, or these auricles very indistinct and easily overlooked (F.
resinosa)............................................................................................................................................8
3. Leaf blade decurrent along the petiole, forming distinct wings several mm wide all along the
petiole, and continuing into the petiole base auricle (Flowers not known)......................
Fagraea sp. A
Wong & Sugau, Sandakania 8 (1996) 92.
Epiphytic shrub. Leaves large, base attenuate to decurrent. Flowers not known.
Borneo (Sarawak). Riverside forest, 100200 m.
Leaf blade not decurrent along the petiole, or only very slightly so and never more than 1 mm
wide, distinct from the petiole base auricle.......................................................................................4
4. Lateral veins on the lower leaf surface distinct and prominent. Flower subtended by an involucre
of large bracts. Corolla tube inside pale floccose (with loose long hairs) at the middle
part.............................................................................................................................
F. macroscypha Baker
Kew Bull. (1896) 65; Merrill l.c. (1921) 492; Masamune l.c. 614; Cockburn l.c. 210; Wong &
Sugau l.c. 77. Synonym: F. involucrata Merr. var. longipetiolata Merr., PEB (1929) 25,
Masamune l.c. 614.
Epiphytic shrub or climber, possibly a strangler. Leaves oblong-lanceolate, 1227 x 3.59.5
cm, thick-coriaceous. Flower terminal, solitary, corolla tubular, slightly widened upwards,
creamy white with broad green stripes on the outside, tube 1214 cm long.
Endemic to Borneo (Sabah, Sarawak, Brunei and Kalimantan). Mixed dipterocarp forest, to
200 m.
Lateral veins on the lower surface obscure or only very slightly distinct.Flowers not subtended by
any involucre or (in F. involucrata) with such an involucre. Corolla tube inside completely
glabrous or (in F. involucrata) pale floccose (with loose long hairs) at the middle
part.....................................................................................................................................................5
5. Leaf apex obtuse, rounded or with only an inconspicuous tip..........................................................6
Leaf apex acute, caudate or distinctly cuspidate...............................................................................7
6. Flowers solitary. Corolla tube 1314 cm long. Calyx lobes 3.54 cm long. Leaf blades typically
very large, 2829 x 12 13 cm................................................................................
F. megalantha Wong & Sugau
Sandakania 8 (1996) 77.
Climber. Leaves oblanceolate, thick coriaceous. Flower large, corolla funnel-shaped.
Borneo (Sarawak); known only from the type. Peat swamp forests.
193
Flowers 35 in a cyme. Corolla tube 69 cm long. Calyx lobes 2.53 cm long. Leaf blades
typically smaller, 1219 x 610 cm.......................................................................................
F. auriculata Jack
Mal. Misc. 2, 7 (1822) 82; Wong & Sugau l.c. 51; sensu Leenhouts, FM 1, 6 (1962) 326, pro
parte.
Climbing or terrestrial shrub or epiphyte. Leaves broad obovate, 1020 x 58 cm, thickcoriaceous. Flowers large, corolla infundibular, corolla tube 69 cm long, 11.5 cm wide,
corolla lobes ovate, 45 cm long, 2.53 cm wide.
Indo-China, Peninsular Malaysia, Java and Borneo (Sarawak and Kalimantan). Lowlands and
mountains, to 1930 m.
7. Flowers solitary, subtended by an involucre of bracts. Corolla tube inside pale floccose at the
middle part. Inflated upper part of corolla tube bell-shaped. Petiole base auricles large and
conspicuous, resembling lobes.......................................................................................
F. involucrata Merr.
J. Str. Br. R. As. Soc. 77 (1917) 233, l.c. (1921) 492, PEB (1929) 251 (excl. var.
longipetiolata Merr.); Masamune l.c. 614 (excl. var. longipetiolata Merr.); Cockburn l.c. 210;
Wong & Sugau l.c. 68. Synonyms: F. macroscypha sensu Heine, Pfl. Clemens Kinabalu
(1953) 91, non Baker (1896); F. uniflora Heine l.c. 92, non Merr. (1917).
Epiphytic shrub or liana, possibly a strangler. Leaves oblong to oblanceolate, 9.510 x 36
cm, coriaceous. Flower terminal, solitary, corolla infundibular, 1415 cm long, greenish
white.
Endemic to Borneo (Sabah, Sarawak and Kalimantan). Montane forests to 1800 m.
Flowers 35 in a cyme, without any involucre. Corolla tube inside completely glabrous. Inflated
upper part of corolla tube trumpet-shaped. Petiole base auricles 12 mm wide only, resembling
rims..........................................................................................................3. F. borneensis (typically)
8. Lateral veins on lower leaf surface distinct and prominent on drying..............................................9
Lateral veins on lower leaf surface indistinct, or sunken on drying...............................................12
9. Cyme subsessile, the flowers individually subtended by bracts 11.5 cm long forming a loose
involucre......................................................................................................................
F. iliasii Wong & Sugau
Epiphytic shrub. Leaves thin-chartaceous. Inflorescence a 3-flowered cyme; pedicels
indistinct (hidden by bracteoles).
Endemic to Borneo (Sarawak); known only from the type. On yellow sandy soil, river bank at
280 m.
Cyme distinctly pedunculate, the flowers not subtended by any unusually large
bracts................................................................................................................................................10
10. Inflorescence a laxly branched cyme 1416 cm long, the primary branches rebranching to 3
orders........................................................................................................................9. F. floribunda
Inflorescence with condensed branches or laxly branched but never exceeding 10 cm long, the
primary branches rebranching only once.........................................................................................11
194
11. Leaves thick-coriaceous, the petioles massive (58 mm thick). Inflorescence with condensed
branches. Calyx lobes in flower 1012 mm long.................................................
F. ridleyi King & Gamble
J. As. Soc. Beng. 74, 2 (1908) 612, non Gandoger (1924); Wong & Sugau l.c. 87.
Liana or big straggling shrub, twigs massive. Leaves broadly obovate to suborbicular.
Peninsular Malaysia, Lingga and Borneo (Sarawak and Brunei). Lowland primary forest, in
kerangas forest, sea-level to 300 m.
Leaves thin-coriaceous, the petioles slender (23 mm thick). Inflorescence with distinct elongate
branches. Calyx lobes in flower 58 mm long........................................................
F. renae Wong & Sugau
Sandakania 8 (1996) 82. Synonym: F. blumei sensu Cockburn l.c. 211, Anderson l.c. 239,
Ashton l.c. 312, pro parte, non G. Don (1837).
Strangling or epiphytic shrub. Leaves broadly elliptic to broadly oblanceolate, lateral veins 3
6 pairs, lower side prominent.
Peninsular Malaysia and Borneo (Sabah and Sarawak). Primary and secondary forests, also
on limestone; lowlands up to 1200 m in mossy forest.
12. Flowers subtended by an involucre of enlarged bracts..................................................................13
Flowers without any involucre at their base..................................................................................15
13. Calyx and most of the involucre bracts elliptic with rounded apices. Petiole bases developing
small, inconspicuous auricles......................................................................................15. F. resinosa
Calyx and most of the involucre bracts with acute-pointed apices. Petiole bases clearly without
auricles.............................................................................................................................................14
14. Leaf blades narrowly elliptic, the apex acute-caudate. Petioles longer, 2.24 cm long. Flowers 1
3, each on a short but distinct and thick pedicel; involucre bracts typically
keeled....................................................................................................................................
F. kuminii Wong & Sugau
Epiphytic shrub or woody climber. Leaves narrowly obovate to oblanceolate. Bracteoles in 2
3 decussate pairs, enclosing the lower part of the calyx, the inner or innermost pairs larger
than the outer ones.
Borneo (Sabah). On undulating land, river banks in disturbed forest on ultramafic soil.
Leaf blades obovate, the apex cuspidate. Petioles shorter, 0.52 cm long. Flowers solitary, the
pedicels hidden by the bracts and inconspicuous; involucre bracts without distinct
keels......................................................................................................................................
F. acutibracteata Wong & Sugau
Woody climber. Bracteoles with acute-acuminate apex; corolla unknown.
Borneo (Sarawak), so far known only from the type. On sandstone rocks at 550 m.
15. Flowers solitary..............................................................................................................................16
Flowers (2)several in a cyme.......................................................................................................19
195
16. Corolla tubular to narrowly infundibular (funnel-shaped), the tube 8.515 cm long............
F. carnosa Jack
Malay Misc. 2, 7 (1822) 81; Wong & Sugau l.c. 57. Synonym: F. uniflora Merr., J. Str. Br. R.
As. Soc. 77 (1917) 235, l.c. (1921) 493, Masamune l.c. 615.
Woody climber or epiphyte. Leaves broadly elliptic to obovate, thick-coriaceous.
Lower Burma, Sumatra, Peninsular Malaysia and Borneo (Sarawak). On sandstone soils in
mixed dipterocarp forest, sea-level to 200 m.
Corolla short-infundibular, the tube 34.5 cm long........................................................................17
17. Corolla lobes larger, 3.54 cm long, 22.5 cm broad. Petioles 1.23 cm and leaves
elliptic....................................................................................................................................
F. stonei Wong & Sugau
Woody climber or epiphytic shrub. Leaves broadly elliptic to lanceolate, thick-coriaceous.
Flower solitary.
Borneo (Sabah and Sarawak). In kerangas forest on white sand podsols, also on basalt ridges.
Corolla lobes smaller, 1.83 cm long, 1.32.5 cm broad. Petioles less than 1 cm long
if leaves elliptic (F. kinabaluensis), or when longer (12.2 cm) then leaves distinctly
obovate.............................................................................................................................................18
18. Leaves obovate, the petioles 12.2 cm long. Upper part of corolla tube at most 1 cm across at the
mouth...........................................................................................................................7. F. dulitensis
Leaves elliptic or only very slightly obovate, the petioles less than 1 cm long. Upper part of
corolla tube more flared, 22.5 cm across at the mouth.....................................12. F. kinabaluensis
19. Inflorescence a tight cluster of many flowers, the branches condensed and hidden by the flowers
themselves (in fruit the axes visible but still short, supporting many fruits).............
F. splendens Blume
Mus. Bot. Lugd. Bat. 1 (1850) 168; Wong & Sugau l.c. 87. Synonyms: F. heterophylla
Blume, l.c. (1850) 168; F. acuminatissima Merr., J. Str. Br. R. As. Soc. 77 (1917) 232, l.c.
(1921) 491, Masamune l.c. 613, Cockburn l.c. 211, Anderson l.c. 239, Ashton l.c. 311; F.
rostrata sensu Merrill l.c. (1921) 493, Masamune l.c. 615, non Blume (1836); F. ceilanica
sensu Leenhouts, FM 1, 6 (1962) 315, pro parte, non Thunb. (1782).
Climber or epiphyte, rarely a small tree to 6 m tall. Leaves oblanceolate or obovate, 1125.5
4.59.5 cm, coriaceous. Inflorescence a 38(10)-flowered sessile cyme, corolla tubular, 4
4.5 cm long, lobes 1.52 cm long.
Sumatra, Peninsular Malaysia, Borneo (Sabah, Sarawak and Brunei). On various types of
soils, including in swamp forest, and especially kerangas forest, from sea-level to 1333 m.
Inflorescence a few-flowered cyme or, if many-flowered, the branches distinct and not hidden by
the flowers.......................................................................................................................................20
20. Leaf apex obtuse-rounded. Leaves drying black or dark brown............................................
196
F. tuyukii Wong & Sugau

Probably epiphytic shrub to small tree. Leaves elliptic to obovate, 48 x 47 cm, stiffcoriaceous. Flower unknown. Infructescence terminal, of a few fruits in a cyme. Fruit
ellipsoid, mucronate apically, 2.53 cm long, 2.52.7 cm wide.
Endemic to Borneo (Sabah); known only from the type. Mixed dipterocarp forests, 700800
m.
Leaf apex acute or cuspidate or caudate. Leaves drying pale to dark greenish brown...................21
21. Corolla tube more than 10 cm long............................................................3. F. borneensis (in part)
Corolla tube less than 8 cm long.....................................................................................................22
22. Corolla tubular, the tube more than twice as long as the lobes...............................................
F. havilandii Wong & Sugau
Sandakania 8 (1996) 64. Synonym: F. gardenioides ssp. borneensis Leenhouts, Bull. Jard.
Bot. Brux. 32 (1962) 425.
Probably shrub or epiphyte. Leaves obovate, 811 x36 cm, coriaceous. Flowers 25 in a
cyme, corolla tubular, tube 5.5 cm long, lobes 11.8 cm long.
Borneo (Sarawak, near Kuching).
Corolla infundibular, the tube less than twice as long as the lobes................................................23
23. Corolla throat 58 mm across........................................................................................................24
Corolla throat 1020 mm across....................................................................................................26
24. Leaf blades decurrent to the base of their petioles, which are broadly winged. Ultimate branches
developing broadened, somewhat corky and very short internodes that form a series of coarse
cicatrices (bands) along the branches ..........................................................
F. oreophila Wong & Sugau
Sandakania 8 (1996) 78. Synonym: F. ceilanica sensu Leenhouts, FM 1, 6 (1962) 315,
Cockburn l.c. 211, pro parte, non Thunb. (1782).
Epiphyte, possibly also a small tree. Leaves obovate, 79 x 2.54 cm, coriaceous.
Inflorescence terminal, 3-flowered; corolla pale yellow or white, tube c. 2.5 cm long, lobes c.
1.51.8 cm long.
Sabah (on Mt. Kinabalu) and Sarawak (Mt. Murud). Mossy montane forest.
Leaf blades not completely decurrent along their petioles, which are always distinctly slender.
Ultimate branches not so, such nodal cicatrices if developing always far
apart.................................................................................................................................................25
25. Leaves small, to 9 x 2.5 cm, the apex caudate. Corolla tube 1820 mm long........................
F. rarissima Wong & Sugau
Sandakania 8 (1996) 82. Synonym: F. ceilanica sensu Leenhouts, FM 1, 6 (1962) 315, pro
parte, non Thunb. (1782).
Shrub. Leaves narrowly elliptic, 48.5 x1.52.5 cm, coriaceous. Inflorescence a 23flowered cyme; corolla salverform, white, tube 22.5 cm long, lobes 1.51.7 cm long.
Borneo (Sarawak and Brunei). On yellow podsols in mixed dipterocarp forests.
Leaves typically larger, 815 x 36.5 cm, the apex acute to cuspidate. Corolla tube 2030 mm
long.................................................................................................................................2. F. blumei
197
26. Corolla tube with a base 57 mm across, gradually flared upwards. Leaves smaller typically, at
most 3 cm wide..................................................................................................
F. longipetiolata Wong & Sugau
Climbing shrub. Leaves elliptic to narrowly obovate, margin slightly recurved when
dry.
Borneo (Sarawak and Brunei). Primary mixed dipterocarp forest.
Corolla tube with a base only 12 mm across, abruptly flared upwards. Leaves larger typically, 3
10 cm wide......................................................................................................................................27
27. Flower calyx lobes small, 24 mm long. Leaves narrowly obovate-elliptic to oblong. Upper leaf
surface parchment-like when dry.........................................................................
F. oblonga King & Gamble
J. As. Soc. Beng. 74, 2 (1908) 612; Wong & Sugau l.c. 78.
Climbing shrub. Leaves with slightly recurved margin, apex obtuse.
Peninsular Malaysia, Sumatra and Borneo (Sarawak). Lowlands to mountains, also on
limestone.
Flower calyx lobes larger, 78 mm long. Leaves broadly ovate to elliptic. Upper leaf surface
coarsely shagreen when dry......................................................................................
F. littoralis Blume
Bijdr. Fl. Ned. Ind. (1826) 1021, Merrill l.c. (1921) 492, Masamune l.c. 614, Wong & Sugau
l.c. 73.
Climbing shrub. Leaf base rounded; midrib prominent, sharp-ridged.
Borneo, all territories. Mostly near river and lakes.
The variety in Borneo is var. borneensis Wong & Sugau, l.c. 73, wrongly attributed by both
Merrill and Masamune to Miquels var. forstenii (a synonym of var. amboinensis Blume)
from Sulawesi.
28. Inflorescence pendulous or typically with all branches very condensed and grouped in distinct
tiers along the main axis. Stigma capitate but at maturity developing an expanded rim that gives it
a peltate structure. Stamens and style not or only slightly exsert in the open
flower...............................................................................................................................................29
Inflorescence rigid and erect, with well-developed primary branches (as long as the lower
internodes of the main axis) that rebranch 36 orders. Stigma capitate throughout, an expanded
rim not developing. Stamens and style long-exsert in the open flower...........................................35
29. Narrowed basal tubular part of corolla very short, much less than a third the length of the corolla
tube, and mostly hidden by the calyx in the open flower...........................................6. F. cuspidata
Narrowed basal tubular part of the corolla longer, more than a third or half the corolla tube, and
clearly extending beyond the calyx in the open flower...................................................................30
30. Corolla lobes in the open flower relatively short, at most up to about a third the length of the
expanded upper part of the corolla tube. Leaves subsessile..........................................17. F. spicata
Corolla lobes in the open flower relatively longer, about half or more the length of the expanded
upper part of the corolla tube. Leaves with distinct petioles...........................................................31
31. Corolla of open flowers relatively narrow (the uppermost part of the tube only 68 mm
wide)................................................................................................................................................32
198
Corolla of open flowers wider (the uppermost part of the tube 10 mm or wider)..........................33
32. Corolla of open flowers 2027 mm long (the narrowed basal tubular part 810 mm long, the
inflated upper part 1011 mm long). Anthers 22.5 mm long. Leaves linear, 1227 cm long, with
714
pairs
of
lateral
veins
faintly
visible.
(Rheophytic
plants
of
the
lowlands)................................................................................................................18. F. stenophylla
Corolla of open flowers 1820 mm long (the narrowed basal tubular part 78 mm long, the
inflated upper part 56 mm long). Anthers c. 1 mm long. Leaves elliptic-obovate, 3.513.5 cm
long, with 45 pairs of lateral veins faintly visible. (Montane plants)
....................................................................................................................................13. F. montana
33. Mature flowers with calyx 46 mm long; inflated upper part of the corolla tube 710 mm long,
shorter than or about as long as the basal tubular part................................................20. F. volubilis
Mature flowers with calyx 79 mm long; inflated upper part of corolla tube 1216 mm long,
much longer than the basal tubular part..........................................................................................34
34. Uppermost part of the corolla tube only 1012 mm wide; corolla lobes 78 mm long. Anthers c.
1.5 mm long. (Leaves typically smallish, the length at least 3 times the
width.)......................................................................................................................19. F. teysmannii
Uppermost part of the corolla tube 2022 mm wide; corolla lobes 1012 mm long. Anthers 3.54
mm long. (Leaves typically large, sometimes small, the length typically less than 3 times the
width.)................................................................................................................14. F. philippinensis
35. Inflorescences exclusively axillary.................................................................................................36
Inflorescences terminal...................................................................................................................38
36. Inflorescence a 3-branched cyme, or (sometimes) reduced unbranched inflorescence, (1)3flowered. Corolla tube 1012 mm long. Leaves coriaceous, the margins never wavy, with lateral
veins 36 pairs...............................................................................................................4. F. caudata
Inflorescence a many-branched, many-flowered cyme. Corolla tube 68 mm long. Leaves
chartaceous with wavy margins, or leaves (sub)coriaceous with lateral veins 912 pairs.............37
37. Inflorescence peduncle 46 cm long. Open flowers with calyx diameter 22.5 mm, corolla lobes
(5)67 mm long and 34 mm wide, and styles exsert for 810 mm. Leaves with 912 pairs of
lateral veins which are flat on the upper surface, and plane margins when
fresh............................................................................................................................10. F. fragrans
Inflorescence peduncle 23.5 cm long. Open flowers with calyx diameter 1.52 mm, corolla lobes
45 mm long and 2.53 mm wide, and styles exsert for 1416 mm. Leaves with 38 pairs of
lateral veins which are often depressed on the upper surface, and wavy margins when
fresh............................................................................................................................11. F. gigantea
38. Cymes with main axis branching 46 orders. Corolla lobes ovate, 2.53 mm long. Leaf
apex typically obtuse-rounded to emarginate. Tree mostly of lowland secondary forest and open
sites, and forest gaps......................................................................................................1. F. belukar
Cymes with main axis typically branching only 34 orders (very exceptionallly an occasional 5th
order of branching present). Corolla lobes lanceolate, c. 3.5 mm long or more. Leaf apex
acuminate to cuspidate-caudate. Tree typically of primary forest in the lowlands or
mountains.........................................................................................................................................39
199
39. Calyx in flower 34 mm long. Corolla lobes 58 mm long. Stamen filaments in open flowers 15
18 mm long. Mature fruits (6)910 mm across. Leaf surfaces drying very coarsely wrinkled and
very coarse to the touch..............................................................................................16. F. rugulosa
Calyx in flower 23 mm long. Corolla lobes 3.56 mm long. Stamen filaments in open flowers 7
11 mm long. Mature fruits smaller, only 3.55(6) mm across. Leaf surfaces drying
smooth..............................................................................................................................................40
40. Flowers sessile to subsessile, the pedicels (if developed) to 1 mm long. Leaf apex acuminate. Leaf
lateral veins obscure to only very faintly visible, never elevated on the lower side. (Leaves
coriaceous. Corolla tube 67 mm long)...........................................................................5. F. collina
Flowers with distinct pedicels 13 mm long. Leaf apex cuspidate to caudate. Leaf lateral veins
distinct and raised on the lower side. (Leaves chartaceous to coriaceous. Corolla tube 611 mm
long)...............................................................................................................................8. F. elliptica
1. Fagraea belukar Wong & Sugau
Fig. 1.
(Malay, belukar = secondary forest; the usual habitat)

Sandakania 8 (1996) 15. Type: Saikeh SAN 72151, Sabah, Beaufort, Beaufort Hill (holotype SAN). Synonym: F. elliptica
sensu Leenhouts, FM 1, 6 (1962) 303, Cockburn l.c. 210, Ashton l.c. 314, Anderson l.c. 239, pro parte, non Roxb. (1824).
Tree to 30 m tall and 60 cm diameter or more, trunk base fluted-cylindric. Bark fissured, greyish
black; inner bark reddish-yellow. Sapwood yellowish white. Leaves coriaceous, upper and lower
surfaces finely shagreen or smooth; obovate to broad-elliptic, 722 x 413 cm; base cuneate, margin
plane to recurved, apex obtuse-rounded to emarginate; midrib prominent on lower side, lateral veins
1012 pairs, faintly visible and immersed (never raised) on the lower side, faintly visible, flat,
immersed or slightly sunken on the upper side, intercostal veins obscure; petioles 34.5 cm long,
stout, the base with axillary scales forming a cup-shaped ochrea tightly clasping the node.
Inflorescence terminal, a many-flowered branched cyme, 13 cm long, 1519 cm wide, main axis
branching 46 orders; peduncles 23.5 cm long. Flowers with pedicels 34 mm long; calyx
campanulate, 12 mm long, 12 mm diameter, divided to the middle; corolla salverform, tube 710.5
mm long, 11.5 mm wide, lobes ovate, 2.53 mm long, 1.5 mm wide; stamens 810 mm long,
filaments 68 mm long, inserted at the mouth of corolla, exserted for 89 mm, anthers oblong, c. 1
mm long; style exsert for 49 mm, stigma small, capitate, very obscurely 2-lobed. Fruits globose,
3.55 mm across, tipped with a minute circular scar of the fallen style; fruit calyx 11.5 mm long, 1
1.5 mm wide. Seeds c. 1 mm diameter, angular, brownish black.
Vernacular names. Sabahtamasuk hutan (Brunei Malay), tembusu bukit (Malay), tembusu tagai
(Malay).
Distribution. Borneo, in all districts including Sabah and Sarawak, and Banka island. Very common
throughout Sabah and Sarawak.
Ecology. Lowland secondary forests, open sites and forest gaps.
Uses. Apparently a durable wood for house and bridge construction.
200
Fig. 1. Fagraea belukar. A, flowering leafy twig; B, flower; C, corolla lobe; D, fruit. (AC from SAN 72151, D from SAN
72256.)
201
2. Fagraea blumei G. Don

(C.L. Blume, 17961862, Dutch Botanist)
Gard. Dict. 4 (1837) 69; Leenhouts, FM 1, 6 (1962) 320, Cockburn l.c. 210, Anderson l.c. 239, Ashton l.c. 312, pro parte;
Wong & Sugau l.c. 54. Type: Blume, s.n. (= Leiden sheet no. 908. 127-758) (L), Java. Synonyms: F. obovata sensu Blume,
l.c. (1826) 1021, nom. illeg., non Wall. (1824), Masamune l.c. 614; F. obovato-javana Blume, Rumphia 2 (1838) 29, t. 75,
incl. var. bebeak Blume; F. vaginata King & Gamble, J. As. Soc. Beng. 74, 2 (1908) 610, pro parte; F. obovata sensu Bakh.
f. in Backer, Bekn. Fl. Java 7 (1948) fam. 170: 11, incl. var. brevicalyx Bakh. f., Blumea 6 (1950) 382.
Tree to 20 m tall, 20 cm diameter, sometimes a climbing shrub or epiphyte. Leaves coriaceous, upper
and lower surfaces shagreen, smooth; elliptic, obovate to oblanceolate, 815 x36.5 cm; base cuneate
to attenuate, margin plane, apex acute to cuspidate; midrib prominent on lower side, lateral veins 23
pairs, obscure to only rather faintly visible on the lower side, obscure on the upper side, intercostal
veins obscure; petioles 13 cm long, base with conspicuous axillary scales loosely clasping the node.
Inflorescence terminal, a few-flowered cyme, 47 cm long; peduncle nil to 2 cm long, smooth or
lenticellate. Flowers with pedicels 0.52 cm long; bracteoles small, 13 mm long, attached halfway
on the pedicel; calyx 1.22.9 cm long, 0.81 cm wide; corolla narrowly to widely infundibular, 4.5
6.5 cm long, divided to about halfway down or slightly deeper, tube 23 cm long, lobes 33.4 cm
long; stamens c. 2 cm long, anthers oblong, 57 mm long; style 2.53 cm long, stigma capitate, 15
mm diameter. Fruits elipsoid, c. 4 cm wide, fruit calyx 1.33 cm long, 1.42 cm wide, lobes patent.
Seeds ellipsoid-rounded.
Distribution. Sumatra, Peninsular Malaysia, Java and Borneo (Sabah and Sarawak). In Sabah,
recorded from the Tambunan, Kota Belud and Ranau districts; in Sarawak, documented from the Miri
district but probably more common.
Ecology. Primary forest, mostly in highlands.
3. Fagraea borneensis Scheff.

(of Borneo)
In Hasskarl, Flora 52 (1869) 309; Merrill l.c. (1921) 491; Masamune l.c. 613; Wong & Sugau l.c. 55. Type: Lobb, s.n.,
Sarawak (K). Synonyms: F. auriculata ssp. borneensis (Scheff.) Leenh., FM 1, 6 (1962) 328, Cockburn l.c. 210, pro parte;
F. nonok Elm., Leafl. Philip. Bot. 3 (1910) 858; F. resinosa sensu Leenhouts, FM 1, 6 (1962) 331, pro parte.
Epiphytic shrub or tree, to c. 13.5 m tall, c. 30 cm diameter. Bark smooth, grey; inner bark green.
Wood whitish. Leaves thick-coriaceous, upper and lower surfaces shagreen, rough; oblanceolate,
elliptic to broad obovate, 825 x 410 cm; base attenuate to cuneate, margin plane to slightly recurved
when dry, apex acute to acuminate; midrib prominent on lower side, lateral veins 68(11) pairs,
indistinct to slightly distinct (not prominent) on both sides, intercostal veins obscure; petioles 23(4)
cm long, base with conspicuous axillary scales loosely clasping the node, auricles indistinct or 12
mm wide. Inflorescence terminal, a 35-flowered cyme, sessile. Flowers with pedicels 1.52 cm
long; bracteoles 1 pair, c. 0.5 cm long, attached to calyx cup base; calyx 33.5 cm long, 1.52 cm
diameter; corolla slender, infundibular, tube 911 cm long, 0.83.5 cm diameter, glabrous inside,
202
upper inflated part trumpet-shaped, lobes ovate, 23.5 cm long, 12 cm wide; stamen filaments 55.5
cm long, inserted c. 44.5 cm down from the corolla mouth, anthers c. 0.5 cm long; style 1111.5 cm
long, stigma peltate, c. 3 mm diameter. Fruits ellipsoid, 55.5(7) cm long, 22.5 cm wide; fruit
calyx lobes 22.5 cm long, 11.5 cm wide, patent. Seeds ellipsoid-rounded.
Distribution. Borneo (Sabah, Sarawak, Brunei and Kalimantan) and the Philippines.
Ecology. Lowland and montane forest, also in swampy areas and on limestone. Once recorded from
seaside rocks.
4. Fagraea caudata Ridl.

(Latin, caudatus = caudate; the attenuate leaf apex)
J. Str. Br. R. As. Soc. 79 (1918) 97; Merrill l.c. (1921) 491; Masamune l.c. 613; Wong & Sugau l.c. 8. Type: Lobb, s.n.,
(1858), Borneo (Sarawak) (K). Synonym: F. fragrans sensu Leenhouts, FM 1, 6 (1962) 304, Ashton l.c. 310, pro parte,
non Roxb. (1824).
Small tree, to 6 m tall, 4.5 cm diameter. Bark fissured and flaky, dark brown; inner bark orange.
Wood pale brown. Leaves coriaceous, upper and lower side smooth; elliptic to obovate, 1.54 x 612
cm; base acute, decurrent, margin plane, apex caudate; midrib prominent on lower side, lateral veins
36 pairs, prominent on lower side, obscure or faint and impressed on upper side, intercostal veins
obscure; petioles 0.71.5 cm long, base with axillary scales adnate to and tightly clasping the node.
Inflorescence axillary, a (1)3-flowered branched cyme, or (sometimes) a reduced unbranched
inflorescence, 69 cm long; peduncle 2.55 cm long, branches (when present) 1.83 cm long.
Flowers with pedicels 510 mm long; calyx cylindric, 44.5 mm long, divided to halfway, 2.53 mm
diameter; corolla salverform, tube 1012 mm long, 45 mm wide, lobes 6.58 mm long, 45 mm
wide, ovate; stamen filaments 1013 mm long, inserted just above the middle of the corolla tube,
anthers brownish green; style exsert for 812 mm, stigma capitate. Fruits broadly ellipsoid, 810 mm
long, 56 mm wide; fruit calyx 3.54 mm long, 3.54 mm wide. Seeds angular.
Distribution. Endemic to Borneo (Sarawak and Brunei). In Sarawak, recorded only from the Kuching
and Miri districts and in Brunei, documented for the Batu Patam area in Belait district.
Ecology. Mixed dipterocarp forest, sea-level to 285 m.
5. Fagraea collina Wong & Sugau

(Latin, collinus = pertaining to hills; the typical habitat)
Sandakania 8 (1996) 19. Type: Aban SAN 50747, Sabah, Ranau, copper mining area (holotype SAN). Synonym: F. elliptica
sensu Leenhouts, FM 1, 6 (1962) 303, Cockburn l.c. 210, Ashton l.c. 314, Anderson l.c. 239, pro parte, non Roxb. (1824).
Shrub to medium-sized tree, to 15(20) m tall, 10(25) cm diameter. Bark fissured, dark brown to
black. Sapwood yellowish. Leaves coriaceous, upper and lower surfaces smooth; elliptic,
oblanceolate-obovate, 415 x 27 cm; base cuneate, margin plane to slightly recurved, apex
acuminate; midrib prominent on lower side, lateral veins 79 pairs, faint to obscure on both sides,
203
intercostal veins obscure; petioles 1.52 cm long, base with axillary scales adnate to the twig and
forming a nodal ochrea. Inflorescence terminal, a many-flowered cyme, 1118 cm long, main axis
branching to 34 orders; peduncle 13.5 cm long, first branch 2.57 cm long. Flowers with pedicels
01 mm long; calyx campanulate, 23 mm long, 1.52 mm diameter, divided to almost halfway or
more; corolla salverform, tube 67 mm long, 11.5 mm wide, lobes ovate to lanceolate, 3.54 mm
long, 11.5 mm wide; stamen filaments 811 mm long, inserted at the corolla mouth, anthers c. 1 mm
long, exsert for 67 mm; style exsert for 57 mm, stigma capitate. Fruits globose, 3.55 mm across.
Seeds angular, c. 1 mm diameter, brownish black.
Distribution. Borneo, so far known only from Sabah (around Ranau and on Gaya Island on the west
coast) and Sarawak (Kuching, Sibu, Bintulu and Kapit districts).
Ecology. Mostly in montane forest to 1700 m, sometimes lowland (Gaya island, Sabah).
6. Fagraea cuspidata Blume

(Latin, cuspidatus = with an abrupt, short point; the leaf apex)
Mus. Bot. Lugd. Bat. 1(1850) 170; Merrill l.c. (1921) 491; Masamune l.c. 614; Wong & Sugau l.c. 28. Type: Blume, s.n.
(Leiden sheets no. 908.127-738, 908.127-740 & 944.202-211), Borneo, Tanjong Java (isotype L). Synonyms: F. robusta
Blume, l.c. (1850) 170; F. crassipes Benth., J. Linn. Soc. Bot 1 (1856) 99; F. cymosa Merr., J. Str. Br. R. As. Soc. 77 (1917)
234, l.c. (1921) 492, Masamune l.c. 614; F. pendula Merr. l.c. (1929) 251, Masamune l.c. 615; F. racemosa sensu Merrill
l.c. (1921) 493, Masamune l.c. 615, Leenhouts, FM 1, 6 (1962) 311, Anderson l.c. 240, Ashton l.c. 314, pro parte, non Jack
ex Wall. (1824).
Small tree to 18 m tall, 12 cm diameter. Bark smooth to slightly fissured, dark grey. Leaves
coriaceous, upper and lower surfaces smooth; ovate-elliptic, 840 x 4.516 cm; base cuneate-rounded
to cordate, margin plane, apex abruptly short-caudate; midrib prominent on lower side, lateral veins
69 pairs, prominent on the lower side, faint to obscure on the upper side, intercostal veins obscure
to prominent; petioles 0.52 cm, base with axillary scales fused to form a nodal ochrea. Inflorescence
terminal, 5.534 cm long, with clusters of condensed branches in distinct tiers along the main axis,
peduncle 2.522 cm long. Flowers with pedicels 735 mm long; calyx 79 mm long, 78 mm wide,
lobes 45 mm long; corolla narrowly funnel-shaped, 3459 mm long, narrowed basal part 56 mm
long, expanded upper part 2224 mm long, 1118 mm wide, lobes semi-orbicular to ovate, 711 mm
long, 6.59 mm wide; stamen filaments exsert for 23 mm, anthers 56 mm long; style exsert for 8
10 mm, stigma capitate. Fruits ovoid-ellipsoid, 1318 mm long, 914 mm wide; fruit calyx lobes
clasping the fruit base. Seeds angular.
Vernacular names. Sabahtodopon puak (Dusun). Sarawaksukang ranyai (Iban), sukong ganyai
(Iban), tembusu gajah (Malay), tinggirang pirak (Kedayan).
Distribution. Known only from Borneo (all districts) and the Philippines (Balabac island). Very
common.
Ecology. Mixed dipterocarp forest, in gaps and clearings, also secondary forests and forest fringes.
Lowlands to 1500 m.
204
7. Fagraea dulitensis Wong & Sugau

(of Mt. Dulit, Sarawak)
Sandakania 8 (1996) 59. Type: Tong S. 34870, Sarawak, 4th Div., Marudi, Ulu Sg. Tinjar, Dulit Range, near Koyan
(holotype SAN, isotypes K, KEP, L, MO, SAR).
Tree, to 5 m tall, 5 cm diameter. Bark brown. Leaves thin-coriaceous, upper and lower
surfaces smooth; obovate, 51.5 x 24.5 cm; base attenuate to cuneate, margin plane, apex
acute or shortly acute-acuminate; midrib prominent on lower side, lateral veins obscure on
both sides, intercostal veins obscure; petioles 12.2 cm long, base with conspicuous axillary
scales loosely clasping the node. Inflorescence a solitary flower. Flowers subsessile or with
pedicels to 0.5 cm long; bracteoles 12 decussate pairs, tiny (only up to 5 mm long); calyx 2
cm long, 11.3 cm diameter, divided more than halfway down (in bud), lobes ovate, c.1.4 cm
long; corolla greenish-yellow, short-infundibular, tube to 4.5 cm long, the upper part to 10
mm wide, lobes white, to 1.8 cm long, 1.5 cm wide; anthers brown; pistil light green. Fruit
unknown.
Vernacular name. Sarawakjatem (Kenyah).
Distribution. Borneo, known only from the type specimen from Sarawak.
Ecology. Recorded in old secondary forest, at c. 113 m.
8. Fagraea elliptica Roxb.

(Latin, ellipticus = elliptic; the leaf shape)
Fl. Ind. ed. Wall. 2 (1824) 32; Leenhouts, FM 1, 6 (1962) 303, Cockburn l.c. 210, Anderson l.c. 239, Ashton l.c. 314, pro
parte; Wong & Sugau l.c. 21. Type: not designated (see Wong & Sugau l.c.). Synonyms: Picrophloeus javanensis Blume,
l.c. (1826) 1020, nom. nud.; Cyrtophyllum speciosum Blume, l.c. (1826) 1022; Willughbeia elliptica (Roxb.) Spreng., Syst.
Veg. 4 (1827) Cur. Post. 71; F. speciosa (Blume) Blume, Rumphia 2 (1838) 35, t. 81, Mus. Bot 1 (1850) 172, Merrill l.c.
(1921) 493, Masamune l.c. 615; F. picrophloea Blume, l.c. (1838) 36, nom. illeg., Merrill l.c. (1921) 492, Masamune l.c.
615; F. kimangu Blume, l.c. (1850) 173; F. valida Miq., Fl. Ind. Bat. 2 (1857) 376; F. sumatrana Miq. l.c. 377; F.
aurantiodora S. Moore, J. Bot. 66 (1928) 105; F. pseudoelliptica Kanehira & Hatusima, Bot. Mag. Tokyo 56 (1942) 161, f.
5; F. javensis (Blume) Bakh. f. in Backer, Bekn. Fl. Java 7 (1948) fam. 170, 12, nom. illeg.; F. pusilliflora Bakh. f. in Backer
l.c. 13, nom. nud.
Shrub to tree, to 25 m tall, 32 cm diameter or more, sometimes with buttresses to 45 cm high. Bark
fissured, pale brown to dark grey-brown; inner bark cream. Leaves chartaceous to coriaceous, upper
and lower surfaces smooth when dry; elliptic, oblanceolate-obovate, 615 x 38 cm; base cuneate,
margin plane to slightly recurved, apex cuspidate-caudate; midrib prominent on lower side, lateral
veins 812 pairs, distinct and raised on the lower side, flattened and indistinct to slightly raised on the
upper side, intercostal veins obscure; petioles 1.54 cm long, base with axillary scales adnate to the
twig and fused to form a nodal ochrea. Inflorescence terminal, a many-flowered branched cyme, 9
12 cm long, 621 cm wide, main axis branching 34 orders (occasionally to 5 orders); peduncle 3
6.5 cm long. Flowers with pedicels 13 mm long; calyx campanulate, 23 mm long, divided to about
halfway, 22.5 mm diameter; corolla salverform, whitish, tube 611 mm long, 11.5 mm wide, lobes
205
lanceolate, 3.56 mm long, 12 mm wide; stamen filaments 710 mm long, inserted at corolla mouth,
anthers oblong, c. 1 mm long; style exsert for 46 mm, stigma small, capitate. Fruits globose, 3.56
mm across, orange to brick-red, fruit calyx lobes 12 mm long. Seeds angular.
Distribution. Sumatra, Java, Borneo, Sulawesi, Maluku and New Guinea. In Borneo, in all territories.
In Sabah, documented for the Ranau and Penampang districts, in Sarawak, for the Miri and Kapit
districts. Uncommon.
Ecology. Lowland primary forest, occasionally montane forest to 1600 m (on Mt. Kinabalu).
9. Fagraea floribunda Wong & Sugau
Fig. 2.
(Latin, floribundus = profusely flowering)

Sandakania 8 (1996) 62. Type: Ashton S. 16743, Sarawak, Ulu Anap, Bt. Mersing (holotype SAN, isotypes BO, K, KEP, L,
MEL, SAR, SING). Synonym: F. blumei sensu Leenhouts, FM 1, 6 (1962) 320, pro parte, non G. Don (1837).
Tree, to 23 m tall, 60 cm diameter, sometimes with stilt-roots. Bark smooth, with white hoop marks.
Leaves subcoriaceous, lower surface shagreen, upper surface smooth; broadly oblanceolate, 513.5
x36 cm; base cuneate-attenuate, margins plane, apex broadly acute; midrib prominent on lower side,
lateral veins 45 pairs, prominent on lower side, impressed or sunken on upper side, intercostal veins
obscure; petioles 23 cm long, base with conspicuous axillary scales loosely clasping the node.
Inflorescence terminal, a compound cyme, laxly branched, short-pedunculate, 1416 cm long,
primary branches rebranching to 3 orders. Flowers with pedicels 0.81 cm long, 34 mm thick;
bracts subtending inflorescence branches 56 mm long; bracteoles 1 pair, small, 12 mm long,
attached halfway on the pedicel; calyx campanulate, 0.91.2 cm long, lobes rounded, 0.30.5 cm
across; corolla creamy yellow, c. 2.7 cm long, lobes 11.4 cm long, 34 mm wide; stamen filaments
11.5 cm long, anthers 34 mm long; style 0.91 cm long, stigma capitate. Fruits oblong, to 5 x 4
cm. Seeds ellipsoid.
Distribution. Borneo (Sarawak, documented only for the Bintulu and Mukah districts).
Ecology. Mixed dipterocarp forest and kerangas forest, also on basalt boulders, up to 700 m.
10. Fagraea fragrans Roxb.

(Latin, fragrans = fragrant; the flowers)
Fl. Ind. ed. Wall., 2 (1824) 32; Merrill l.c. (1921) 492; Masamune l.c. 614; Cockburn l.c. 211; Leenhouts, FM 1, 6 (1962)
304, Anderson l.c. 240, Ashton l.c. 310, pro parte; Wong & Sugau l.c. 11. Type: Hunter, s.n. (= Wallich, Cat. no. 1597E),
Pullo Penang (K). Synonyms: Cyrtophyllum peregrinum Reinw., Syll. Pl. Nov. Soc. Bot. Ratisb. 2 (1826) 9, Blume, Bijdr.
Fl. Ned. Ind. (1826) 1022; F. peregrina Blume, Rumphia 2 (1838) 34, t. 80, Mus. Bot. Lugd. Bat. 1 (1850) 172; F.
cochinchinensis A. Chevalier, Cat. Pl. Jard. Bot. Saigon (1919) 65, pro specim., excl. basionyms.
Medium-sized to big tree, to 35 m tall, 1.2 m diameter. Bark irregularly fissured, dark brown to
greyish to black; inner bark yellowish, fibrous. Wood pale yellow. Leaves subcoriaceous,
206
Fig. 2. Fagraea floribunda. A, flowering leafy twig; B, flower; C, fruit. (A, B from S. 16743; C from S. 19455.)
207
upper and lower surface smooth; elliptic, 414 x 1.55 cm; base acute, decurrent on the petiole,
margin plane, apex cuspidate; midrib prominent on the lower side, lateral veins 912 pairs, distinct,
flat to only slightly prominent on the lower side, flat and indistinct to obscure on the upper side,
intercostal veins obscure; petioles 0.51.8 cm long, base with axillary scales adnate to the twig and
fused to form a nodal ochrea. Inflorescence axillary, a many-flowered cyme, 6.512 cm long, 47 cm
wide, main axis branching 23 orders; peduncle 46 cm long. Flowers with pedicels 57 mm long;
calyx campanulate, 22.5 mm long, 22.5 mm diameter; corolla infundibular, pale yellow, fragrant,
tube 68 mm long, 1.52 mm wide, lobes ovate, (5)67 mm long, 34 mm wide; stamens exsert,
filaments 1417 mm long, inserted just above the middle of the tube, anthers oblong-linear, c. 1.5 mm
long; style exsert for 810 mm, stigma faintly 2-lobed. Fruits ovoid-globose, 56 mm long, 56 mm
wide, orange then scarlet; fruit calyx 22.5 mm long, 2.53 mm wide. Seeds minute, angular.
Vernacular names. Sabahbanati (Keningau Murut), ombinaton (Dusun), tambiaton, tambinaton
(Dusun), temasuk, temasuk laut, temasuk pasir (Brunei Malay). Sarawaktembusu (Malay).
Distribution. Bengal, Myanmar, Thailand, Indo-China, Andamans, Sumatra, Java, Peninsular
Malaysia, Mindoro, Balabac, Palawan, Borneo and Sulawesi. In Borneo, in all territories. In Sabah
and Sarawak common only in the west coast districts, elsewhere mostly planted.
Ecology. Lowland forests, especially secondary or disturbed forest, sometimes in coastal or beach
forest, or kerangas forest. Sea-level to c. 800 m.
Uses. Often planted along roadsides as an ornamental and shade tree. The timber is hard and very
durable.
11. Fagraea gigantea Ridl.

(Latin, giganteus = very big; the tree)
J. Str. Br. R. As. Soc. 79 (1918) 98; Cockburn l.c. 211; Wong & Sugau l.c. 11. Lectotype (Wong & Sugau l.c.): Ridley 5818,
Singapore, Gardens Jungle (SING). Synonyms: F. speciosa sensu Ridley, J. Str. Br. R. As. Soc. 50 (1908) 122, non Blume
(1838); F. sororia J.J. Smith ex Cammerl., Bull. Jard. Bot. Btzg. 3, 5 (1923) 319, pl. 5; F. fragrans sensu Leenhouts, FM 1, 6
(1962) 304, Anderson l.c. 240, Ashton l.c. 310, pro parte, non Roxb. (1824).
Small to very large tree, to 45 m tall, 1.2 m diameter, trunk sometimes fluted, buttresses short, to 1 m
high. Bark deeply fissured, dark brown. Wood orange to pale yellow. Leaves chartaceous to
subcoriaceous, upper and lower side smooth; elliptic-obovate, 414 x 1.55.5 cm; base acute, margin
wavy, apex caudate; midrib prominent on lower side, lateral veins 38 pairs, prominent to faint on the
lower side, often depressed on the upper side, intercostals veins obscure; petioles 1.52.5 cm long,
base with axillary scales adnate to the twig and fused to form a nodal ochrea. Inflorescence axillary,
a many-flowered branched cyme, 2.55.5 cm long, 35 cm wide, main axis branching 23 orders;
peduncle 23.5 cm long. Flowers with pedicels 47 mm long; calyx 33.5 mm long, 1.52 mm
diameter; corolla salverform, tube 68 mm long, 1.5 mm wide, lobes elliptic, 45 mm long, 2.53 mm
wide; stamen filaments 1215 mm long, inserted just above the middle of the tube, anthers c. 1 mm
long; style exsert for 1416 mm, stigma capitate. Fruits globose, 35 mm across, yellowish, fruit
calyx 33.5 mm long, 33.5 mm wide. Seeds minute, angular, dark brown.
208
Vernacular names. Sabahtemasuk, temasuk hutan (Brunei Malay).

Distribution. Sumatra, Peninsular Malaysia and Borneo (all territories). In Sabah, documented for
the Keningau, Sandakan and Tawau districts; in Sarawak, known in the Kuching, Baram and Limbang
districts.
Ecology. Mixed dipterocarp forests, lowlands and hills up to 450 m.
Uses. The timber is hard and sometimes used for construction.
12. Fagraea kinabaluensis Wong & Sugau
Fig. 3.
(of Mt. Kinabalu)

Sandakania 8 (1996) 69. Type: Aban & Meijer SAN 93260, Sabah, Keningau, Crocker Range FR, mile 16 (holotype SAN;
isotype L). Synonym: F. ceilanica sensu Leenhouts, FM 1, 6 (1962) 315, Cockburn l.c. 211, pro parte, non Thunb. (1782).
Shrub or tree to 6 m tall, sometimes a climber or epiphyte, possibly also a strangler. Leaves thincoriaceous, upper and lower surfaces shagreen; narrowly elliptic to slightly obovate, 511 x 23.5 cm;
base acute to attenuate, margin slightly recurved when dry, apex acuminate to caudate; midrib
prominent on lower side, lateral veins obscure on both sides, intercostal veins obscure; petioles less
than 1 cm long, base with inconspicuous axillary scales. Inflorescence a solitary terminal flower.
Flowers with pedicels 23 mm long; bracteoles two pairs, small, 23 mm long; calyx cup-shaped,
2.22.6 cm long, 1.21.3 cm diameter, divided to its middle, lobes 1.71.8 cm long; corolla shortinfundibular, divided more than halfway down, tube 3-4 cm long, upper part 22.5 cm across, lobes
2.53 cm long, 1.52.5 cm wide. Fruits ellipsoid, mucronate, 33.5 cm long, 2.52.7 cm across; fruit
calyx lobes patent. Seeds rounded, brown.
Distribution. Endemic to Borneo (Sabah). In Sabah, documented only on the Crocker Range
(including Mt. Kinabalu) and its foothills, and Mt. Silam on the east coast.
Ecology. Montane forests.
13. Fagraea montana Wong & Sugau

(Latin, montanus = of mountains; the typical habitat)
Sandakania 8 (1996) 31. Type: Aban SAN 50722, Sabah, Ranau, copper mining area (holotype SAN, isotypes K, L).
Synonyms: F. minor sensu Merrill l.c. (1921) 492, Masamune l.c. 614, non Reinw. Ex
Blume (1826); F. racemosa sensu Leenhouts, FM 1, 6 (1962) 311, Cockburn l.c. 210, pro parte, non Jack ex Wall. (1824).
Tree to 40 m tall, 90 cm diameter, buttresses to 1.3 m high. Bark fissured to scaly, brown; inner bark
fibrous, laminated, yellow to reddish. Wood pale yellow. Leaves coriaceous, upper and lower surface
smooth; elliptic-obovate, 3.513.5 x 1.55.5 cm; base cuneate, margin plane to slightly recurved, apex
abruptly caudate; midrib prominent on lower side, lateral veins 45 pairs, faint to obscure on both
sides, intercostal veins obscure; petioles 0.72 cm long, base with axillary scales fused to form a
nodal ochrea. Inflorescence terminal, 4.514 cm long,
209
with clusters of condensed branches in distinct tiers along the main axis; peduncle 1.55.5 cm long.
Flowers with pedicels 612 mm long; calyx 44.5 mm long, 46.5 wide, lobes 2.53 mm long;
corolla funnel-shaped, 1820 mm long, narrowed basal part 78 mm long, expanded upper part 56
mm long, 67.5 mm wide, lobes ovate, 4.55 mm long, 33.5 mm wide; stamens inserted at the base of
the inflated part of the corolla, anthers 1 mm long, included or just visible at the throat; style exserted
for 45 mm, stigma capitate. Fruits ovoid-globose, 915 mm long, 811 mm wide; fruit calyx lobes
patent to clasping the fruit base. Seeds angular.
Vernacular name. Sarawakbira parak (Kelabit).
Distribution. Endemic to Borneo (Sabah and Sarawak). In Sabah, known only in the Crocker Range
(including Mt. Kinabalu) and its foothills; in Sarawak, documented for the Baram and Kapit districts.
Ecology. Montane forests, usually 10001600 m.
14. Fagraea philippinensis Wong & Sugau

Sandakania 8 (1996) 35. Type: Aherns Coll., For. Bur. 3270, Philippines, Luzon, Rizal (holotype K). Synonym: F.
racemosa sensu Leenhouts, FM 1, 6 (1962) 311, Anderson l.c. 240, Ashton l.c. 314 (as Form 4), pro parte, non Jack ex
Wall. (1824).
Small tree, 310 m tall, to 13 cm diameter. Bark fissured, grey. Leaves coriaceous, upper and lower
surfaces smooth; elliptic-oblong, 823 x 3.510.5 cm; base cuneate rounded or cordate, margin plane
to recurved, apex acuminate-short caudate; midrib prominent on lower side, lateral veins 48 pairs,
slightly prominent on the lower side, faint to obscure on the upper side, intercostal veins obscure;
petioles 0.61.5 cm long, the base with axillary scales forming a nodal ochrea. Inflorescence
terminal, 4.519 cm long, with clusters of condensed branches in distinct tiers along the main axis;
peduncle 2.58.5 cm long. Flowers with pedicels 26 mm long; calyx 79 mm long, 58 mm wide,
lobes 46 mm long; corolla salverform, 3446 mm long, narrowed basal part 712 mm long,
expanded upper part 1216 mm long, 2022 mm wide, lobes ovate, 1012 mm long, 910 mm wide;
stamen filaments inserted at the base of the inflated part, anthers included, 3.54 mm long; style
exsert for 57 mm. Fruits ellipsoid, 911 mm long, 79 mm wide; fruit calyx lobes clasping the fruit
base. Seeds angular.
Distribution. Philippines, Borneo (Sarawak and Kalimantan). In Sarawak, documented for the
Kuching and Limbang districts.
Ecology. Mixed dipterocarp forest and kerangas forest, to 600 m.
15. Fagraea resinosa Leenh.

(Latin, resinosus = resinous; the shoot tips and flower buds)
Bull. Jard. Bot. Brux. 32 (1962) 429; Wong & Sugau l.c. 86. Type: Hallier 3162, Amai Ambit (holotype L; isotype BO).
211
Tree to 25 m tall, 80 cm diameter, also recorded as an epiphyte or climber. Bark smooth, sometimes
covered with big lenticels, whitish brown; inner bark pale yellow. Leaves thick coriaceous, upper
surface slightly coarse, lower surface shagreen; broadly obovate, 1018 x 48 cm; base cuneate,
margin slightly recurved when dry, apex acute with short acumen; midrib prominent on the lower
side, lateral veins 78(11) pairs, faint to sunken on both sides, intercostal veins obscure; petioles 1
2.5 cm long, base with conspicuous axillary scales loosely clasping the node, auricles indistinct or
absent. Inflorescence terminal, with a solitary or a pair of flowers, rarely a sessile 3-flowered cyme.
Flowers with pedicels 2.53 cm long; bracteoles large, 23 pairs, with rounded apices, attached to
calyx cup base and forming an involucre; calyx 2.53 cm long, c. 1.5 cm diameter; corolla tubular,
white, fragrant, tube 89 cm long, contracted lower part c. 0.6 cm diameter, expanded upper part c. 2
cm wide, lobes ovate, c. 3 cm long, 2.5 cm wide; anthers broadly oblong, c. 6 mm long; stigma dishshaped, c. 2.5 mm diameter. Fruits ellipsoid, 5.56 cm long, 3.54 cm wide, fruit calyx 44.5 cm
long, 2.53 cm wide. Seeds ellipsoid-rounded.
Vernacular name. Sarawakterongau paya (Kelabit).
Distribution. Endemic to Borneo (Sabah, Sarawak, Brunei and Kalimantan). In Sabah, documented
only for the Ranau district; in Sarawak, recorded for the Kuching, Kapit, Miri and Limbang districts.
Ecology. Montane forest, up to 1500 m; also recorded at c. 1000 m on limestone on Mt. Api in
Sarawak.
16. Fagraea rugulosa Wong & Sugau
Fig. 4.
(Latin, rugulosus = wrinkled; the coarse leaf surface)

Sandakania 8 (1996) 22. Type: Chai & Illias S. 27929, Sarawak, 5th Div., Ulu Lawas, near Sg. Telau, Kota FR (holotype
SAN; isotypes A, BO, K, KEP, L, SING). Synonym: F. elliptica sensu Leenhouts FM 1, 6 (1962) 303, Ashton l.c. 314,
Anderson l.c. 239, pro parte, non Roxb. (1824).
Tree to 15 m tall, 25 cm diameter. Bark lightly fissured, dark brown; inner bark dark brown. Leaves
coriaceous, upper and lower surfaces very coarsely shagreen and rough; elliptic-obovate, 1121 x 5
9 cm; base cuneate, margin recurved when dry, apex cuspidate; midrib prominent on lower side,
lateral veins 912 pairs, faint to obscure on both sides, intercostal veins obscure; petioles 23.5 cm
long, base with axillary scales adnate to the twig and fused to form a nodal ochrea. Inflorescence
terminal, a many-flowered branched cyme, c. 6 cm long, c. 15 cm wide, main axis branching 34
orders, first branch 2.56 cm long; peduncle 12 cm long. Flowers with pedicels 23 mm long; calyx
campanulate, 34 mm long, 23.5 mm diameter, divided almost to its base; corolla salverform, tube
1011 mm long, 11.5 mm wide, lobes lanceolate, 58 mm long, 22.5 mm wide; stamen filaments
1518 mm long, inserted at corolla mouth, anthers c. 1 mm long; style exsert for 1011 mm; stigma
capitate. Fruits globose, (6)910 mm across; fruit calyx 23 mm long, 23 cm wide. Seeds angular,
c. 1 mm diameter.
Vernacular name. Sabahtamasuk jantan (Brunei Malay).
Distribution. Borneo: SW Sabah, Sarawak (Lawas and Baram area only) and Brunei.
Ecology. Mixed dipterocarp and kerangas forests, up to 450 m.
212
Fig. 4. Fagraea rugulosa. A, flowering leafy twig; B, detail of rough lower leaf surface; C, ultimate cyme-like unit of the
inflorescence, in bud; D, fruit. All from S. 27929, except D from S. 16603.)
213
17. Fagraea spicata Baker

(Latin, spicatus = spike-like; the inflorescence)
Kew Bull. (1896) 25; Merrill l.c. (1921) 493; Masamune l.c. 615; Wong & Sugau l.c. 38. Type: Creagh, s.n., British North
Borneo, East Coast (holotype K). Synonyms: F. congestiflora Elmer, Leafl. Philip. Bot. 8 (1915) 2741; F. racemosa sensu
Leenhouts, FM 1, 6 (1962) 311, Cockburn l.c. 210, Anderson l.c. 240, Ashton l.c. 314, pro parte, non Jack ex Wall. (1824).
Tree, to 17 m tall and 30 cm diameter or more. Bark smooth to fissured, brown. Leaves chartaceous
to thin coriaceous, upper and lower surfaces smooth; ovate-elliptic, 1030 x 514 cm; base cordate,
margin plane, apex acuminate-short caudate, midrib prominent on lower side, lateral veins 915 pairs,
prominent on lower side, faint to obscure on upper side, intercostal veins obscure; petioles indistinct,
0.20.5 cm long, base with axillary scales fused to form a nodal ochrea. Inflorescence terminal, 413
cm long, with clusters of condensed branches in distinct tiers along the main axis; peduncle 13 cm
long. Flowers with pedicels 27 mm long; calyx campanulate, 56 mm long, 3.55.5 mm wide, lobes
1.52.5 mm long; corolla infundibular, 3241 mm long, narrowed basal part 1112 mm long,
expanded upper part 2028 mm long, 1217 mm wide, lobes ovate, 5.57 mm long, 56.5 mm wide;
stamen filaments short, inserted at base of the inflated part, anthers included, 2.54 mm long; style 3
3.5 cm long, included or barely exsert, stigma capitate. Fruits ovoid-globose, 812 cm long, 810
mm wide, fruit calyx lobes patent. Seeds angular.
Vernacular names. Sabahtodopon puak (Dusun). Sarawaksira (Kelabit).
Distribution. Borneo (Sabah, Sarawak, Brunei and Kalimantan) and the Philippines. In Sabah,
documented only for the east coast districts, but probably more widespread; in Sarawak known for the
Bintulu, Kapit and Miri districts.
Ecology. Lowlands to montane forest (to c. 3200 m), frequently in gaps and also secondary forests.
18. Fagraea stenophylla Becc. ex Merr.

(Greek, stenos = narrow, phullon = leaf)
J. Str. Br. R. As. Soc. 77 (1917) 236, l.c. (1921) 493; Masamune l.c. 615; Wong & Sugau l.c. 38. Type: Native coll., Bur.
Sci. 2828, Sarawak, Upper Baram, Selongo (holotype PNH, destroyed; isotypes A, UC). Synonyms: F. eucalyptifolia
Cammerl., Bull. Jard. Bot. Btzg. 3, 5 (1923) 312, f. 2; F. racemosa sensu Leenhouts, FM 1, 6 (1962) 311, Cockburn l.c. 210,
Anderson l.c. 240, Ashton l.c. 314 (as Form 5), pro parte, non Jack ex Wall. (1824).
Small tree to 4 m tall. Bark smooth. Leaves coriaceous, upper and lower surfaces smooth; linear, 12
27 x 0.72.7 cm; base cuneate, margin recurved when dry, apex acuminate to caudate; midrib
prominent on lower side, lateral veins 714 pairs, faint to obscure on both sides, intercostal veins
obscure; petioles 1.32.2 cm long, base with axillary scales fused to form a nodal ochrea.
Inflorescence terminal, 3.515 long, with clusters of condensed branches in distinct tiers along the
main axis; peduncle 1.55.5 cm long. Flowers with pedicels 36 mm long; calyx campanulate, 56
mm long, 56 mm wide, lobes 34 mm long; corolla funnel-shaped, 2027 mm long, narrowed basal
part 810 mm long, expanded upper part 1011 mm
214
long, 78 mm wide, lobes semi-orbicular, 79 mm long, 4.55 mm wide; stamen filaments short,
inserted at the base of the inflated part of the corolla tube, anthers included, 22.5 mm long; style
included, stigma capitate. Fruits ellipsoid, 1014 mm long, 69 mm wide, fruit calyx lobes clasping
the fruit base to patent. Seeds angular.
Distribution. Endemic to Borneo (Sarawak, Kalimantan, Sabah and Brunei). In Sabah, documented
for the Sipitang, Penampang and Keningau districts in the SW; in Sarawak, recorded for the Kapit,
Miri and Limbang districts.
Ecology. A rheophyte, on banks of swift-running streams in the lowlands to 180 m.
19. Fagraea teysmannii Cammerl.

(J.E. Teysmann, 18081882, Curator of the Bogor Botanic Gardens)
Bull. Jard. Bot. Btzg. 3, 5 (1923) 314. f. 3; Wong & Sugau l.c. 40. Lectotype (Wong & Sugau l.c.): Teysmann, s.n.,
Karimata, Soengei Tajan (L). Synonym: F. racemosa sensu Leenhouts, FM 1, 6 (1962) 311, Anderson l.c. 240, Ashton l.c.
314 (as Form 5), pro parte, non Jack ex Wall. (1824).
Tree, to 10 m tall, c. 20 cm diamater. Bark smooth, brown. Leaves coriaceous, upper and lower
surfaces smooth; elliptic, oblong, 4.516 x 1.56 cm; base cuneate to rounded, margin recurved when
dry, apex acuminate; midrib prominent to flattened on the lower side, lateral veins 47 pairs, faint to
obscure on both sides, intercostal veins obscure; petioles 0.61.7 cm long, base with axillary scales
adnate to the twig and fused to form a nodal ochrea. Inflorescence terminal, 210 cm long, with
clusters of condensed branches in distinct tiers along the main axis; peduncle 15 cm long. Flowers
with pedicels 27 mm long; calyx 78 mm long, 56 mm wide, lobes 34 mm long; corolla
infundibular, 2639 mm long, narrowed basal part 67 mm long, expanded upper part 1314 mm
long, 1012 mm wide, lobes semi-orbicular, 78 mm long, 56 mm wide; stamen filaments short,
inserted at the base of the inflated part of the corolla tube, anthers included, c. 1.5 mm long; style
included or barely exsert, 2.53 cm long, stigma capitate. Fruits ovoid-ellipsoid, 78 mm long, 57
mm wide, fruit calyx clasping the fruit base. Seeds angular.
Vernacular names. Sarawakkapan, uchip bali (Kenyah).
Distribution. Endemic to Borneo (Sabah, Sarawak and Kalimantan). In Sabah, documented for the
Tambunan district, in Sarawak for the Kapit district.
Ecology. In montane forests, to 1250 m.
20. Fagraea volubilis Wall.
Fig. 5
(Latin, volubilis = twining; a habit wrongly attributed to this species)

In Roxb., Fl. Ind. 2 (1824) 36; Wong & Sugau l.c. 40. Type: Jack, s.n., E Bencoolen (holotype K, sheet marked 1600 E.
Bencoolen on bottom left).
Tree to 14 m tall, 20 cm diameter or more. Bark fissured or cracking, dark brown; inner bark pale
brown. Sapwood yellowish. Leaves coriaceous, upper and lower surfaces smooth; ovate, elliptic to
oblong, 924 x 415 cm; base cuneate, rounded to subcordate, margin plane, apex
215
Fig. 5. Fagraea volubilis var. microcalyx. A, flowering leafy twig; B, detail of part of inflorescence; C, fruit. (All from S.
35683.)
216
acuminate to short caudate; midrib prominent on lower side, lateral veins 511 pairs, prominent on the
lower side, faint to obscure on the upper side, intercostal veins obscure; petioles 12 cm long, stout,
base with axillary scales fused to form a nodal ochrea. Inflorescence terminal, 230 cm long, with
clusters of branches in distinct well-spaced tiers along the main axis, these branches typically
condensed but rarely elongate; peduncle 2.517 cm long. Flowers with pedicels 313 mm long; calyx
46 mm long, 3.56 mm wide, lobes 1.53.5 mm long; corolla broadly infundibular, 2634 mm long,
narrowed basal part 1012 mm long, expanded upper part 810 mm long, 1315 mm wide, lobes
semi-orbicular to ovate, 712 mm long, 7.59 mm wide; stamen filaments inserted at the base of the
inflated part of the corolla tube, anthers included, 2.53 mm long; style barely exsert for 23 mm,
stigma capitate. Fruits ovoid to ellipsoid, 1116 mm long, 812 mm wide, tipped with persistent style
base; fruit calyx lobes patent. Seeds tiny, angular, black.
Key to varieties
Calyx lobes 33.5 mm long..........................................................................................................
var. volubilis
Synonyms: F. morindaefolia Blume, Rumphia 2 (1838) 32, t. 73 f. 2, t. 79; F. coarctata
Blume l.c. (1838) 33, Merrill l.c. (1921) 491, Masamune l.c. 613; F. scholaris Blanco, Fl.
Filip. ed. 2 (1845) 93, ed. 3, 1 (1877) 171; F. appendiculata Blume, Mus. Bot. Lugd. Bat. 1
(1850) 169; F. subreticulata Blume l.c. (1850) 171; F. latifolia Miq., Fl. Ind. Bat. 2 (1857)
369; F. rodatzii Laut. & Schum., Fl. Schutzgeb. (1900) 499; F. grandifolia Merr., J. Str. Br.
R. As. Soc. 77 (1917) 231, l.c. (1921) 492, Masamune l.c. 614; F. racemosa var. pauciflora
King & Gamble, J. As. Soc. Beng. 74, 2 (1908) 609; F. pauciflora (King & Gamble) Ridl.,
FMP 2 (1923) 419, f. 110; F. racemosa sensu Leenhouts, FM 1,6 (1962) 311, Cockburn l.c.
210, Anderson l.c. 240, Ashton l.c. 314 (as Form 5), pro parte, non Jack ex Wall. (1824).
The Andaman and Nicobar islands, Sumatra, Peninsular Malaysia, Java, Borneo (Sabah,
Sarawak, and Kalimantan), the Philippines, Sulawesi, Maluku and New Guinea. In Sabah,
known from the Sipitang, Beaufort, Papar, Tenom, Keningau, Kota Belud, Labuk Sugut and
Sandakan districts; in Sarawak, from the Kuching and Miri districts. Lowland rain forest,
including secondary forest and forest fringes, up to 1100 m, also in kerangas and peat swamp.
Calyx lobes 1.52.5 mm long.......................................................................................................

var microcalyx Wong & Sugau
l.c. 43. Synonyms: F. ligustrina Blume, Rumphia 2 (1838) 33 (including var. disparifolia
Blume), Merrill l.c. (1921) 492, Masamune l.c. 614; F. cordifolia Blume, l.c. (1838) 33,
Merrill l.c. (1921) 491, Masamune l.c. 613; F. gracilis Cammerl., Bull. Jard. Bot. Btzg. 3, 5
(1923) 316; F. racemosa sensu Leenhouts, FM 1, 6 (1962) 311, Anderson l.c. 240, Ashton l.c.
314 (as Form 3), pro parte, non Jack ex Wall. (1824).
Borneo (Sarawak, Brunei and Kalimantan) and Maluku. In Sarawak, documented for the
Kuching, Sibu and Bintulu districts. Lowlands; peat swamps, mixed dipterocarp forest and
kerangas forest.
Vernacular names. Sabahtodopon puok (Dusun). Sarawaksokong ranyai (Iban), tembusu
(Malay).
217
2. GENIOSTOMA J.R. Forst. & G. Forst.

(Greek, geneion = bearded, stoma = mouth; the hairy corolla throat)
Char. Gen. Pl. 12 (1776) t. 12; Valeton, Bull. Inst. Bot. Btzg. 12 (1902) 1; Leenhouts, FM 1, 6 (1962) 369; A.C. Smith &
Stone, Contrib. U.S. Nat. Herb. 37 (1962) 1; Conn, Blumea 26 (1980) 245.
Shrubs or treelets. Leaves pinnately nerved; petioles distinct or very short, the base expanded to form
a distinct cup-like ochrea. Inflorescence axillary, sometimes at leafless nodes, a cyme or sometimes a
solitary flower; minute bracteoles often present. Flowers (4-)5-merous, gynodioecious (with female
and bisexual flowers together); sepals fused at the very base, margin ciliate; corolla campanulate to
rotate, greenish white, lobes imbricate or contorted in bud, outside glabrous or short-hairy, inside
glabrous or densely woolly, especially at the throat; stamens inserted in the throat, exsert, filaments
usually short, anthers 2-locular; ovary 2-locular, ovules many, style often very short, stigma clubshaped or ellipsoid to globular, about as large as the ovary. Fruit a capsule, splitting into 2 parts.
Seeds numerous, ellipsoid to subglobular, intruded on the hilar side, minutely warty, embedded in a
juicy pulp.
Distribution. About 2040 species, mainly in the Pacific region, extending to S Japan (Kyushu) and
to Australia, Lord Howe Island, New Zealand, New Caledonia and the Society Islands in the east.
Four species occur in Malesia, 3 of which are restricted to New Guinea; 1 species in Sabah.
Geniostoma rupestre J.R. Forst. & G. Forst.
Fig. 6.
(Latin, rupestris = rock-dwelling)

Char. Gen. Pl. 12 (1776) t. 12; Valeton l.c. 12, 17, f. 1. Lectotype (A. C. Smith & Stone l.c.): J. R. & G. Forster 30, s. dat.,
Tanna, New Herbrides (BM; J. R. & G. Forster, s.n., s. dat., Tanna, New Herbrides at K is believed to be the isolectotype).
Synonyms: G. micranthum DC., Prod. 9 (1845) 27; G. montanum Zoll. & Moritzi in Moritzi, Syst. Verz. (1846) 58; G.
lasiostemon Blume, Mus. Bot. Lugd. Bat. 1 (1850) 239; G. cumingianum Benth., J. Linn. Soc. Bot. 1 (1856) 97, Merr., En.
Philip. 3 (1923) 310; G. avene Valeton, Bull. Inst. Bot. Btzg. 12 (1902) 23 & 16; G. celebicum Valeton l.c. 19 & 15, f. 7 &
13; G. miquelianum Koord. & Valeton ex Valeton l.c. 22 & 14, f. 11, 12 & 16; G. moluccanum Valeton, l.c. (1902) 19 & 15,
f. 4, Cammerl., Bull. Jard. Bot. Btzg III, 5 (1923) 297; G. oblongifolium Koord. & Valeton ex Valeton l.c. (1902) 20 & 15, f.
5 6 & 14; G. philippinensis Merr., Philip. J. Sc. 3 (1908) Bot. 259; G. batanense Merr., ibid. 3 (1909) Bot. 427, l.c. (1923)
309; G. stenophyllum Merr., Philip. J. Sc. 7 (1912) Bot. 329, non Gilg. & Bened. (1916); G. brevipes Merr., Philip. J. Sc. 9
(1914) Bot. 384; G. laxa Elm., Leafl. Philip. Bot. 8 (1915) 2746; G. mindanaense Elm., Leafl. Philip. Bot. 8 (1915) 2747; G.
longipes Merr., Philip. J. Sc. 12 (1917) Bot. 296; G. pachyphyllum Merr., Philip. J. Sc. 14 (1919) 448; G. lancilimbum Merr.,
Philip. J. Sc. 17 (1921) 304; G. ramosii Merr., Philip. J. Sc. 17 (1921) 304; G. fasciculata Quis. & Merr., Philip. J. Sc. 37
(1928) 190.
Shrub or small tree, to 4(10) m tall. Branches terete, rarely quadrangular, glabrous or the young
twigs shortly yellowish brown hairy. Leaves chartaceous to thin-coriaceous, glabrous, ovate to
elliptic, 46 x 1.52.5 cm; base cuneate, apex gradually tapering to acuminate, margins strongly
recurved when dry; midrib prominent, lateral veins 67 pairs, intercostal veins loosely reticulate and
flat to slightly prominent on both surfaces; petioles 0.51.5 cm long, the expanded bases forming a
distinct cup-like ochrea 12 mm high around the node. Inflorescence several-flowered, 0.51.5 cm
long, glabrous; bracteoles small. Flowers with
218
Fig. 6. Geniostoma rupestre. A, flowering leafy twigs; B, detail of node, showing cup-shaped ochrea; C, flower. (All from
RSNB 4708.)
219
pedicels 15 mm long; calyx 0.52 mm long, lobes ovate to triangular, acute, outside glabrous,
margin ciliate; corolla white, 1.54.5 mm long, outside glabrous or rarely fairly densely papillosehairy, inside densely to sparsely hairy at the throat, lobes 0.52.5 mm long, blunt to acute; stamens
with filaments 0.51 mm long, glabrous or hairy, anthers 0.71.2 mm long; ovary glabrous to densely
shortly hairy, style very short (less than 0.2 mm). Fruits somewhat flattened-globose, rarely ellipsoid,
ovoid, or obovoid 48(12) x 46(9) mm, black when ripe. Seeds ellipsoid, dark brown.
Distribution. Widely distributed in Malesia (except in Peninsular Malaysia), E Queensland and the W
Pacific. In Borneo, so far recorded only from Sabah, at about 2000 m on Mt. Kinabalu.
Ecology. The species has a wide ecological amplitude, recorded from sea-level to 2800 m elevation,
in both forest as well as exposed sites. It is, however, apparently uncommon in Borneo.
3. NORRISIA Gardner
(William Norris, 17931859, a British judge in colonial SE Asia)
In Hooker, J. Bot. Kew Misc. 1 (1849) 326; Merrill, EB (1921) 490; Masamune, EPB (1942) 616; Leenhouts, FM 1, 6
(1962) 293; Cockburn, TS 1 (1976) 207; Anderson, CLTS (1980) 240; Ashton, MNDTS 2 (1988) 317.
Trees. Twigs often terete, covered with brown hairs when young. Leaves elliptic to oblong, base
attenuate to cuneate, apex rounded to acute or acuminate; pinnately veined; petiole bases connected by
a ridge-like thickening or forming a low inconspicuous ochrea not more than 1 mm high.
Inflorescence terminal, densely brown-hairy, a dichasially-branched to compound cyme, manyflowered; bracts narrowly triangular. Flowers fragrant, subsessile, with two pairs of decussate
bracteoles at the base; calyx cupular, tube very short, lobes broadly rounded, densely hairy outside,
glabrous inside; corolla salver-shaped, creamy to yellowish, densely hairy outside, lobes valvate in
bud; stamens inserted between the corolla lobes; ovary obovoid, densely short-hairy, 2-locular, ovules
many, style terete, stigma knob-like and slightly bilobed. Fruits capsular, splitting down the septa
into 2 parts. Seeds few to many, small, spindle-shaped, glabrous, smooth; endosperm fleshy.
Distribution. Two closely related species, distributed from Sumatra, Peninsular Malaysia,
Borneo to the Philippines. Both species in Sabah and Sarawak.
Key to Norrisia species

Leaf lateral veins not looping except for the distal-most 23 pairs, which arch and join close
to the margin; midrib on upper side distinctly short-hairy (x10 magnification). Mouth of the
corolla villous-hairy. Anthers suborbicular, c. 0.5 mm across. Fruits larger, 410 mm
long............................................................................................................................1. N. maior
220
Leaf lateral veins distinctly arched and joined at some distance from the margin; midrib on
upper side glabrous. Mouth of the corolla only sparsely hairy to glabrous. Anthers oblong, c.
0.8 mm long. Fruits smaller, 2-3.5 mm long........................................................ 2. N. malaccensis
1. Norrisia maior Soler.
(Latin, maior = bigger; the fruit compared with N. malaccensis)
In Engl. & Prantl, Nat. Pfl. Fam. 4, 2 (1892) 37; Cockburn I.e. 212; Anderson I.e. 240; Ashton I.e. 317. Type: s.
coll., s.n., Malacca (K). Synonyms: N. malaccensis sensu Merr., I.e. (1921) 490, non Gardner (1849); N.
malaccensis var. major Ridl., FMP 2 (1923) 414.
Tree to 50 m tall, 75 cm diameter; buttresses to 3.2 m high. Bark smooth to slightly fissured,
dark brown; inner bark pale brown. Sapwood pale yellow to white. Leaves thin-coriaceous,
elliptic, 2.5-4.5 x2.5-9.5 cm; base attenuate to cuneate, apex acuminate, acute or rounded;
midrib prominent and glabrous to hairy on lower side, flat to slightly channelled and distinctly
short-hairy on upper side, lateral veins 6-10 pairs, the distalmost 23 pairs arching and
joining close to the margin, intercostal veins subreticulate; petioles 0.3-0.5 cm long.
Inflorescence 5.5-10.5 cm long, peduncle 45 cm long; bracts to 2 mm long. Flowers
yellowish green, calyx 0.5-1 cm high, divided to halfway down, densely hairy; corolla 6-8 mm
long, inside villous-hairy at the mouth, outside minutely hairy, lobes 1-2 mm long, inside
glabrous; stamens 4-5 mm long, anthers suborbicular, c. 0.5 mm diameter, ovary c. 1.5 mm
long; style 7-8 mm long. Fruits obovoid, 4-10 x 2-4 mm. Seeds c. 5-20 per locule, c. 4 mm
long.
Vernacular names. Sabahsimpapait (Putatan Dusun). Sarawakbannang (Padawan
Bidayuh), belet (Kenyah), bi'is (Bau Bidayuh), empaling (Iban), gunnong (Sadong Bidayuh),
mepa (Berawan, Tutoh Punan), nyvang (Kayan).
Distribution. Sumatra (Palembang), Banka, Riouw, Peninsular Malaysia and Borneo (Sabah,
Sarawak, Brunei and Kalimantan). In Sabah, recorded for all W coast districts and the Kota
Belud, Keningau, Telupid, Sandakan and Kinabatangan districts. In Sarawak, common
throughout.
Ecology. Primary and secondary forest, often along river banks, also in swamp forest, to 450 m.
Flowering recorded in January and June to August, fruiting in February and September.
Uses. The timber is locally used for construction in Peninsular Malaysia.
2. Norrisia malaccensis Gardner
Fig. 7.
(of Malacca)
In Hooker, J. Bot. Kew Misc. 1 (1849) 327. Type: Griffith, s.n., Malacca (K). Synonyms: Antonia grijfithii
Wight, 111. Ind. Bot. 2 (1858) 172, ;. 156b; N. philippinensis Elm. Leafl. Philip. Bot. 4 (1912) 1482.
Tree to 30 m tall, 50 cm diameter. Bark smooth, dark grey; inner bark pale brown. Sapwood
yellowish. Leaves thin-coriaceous, elliptic to oblong, 4-6 x 1.8-2.6 cm; base attenuate to
cuneate, apex acute to acuminate; midrib prominent and glabrous to sparsely hairy on lower
221
side, flat to slightly channelled and glabrous on upper side, lateral veins 6-8 pairs, distinctly
arching and joining at some distance from the margin, intercostal veins reticulate; petioles
slender, 0.4-1 cm long. Inflorescence 4-1.5 cm long; peduncle 1.8-2 cm long; bracts to 2.5 mm
long. Flowers pale yellow; calyx 0.5-1 mm long, divided nearly to the base, densely short-hairy;
corolla 2-A mm long, inside sparsely hairy to glabrous, outside usually sparsely hairy, lobes 1.52 mm long, inside glabrous; stamens 4-5 mm long, anthers oblong, c. 0.8 mm long; ovary c. 1
mm long, style 6-7.5 mm long. Fruits obovoid, 2-3.5 x c. 1.9 mm. Seeds few, c. 2 mm long.
Distribution. Sumatra (W coast), Peninsular Malaysia, the Philippines (Sibuyan and
Mindanao) and Borneo (Sabah only, here newly recorded with SAN 92976 and SAN 131857
from Bukit Tangkunan in Beluran).
Ecology. Recorded in mixed dipterocarp forest at about 400 m.
223
224
__________________________________________________________________________________________
LYTHRACEAE
P.C. Yii
Kuching, Malaysia
Hooker f., Fl. Br. Ind. 2 (1879) 565; King, J. As. Soc. Beng. 67, 1 (1898) 345; Merrill, EB (1921) 417; Ridley, FMP 1 (1922)
819; Masamune, EPB (1942) 512; Backer & Bakhuizen f., FJ 1 (1963) 251; Keng, OFMSP (1969) 153; Everett & Whitmore,
TFM 2 (1973) 276; Shea in Cockburn, TS 1 (1976) 212; Anderson, CLTS (1980) 240; Corner, WSTM 3rd ed. 1 (1988) 470;
Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 211.
Herbs, shrubs or trees, occasionally scrambling shrubs or vines (not in Borneo). Young twigs usually
quadrangular, gradually becoming terete, smooth or sometimes spiny. Leaves simple, entire, mostly
opposite-decussate and crowded at the ends of twigs; stipules minute or absent. Inflorescences
axillary or terminal racemes or panicles. Flowers bisexual, radially or rarely bilaterally symmetrical;
calyx bell-shaped or tubular, smooth or ridged, free from ovary, persistent in fruit, calyx lobes 36,
valvate; petals as many as calyx lobes, crinkled and usually caducous or absent (Ammannia, Peplis &
Rotala, not in Borneo); stamens twice as many as petals or numerous, inserted on the inside of
receptacle, filaments equal or unequal, anthers 2-locular; ovary superior or semi-superior, 26locular, style one, stigma capitate; ovules numerous. Fruit a loculicidal capsule, adnate to the base of
the persistent calyx. Seeds numerous, straight and small, with or without wings; endosperm absent.
Distribution. About 25 genera and 550 species, chiefly tropical. In Sabah and Sarawak, represented
by 3 genera and 7 species.
Taxonomy. The Lythraceae are closely related to the Crypteroniaceae and Sonneratiaceae. The
Crypteroniaceae are segregated from the Lythraceae in their tiny apetalous flowers with a 23carpellate ovary. The Sonneratiaceae are distinct in their partially united carpels containing numerous
ovules that are inserted on the septa.
Key to genera
1. Stems and branches usually twisted. Leaves fleshy, with indistinct veins. Flowers
axillary................................................................................................................2. Pemphis
Stems and branches not so. Leaves coriaceous, with distinct veins. Flowers in terminal
panicles................................................................................................................................2
2. Flower 4-merous, stamens usually 8 or fewer. Fruit an indehiscent capsule. Seeds not
winged...................................................................................................................................
225
Lawsonia L.
Sp. Pl. (1753) 349; Hooker f. l.c. 573; Merrill l.c. 418; Masamune l.c. 512; Backer &
Bakhuizen f. l.c. 256; Corner l.c. 475.
A monotypic genus (L. inermis) , distributed from the east coast of Africa to SW Asia and
widely cultivated throughout tropical SE Asia.
Shrubs or treelets. Twigs usually spiny or rarely without spines. Leaves short-stalked. Flowers
white, intensely fragrant, in axillary and terminal leafy panicles; calyx obconical, deeply
lobed, lobes 4; petals 4; stamens 8, rarely 4, alternate with petals; ovary 4-carpellate, styles
filiform, stigmas small and capitate; ovules numerous. Fruit an indehiscent capsule, globose
or depressed globose, subtended by a shallow persistent calyx tube.
In Sabah and Sarawak, the species is widely cultivated for its red dye.
Flower 6-merous or more, stamens numerous. Fruit a dehiscent capsule. Seeds winged......
..................................................................................................................1. Lagerstroemia
1. LAGERSTROEMIA L.
(M. Lagerstroem, 16911759, Swedish patron of science)
bungor (Malay)
Arn. Acad. 4 (1759) 137; Hooker f., Fl. Br. Ind. 2 (1879) 575; King, J. As. Soc. Beng. 67, 1 (1898) 350; Merrill, EB (1921)
417; Ridley, FMP 1 (1922) 822; Masamune, EPB (1942) 512; Backer & Bakhuizen f., FJ 1 (1963) 255; Furtado & Montien,
Gard. Bull. Sing. 24 (1969) 186; Everett & Whitmore, TFM 2 (1963) 277; Shea in Cockburn, TS 1 (1976) 214; Anderson,
CLTS (1980) 240; Corner, WSTM 3rd ed. 1 (1988) 472; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 211.
Deciduous or semi-deciduous trees or shrubs; crown usually bushy and spreading; bole often fluted;
buttresses small or absent. Bark smooth, papery, grey to light fawn-brown mottled, flaking; inner
bark fibrous or laminated, grey fawn, staining purple. Sapwood white to pale yellowish brown.
Axillary buds conspicuous, elongated and pointed. Leaves opposite, entire, coriaceous, often
withering orange or red before falling. Inflorescence a large panicle, terminal or in the axils of upper
leaves. Flowers large, showy, pink-mauve, fading white, scentless; calyx funnel- or bell-shaped,
coriaceous, smooth or ridged, lobes acute or acuminate, valvate; petals 6 or more, delicate, crinkled;
stamens numerous, in few rows, filaments slender, anthers minute; ovary globose or ellipsoid, fused
to the base of calyx cup. Fruit a large woody capsule, splitting longitudinally into 6 parts at the
apical end. Seeds numerous, elongated, with apical wing.
Distribution. About 55 species, chiefly in tropical and subtropical parts of Madagascar, Asia, New
Guinea and Australia. In Borneo, represented by 6 species, of which only 3 species are native to
Sabah and Sarawak. Lagerstroemia borneensis occurs only in Kalimantan, while L. indica and L.
floridunda have been introduced as ornamental plants.
Ecology. Lagerstroemia species prefer seasonally dry parts of continental SE Asia; in Sabah and
Sarawak, they occur from coastal forest within reach of sea-spray to lowland forest farther inland.
226
LYTHRACEAE (YII)
Uses. Rarely reaching timber size and, therefore, only occasionally used for domestic purposes. Three
species, namely L. floribunda, L. indica and L. speciosa are widely cultivated as ornamentals
throughout the country.
Key to Lagerstroemia species

1. Shrub. Leaves sessile or almost so.......................................................................................
L. indica L.
l.c. 137; Hooker f. l.c. 575; Backer & Bakhuizen f. l.c. 256; Furtado & Montien l.c. 190;
Corner l.c. 474.
Shrubs. Leaves chartaceous, obovate or ovate, 310 x 24 cm. Inflorescences to 20 cm long,
720 cm across. Flower buds subglobose, 56 mm across, shortly tipped at the apex, ridges
56, superficial and soon disappearing in the upper half; calyx 46-lobed, lobes erect,
triangular; petals suborbicular, wavy and curled at the margins; stamens numerous, only 46
are stouter and longer, others unequal; ovary subglobose, glabrous, with a long slender style.
Capsules globose, c. 10 x8 mm, 46-valved; persistent calyx funnel-shaped or cup-shaped.
Native to the Himalayan region, China, Indo-China and Japan; widely cultivated in parks and
roadsides throughout Sabah and Sarawak.
Tree. Leaves distinctly stalked............................................................................................2

2. Lower leaf surface glabrous................................................................................................3
Lower leaf surface hairy on the veins..................................................................................4
3. Leaves and calyces often pustulate. Persistent calyx lobes in fruit spreading.......................
......................................................................................................................2. L. pustulata
Leaves and calyces not so. Persistent calyx lobes in fruit recurved.................3. L.
speciosa
4. Upper leaf surface glabrous, lower leaf surface minutely hairy. Inflorescences minutely
velvety hairy................................................................................................1. L. piriformis
Both leaf surfaces and inflorescences rusty stellate-tomentose.............................................
L. floribunda Jack
Mal. Misc. 1 (182022) 38; Hooker f. l.c. 577; King l.c. 345; Ridley l.c. 823; Backer &
Bakhuizen f. l.c. 256; Furtado & Montien l.c. 329; Everett & Whitmore l.c. 279; Shea in
Cockburn l.c. 216; Corner l.c. 474.
Tree to 13 m tall. Leaves oblong-elliptic, 623 x 410 cm; young leaves with stellate hairs
above and along veins beneath, late glabrescent; base broadly cuneate or somewhat rounded,
apex obtuse or rarely acute; petioles 37 mm long. Inflorescences 2040 cm long, rusty
stellate-tomentose. Flower buds turbinate, 1012-ridged; calyx, 6-lobed; petals oblong,
gradually cuneate towards the base, margin wavy; stamens numerous, unequal; ovary
subglobose, densely brown-tomentose. Capsules ellipsoid- oblong, 1216 x 711 mm,
sparsely tomentose all over; persistent calyx cup-shaped, lobes oblique or recurved.
Native to subtropical and tropical SE Asia, from S China to Mynmar, Thailand, Cambodia,
Indo-China and Peninsular Malaysia. In Sabah and Sarawak, widely cultivated as ornamental
trees.
227
1. Lagerstroemia piriformis Koehne

(Latin, piriformis = pear-shaped; the fruit)
In Engler, Bot. Jahrb. 4 (1883) 23; Furtado & Montien l.c. 248; Shea in Cockburn l.c. 216. Type: Cuming 1675, Philippines
(n.v.). Synonym: L. crassifolia Furtado & Montien l.c. 232.
Tree to 15 m tall, 30 cm diameter. Leaves coriaceous, greyish green and glabrous above, brownish
green and minutely hairy along the veins beneath; elliptic or ovate, 58 x 34.2 cm; base asymmetric
or rarely symmetric and cuneate, shortly decurrent, apex acute or acuminate; lateral veins 68 pairs;
petioles 36 mm long. Inflorescences 720 cm long, 715 cm across, minutely velvety hairy. Flower
buds 912 x 5 mm, obscurely apiculate at the apex, ridges 6, superficial, straight in the developing
buds, slightly angular in the lower parts, slightly thickened along the suture at the top, finely
furrowed; calyx c. 8 x 5 mm, lobes c. 3 mm long; petals obovate, c. 6 x 4 mm, margins frilly or wavy;
stamens numerous, only 6 thicker and longer ones with filaments; ovary ovoid, glabrous, style c. 1.5
mm long. Fruits broadly ellipsoid, c. 15 x 11 mm, conspicuously beaked; persistent calyx c. 10 x 10
mm, abruptly narrowed at the base, slightly ridged at the lower half of the tube, lobes c. 4 x 3 mm,
recurved.
Distribution. Borneo, Philippines, New Guinea and Melanesia. In Borneo, known from Sabah from
two collections (SAN 47262 and SAN 51859) from primary forest at Gomantong, Sungei Lambah,
Sandakan district and from Lamag, Kinabatangan district, respectively.
Ecology. Low-lying primary forest on limestone or black volcanic soils.
2. Lagerstroemia pustulata Furtado & Montien
Fig. 1.
(Latin, pustulatus = blistered; the leaf surface)

l.c. 222; Shea in Cockburn l.c. 216; Whitmore, Tantra & Sutisna l.c. 212. Type: Muin Chai SAN 26006, Borneo, Sabah,
Lahad Datu, Segam Sarai (holotype SING; isotype SAN, SAR).
Tree to 17 m tall, 60 cm diameter. Leaves oblong-elliptic 1017 x46 cm; base cuneate, margin
slightly recurved, apex acuminate; glabrous with distinct pustules on both surfaces; lateral veins 10
12 pairs; petioles 510 mm long. Inflorescences to 32 cm long, greyish hairy. Flowers: calyx
pustulate and hairy, 1012 mm deep (including 46 mm obconical base), ribs 6, distinct, lobes c. 5 x 2
mm, distinctly recurved; petals 6, suborbicular or obovate, c. 15 mm long (including 5-mm claw);
stamens numerous, erect, inserted at the bottom of the calyx tube; ovary glabrous, ovoid, gradually
narrowed into a conical apex, style long. Fruits oblong, c. 25 x 2022 mm, abruptly apiculate at the
apex; persistent calyx hard, abruptly narrowed into 812 mm long obconical base, c. 5 mm deep,
lobes spreading, slightly thicker at the margins.
Vernacular name. Sabahperlimpong hutan (Dusun).
Distribution. Only known from the type collection from Segam Sarai, Lahad Datu district, Sabah.
Ecology. Lowland forest along river at c. 70 m.
228
LYTHRACEAE (YII)
Fig. 1. Lagerstroemia pustulata. A, flowering leafy twig; B, flower with petals removed; C, petal; D, longitudinal section of
flower; E, stamen; F, mature fruit. (All from SAN 26006.)
229
3. Lagerstroemia speciosa (L.) Pers.

(Latin, speciosus = showy; the flowers)
Synops. 2 (1807) 72; Merrill l.c. 417; Masamune l.c. 512; Furtado & Montien l.c. 264; Everett & Whitmore l.c. 280; Shea in
Cockburn l.c. 216; Anderson l.c. 240; Corner l.c. 474; Whitmore, Tantra & Sutisna l.c. 212. Basionym: Munchausia
speciosa L. in Munchausen, Hausvater 5, 1 (1770) 357, t. 2. Type: Katou-Adamboe Rheede, Hort. Mal. 4 (1683) 45, t. 22,
p.p. Synonym: L. flos-reginae Retz., Obs. 5 (1789) 25, Hooker f. l.c. 577, King l.c. 352, Ridley l.c. 823, Corner, Gard. Bull.
Str. Settl. 10 (1939) 272, Backer & Bakhuizen f. l.c. 256, Whitmore, Tantra & Sutisna l.c. 212.
Tree to 10 m tall, 30 cm diameter. Leaves elliptic-oblong, 519 x 48 cm; base cuneate or almost
rounded, apex acute or obtuse; glabrous, greyish green above and brown beneath; lateral veins 815
pairs, distinctly looped and joined to the next pair; petioles 49 mm long. Inflorescences 1540 cm
long, 1020 cm across, covered with caducous, ashy or rusty hairs; pedicels c. 1.5 mm long. Flower
buds subglobose or pyriform, 1015 x 610 mm, shortly apiculate at the apex, 1214-ridged, ridges of
the same length; calyx ashy or rusty hairy, lobes recurved; petals 6, suborbicular, 1530 x 1020 mm,
tapering towards the slender claw; stamens numerous, subequal; ovary globose, glabrous or slightly
scaly, style filiform, stigma capitate. Fruits woody, globose, glabrous, c. 22 mm across; persistent
calyx glabrous or slightly hairy, lobes recurved.
Vernacular names. Sabahtibabah (Dusun Banggi). Sarawakbongor biru (Malay).
Distribution. China, Indo-China, Peninsular Malaysia, Sumatra, Borneo and the Philippines. Some
doubt has been expressed as to whether L. speciosa is native to Sabah and Sarawak, since most of the
collections are from towns or forest close to habitation. However, they are common in some places
beside large rivers, such as the Kinabatangan in Sabah, and appear to grow wild there.
Ecology. Lowland secondary forest on sandy soil.
2. PEMPHIS J. R. Forst. & G. Forst.

(Greek word for bladder; the shape of the fruit stalk)
Char. Gen. (1776) 68, t. 34; Hooker f., Fl. Br. Ind. 2 (1879) 572; King, J. As. Soc. Beng. 67, 1 (1898) 348; Merrill, EB
(1921) 418; Ridley, FMP 1 (1922) 821; Masamune, EPB (1942) 512; Backer & Bakhuizen f., FJ 1 (1963) 255; Burkill,
EPMP (1966) 1716; Everett & Whitmore, TFM 2 (1963) 276; Shea in Cockburn, TS 1 (1976) 214; Anderson, CLTS (1980)
240; Corner, WSTM 3rd ed. 1 (1988) 475.
Shrubs or rarely trees; stems and branches usually twisted. Leaves decussate, succulent, subsessile.
Flowers axillary, solitary or rarely in pairs, short-stalked, radially symmetrical, 6-merous; calyx bellshaped or tubular with broadly triangular lobes; stamens 1218, with shorter and longer ones
alternating, longer ones opposite the sepals, anthers dorsifixed; ovary sessile, globose, 1-locular, style
filiform, stigma small, capitate, ovules numerous. Fruits obovoid or ellipsoid capsules, inserted inside
the calyx tube, splitting transversely, the upper half falling as a lid. Seeds cuneate and slightly
flattened, with thick, almost corky marginal wings.
230
LYTHRACEAE (YII)
Fig. 2. Pemphis acidula. A, flowering leafy twig; B, detail of leaf; C, flower; D, longitudinal section
of flower; E, stamens; F, fruit; G, fruit cap; H, seed. (All from S. 21437.)
231
Distribution. A monotypic genus, distributed from the Zanzibar coast to the W Pacific.
Ecology. Confined to exposed rocky coasts or limestone cliffs where coral abounds.
Pemphis acidula J.R. Forst. & G. Forst.
Fig. 2.
(Latin, acidulus = rather sour; the taste of the fruits)

l.c. 68 t. 34; Hooker f. l.c. 572; King l.c. 348; Merrill l.c. 418; Ridley l.c. 821; Masamune l.c. 512; Backer & Bakhuizen f. l.c.
255; Everett & Whitmore l.c. 276; Shea in Cockburn l.c. 214; Anderson l.c. 240; Corner l.c. 475.
Shrub or tree to 10 m tall, 30 cm diameter; stem usually short and stocky. Bark dark grey-brown,
finely fissured; inner bark light reddish, laminated. Twigs pale-colured, nodes prominent. Leaves
thick and fleshy; elliptic-oblong, 13.2 x 0.30.5 cm, densely and finely hairy; base acute, margin
entire, apex acute, rarely obtuse. Flower erect, scentless; pedicels 510 mm long; calyx c. 7 mm
deep; petals 67 x34 mm, white, elliptic and crinkled. Fruits c. 1.5 x 1 cm, reddish green and
ripening brown.
Vernacular name. Sarawakmentigi (Malay).
Distribution. As for the genus. In Sabah, known from from the Selingan, Balambangan and Malawali
islands and in Sarawak from Sampadi island and Teluk Bandung on Santubong Peninsula.
232
MALVACEAE
Balu Perumal
World Wide Fund for Nature (WWF),
Merrill, EB (1921) 374; Ridley, FMP 1 (1922) 253; Masamune, EPB (1942) 452; Backer & Bakhuizen f., FJ 1 (1963); van
Borssum Waalkes, Blumea 14 (1966) 23; Hsuan Keng, OFMSP (1969) 161; Kochummen, TFM 1 (1972) 309; Anderson,
CLTS (1980) 242; Corner, WSTM 3rd ed. 2 (1988) 479.
Annual or perennial herbs, shrubs or trees. Bark usually very fibrous, with slime canals. Indumentum
nearly always of stellate hairs or scales and simple hairs, sometimes of simple or stalked glandular
hairs. Leaves spirally arranged, simple, entire to deeply lobed, mostly palmately veined, nectaries
commonly found on the veins below; stipules present. Flowers radially symmetrical, bisexual, 5merous, solitary and axillary, or in terminal and/or axillary racemes or panicles, often in condensed
clusters or solitary by reduction of the inflorescence; epicalyx usually present, usually persistent, with
3 to many free or connate leafy segments; calyx connate at base, lobes valvate, persistent; disc none,
but nectariferous tissue present at the base of calyx; corolla contortate (especially in bud), adnate at
base to the staminal tube and falling off with it; stamens many, connate into a long tube, developing
centrifugally, anthers dorsifixed, 1-locular, pollen grains spiny, very large (visible to unaided eye);
ovary superior, (3)5many-locular, ovules 1many per locule, placentation axile; styles as many as
or twice the number of ovary locules, mostly united to various degrees, stigmas as many as styles, on
distinct style-arms or almost united. Fruit a schizocarp or a capsule, breaking into separate parts,
loculicidally dehiscent, or indehiscent. Seeds usually hairy; endosperm usually oily; embryo mostly
curved; cotyledons usually folded.
Distribution. About 116 genera and 1550 species, distributed throughout the world, but mainly found
in the tropics and subtropics. In Sabah and Sarawak, represented by 7 genera comprising 13 species
and a hybrid (Hibiscus xarcheri W. Watson). Pavonia rigida, although recorded for Kalimantan, has
not been collected from Sabah or Sarawak.
Ecology. Generally sun-loving (or heliophilous) and weedy plants, commonly found near coastal
areas and in lowland forest.
Uses. The most important economic product of the family is cotton, produced by Gossypium. The
bark of certain species of Hibiscus yields valuable fibres that are used for cordage or manufactured
into textiles. Oil is obtained from the seed of several species and the residue is made into oil-cakes for
stock feed. Several shrubs are cultivated for their showy flowers; of these the bunga raya, Hibiscus
rosa-sinensis, is widely grown throughout the tropics. Though the possible origin of the species is
Tanganyika, bunga raya has been designated the national flower of Malaysia. Some species are
cultivated for vegetables, e.g., the Ladies Fingers, Abelmoschus esculentus. A few others are
medicinal, e.g., species of Sida. The timber is not of economic importance but that of a few species is
used locally.
233
Taxonomy. The Malvaceae are very closely related to the Bombacaceae. Spiny pollen grains, 1locular anthers and a well-developed staminal tube are the main characters distinguishing the
Malvaceae. Moll & Janssonius (quoted in Desch, Malay. For. Rec. 15, (1954) 572) have also
described differences in the wood characters.
Key to genera
1. Apex of staminal tube 5-toothed, filaments projecting from its whole surface or from a
greater part of it; epicalyx present.......................................................................................2
Apex of staminal tube without teeth, but splitting into numerous filaments; epicalyx
absent..................................................................................................................................6
2. Styles twice as many as carpels; always 10. Fruit a schizocarp.............................................
Urena L.
Gen. Pl. ed. 5 (1754) 764; Merrill l.c. 374; Ridley l.c. 256; Masamune l.c. 453; Backer & Bakhuizen f. l.c. 428; van
Borssum Waalkes l.c. 137.
A monotypic genus (U. lobata) with pantropical distribution.

Annual or perennial low shrubs. Leaves palmately lobed and veined, often twice divided or undivided,
with extrafloral nectaries on the veins beneath. Flowers axillary, solitary or in clusters; calyx bellshaped or tubular; staminal tube bearing anthers on the upper half; ovary 5-locular, each locule with
one ovule. Fruit breaking-up into 5 dehiscent or indehiscent, 1-seeded mericarps at maturity.
In Sabah and Sarawak, common on roadsides, waste ground, fallow fields, plantations, and secondary
vegetation, from sea level to c. 2000 m.
Style 1, stigmatic arms as many as carpels, usually 5. Fruit a

capsule..................................................................................................................................3
3. Style divided at apex into 5-ribbed or lobed stigmas. Calyx rim minutely toothed to 5lobed....................................................................................................................................4
Style not divided at apex into ribbed or lobed stigmas. Calyx rim entire or only minutely
toothed.................................................................................................................................5
4. Calyx 5-lobed, not splitting on one side during anthesis, free from corolla, persistent after
flowering.............................................................................................................1. Hibiscus
Calyx minutely 5-toothed, splitting on one side during anthesis, joined to corolla and
falling with it after flowering.................................................................................................
Abelmoschus Medic.
Malv. (1787) 46; Merrill l.c. 375; Masamune l.c. 452; Backer & Bakhuizen f. l.c. 434; van
Borssum Waalkes l.c. 89; Kochummen l.c. 310.
Six species in S and SE Asia and Australia.
Herbs or low shrubs. Leaves palmately lobed and veined. Flowers in axillary or terminal
racemes; calyx at anthesis spathaceous and falling together with the corolla; staminal tube
bearing anthers almost throughout; ovary 5(69)-locular, each locule with many ovules. Fruit
a loculicidal capsule.
Two species in Sabah and Sarawak, viz. A. moschatus and A. esculentus. A. moschatus is
common in open places, especially on waste ground, roadsides, and secondary vegetation, to
1000 m. A. esculentus is cultivated as a vegetable.
234
MALVACEAE (BALU PERUMAL)
5. Epicalyx with 38 segments, small and narrow, mostly caducous. Seed coat glabrous or
short-hairy. Ovary and fruit 5merous..............................................................2. Thespesia
Epicalyx with 3 segments, large and leaf-like, cordate, persistent. Seed coat densely longhairy or woolly. Ovary and capsule 35-merous...................................................................
Gossypium L.
Gen. Pl. ed. 5 (1754) 764; Merrill l.c. 375; Masamune l.c. 452; Backer & Bakhuizen f. l.c.
436; van Borssum Waalkes l.c. 119.
About 2070 species; pantropical.
Straggling shrubs. Leaves mostly palmately lobed or divided. Flowers solitary or in 24flowered racemes, terminal or axillary; calyx cup-shaped; staminal tube bearing anthers
throughout; ovary 35-locular, each locule with many ovules. Capsule loculicidally dehiscent
into 35 valves.
In Sabah and Sarawak, one species, G. barbadense L. var. acuminatum (Roxb.) Mast. is
occasionally cultivated in gardens.
6. Seeds 1 per mericarp; mericarp wall enveloping the seeds to form a globose to flatrounded fruit (schizocarp). Flowers less than 2.5 cm across.................................................
Sida L.
Gen. Pl. ed. 5 (1754); Merrill l.c. 374; Ridley l.c. 254; Masamune l.c. 453; Backer &
Bakhuizen f. l.c. 426; van Borssum Waalkes l.c. 177.
About 150 species, distributed in the tropics and subtropics of the world.
Annual or perennial herbs or low shrubs. Leaves simple, rarely divided or lobed, pinnately or
palmately veined, without extrafloral nectaries. Flowers axillary, solitary or clustered;
epicalyx lacking or present; calyx bell-shaped; corolla rotate; staminal columm usually
shorter than the petals; carpels 514, each with one ovule. Fruits globose or flat-rounded;
mericarps often more or less 3-angular, indehiscent or dehiscent at apex only, in falling
leaving a blunt slender columella. Seeds closely enveloped by the wall of the mericarps.
Two species in Sabah and Sarawak; S. rhombifolia L. with subsp. rhombifolia and subsp.
retusa (L.) Borss., and S. acuta Burm. f. subsp. acuta. Common in open places, especially in
the lowlands.
Seeds 2 or more per mericarp; mericarp arranged in a longitudinal series without

constriction to form a globose to cylindrical fruit (schizocarp). Flowers at least about 2.5
cm across...............................................................................................................................
Abutilon Mill.
Gard. Dict. ed. 4, 1 (1754); Merrill l.c. 374; Masamune l.c. 452; Backer & Bakhuizen f. l.c.
422; van Borssum Waalkes l.c. 159.
About 150 species; in the tropics and subtropics.
Annual or perennial herbs or low shrubs. Leaves divided or undivided, mostly cordate at base,
palmately veined, without extrafloral nectaries. Flowers axillary, solitary or in paniculate
clusters; epicalyx absent; calyx usually bell-shaped; corolla rotate, bell-shaped or rarely
tubular; staminal column usually much shorter than the petals; carpels 540, each with 29
ovules. Fruits globose or cylindrical, rarely discoid; mericarps 540, follicular, dehiscent, in
falling leaving a blunt, slender columella. Seeds 29 per mericarp, reniform.
One species in Sabah and Sarawak, A. indicum (subsp. indicum). Common on waste ground.
235
1. HIBISCUS L., nom. cons.

(a Greek plant-name, possibly referring to the hollyhock)
Gen. Pl. 5 (1754) 310; Merrill, EB (1921) 374; Ridley, FMP 1 (1922) 257; Masamune, EPB (1942) 452; Backer &
Bakhuizen f., FJ 1 (1963) 429; van Borssum Waalkes, Blumea 14 (1966) 25; Kochummen, TFM 1 (1972) 311; Anderson,
CLTS (1980) 242; Corner, WSTM 3rd ed. 2 (1988) 480.
Herbs, shrubs or trees. Indumentum of stellate hairs and/or scales. Leaves simple, lobed or entire,
spirally arranged, often with extrafloral nectaries. Flowers axillary, solitary, often in racemes or
panicles; pedicels mostly articulate, at apex rarely thickened into an obconical or discoid
hypanthium; epicalyx usually persistent in fruit, 3many-lobed, lobes free or shortly connate at base;
calyx bell-shaped, distinctly 5-lobed, rarely splitting on one side, mostly with distinct venation,
sometimes with nectaries, persistent; corolla mostly large and showy, of various colours, often yellow
with a dark purple centre; stamens connate into a tube that adnates to the base of corolla, mostly as
long as or shorter than the petals, rarely longer, bearing anthers throughout or only in the upper half;
ovary mostly 5-locular, rarely 10-locular as a result of 5 false dissepiments, style 1, distally 5branched, stigma discoid, capitate or obscure, ovules 3 to many per locule. Fruit a capsule, splitting
loculicidally into 510 valves. Seeds 3 to many per locule, globose or reniform; testa glabrous or
hairy.
Distribution. At least 250 species; in the tropics and subtropics of the Old and New World; only 2 or
3 species in the temperate zone. Five species and 1 hybrid in Sabah and Sarawak, of which H.
borneensis, H. macrophyllus, H. surattensis and H. tiliaceus are indigenous.
Ecology. Most species are sun-loving and prefer lower altitudes. Herbaceous and shrub species occur
particularly on waste ground, along roadsides or planted as ornamentals in the gardens. Tree species
occur especially in secondary forest. Flowers are known to be pollinated by birds.
Key to Hibiscus species

1. Scrambling spiny shrub. Leaves lobed................................................................................2
Tree, not spiny. Leaves not lobed........................................................................................3
2. Epicalyx segments long-spathulate, their apices broadened..................................................
H. surattensis L.
Sp. Pl. (1753) 696; Merrill l.c. 375; Masamune l.c. 453; van Borssum Waalkes l.c. 57.
Herb or low scrambling shrub, to 2 m tall, often rooting at the nodes; stems slender, with long
internodes, densely covered with recurved prickles, often tinged red. Leaves palmately 35lobed, base 57-veined, shallowly cordate or obtuse, lobes coarsely toothed; extrafloral
nectaries absent; veins prickly beneath; stipules broad and leafy,
at base auricled. Flowers axillary, solitary; epicalyx segments 912, spreading; calyx bellshaped, accrescent; corolla yellow and mostly with a dark purple centre; staminal column
much shorter than the petals, bearing anthers throughout; ovary conical, densely hairy;
stigmas discoid, hairy. Capsule ovoid to globose, outside densely covered with stiff, simple
hairs. Seeds many, reniform.
Throughout the tropics of the Old World. In Sabah and Sarawak, common in young
secondary forest, occasionally on waste ground near houses or villages, from the lowlands to
1200 m.
236
Epicalyx segments linear, their apices not broadened...........................................................

H. radiatus Cav.
Diss. 3 (1987) 150, t. 54, fig. 2; van Borssum Waalkes l.c. 60.
Herb or low scrambling shrub, to 1 m high. Stems slender, typically with recurved prickles.
Leaves palmately 35-lobed, base 57-veined, shallowly heart-shaped or truncate, lobes
coarsely toothed; extrafloral nectaries absent; veins prickly beneath; stipules linear. Flowers
axillary, solitary; epicalyx segments 10, spreading; calyx bell-shaped, accrescent; corolla
yellow with a dark purple centre; staminal column much shorter than petals, bearing anthers
throughout; ovary globose, densely hairy, stigmas capitate. Capsules ovoid to globose, outside
with dense long appressed bristles. Seeds c. 4 per locule, trigonous.
S and SE Asia; in Malesia sometimes planted, also a weed. In Sabah, recorded in the
Kinabatangan and Tongod areas.
3. Twigs and leaf stalks with dense coarse shiny stellate hairs (hair arms to 810 mm
long)......................................................................................................2. H. macrophyllus
Twigs and leaf stalks glabrous or with short-tomentose hairs (hairs fine, at most 23 mm
long)....................................................................................................................................4
4
Leaves glaucous on the lower surface, with extrafloral nectaries in the axils of inner veins
beneath. Ovary and fruit 10-locular (5 true and 5 false dissepiments). Staminal tube
shorter than the petals. Rocky and sandy coasts (cultivated inland)...............3. H. tiliaceus
Leaves not glaucous on the lower surface, without extraflora nectaries. Ovary and fruit 5locular, without false dissepiments. Staminal tube as long as the petals. Inland
forest...........................................................................................................1. H. borneensis
1. Hibiscus borneensis Airy Shaw

(of Borneo)
In Hooker, Ic. Pl. 34 (1939) t. 3377; Van Borssum Waalkes l.c. 49. Type: Native Collector (Oxford Univ. Exp.) 1541,
Borneo, Sarawak, Dulit Trail (holotype K; isotypes GH, L, SING).
Tree to 30 m tall, c. 80 cm diameter, with small buttresses; crown narrow, appearing cylindric, lowbranched. Bark smooth to minutely fissured, tough and fibrous. Twig somewhat angular, covered
with lenticels and minute stellate hairs. Stipules large, 810 x 10 mm, orbicular to reniform, auricled,
stellate-hairy. Leaves coriaceous, not glaucous beneath, on both surfaces with scattered minute
stellate hairs, early glabrescent; ovate to orbicular, 1022 x 816 cm; base cordate or truncate, rarely
acute, margins coarsely, often doubly crenate, apex acute to acuminate; at base 57-veined, with the
veins and intercostal veins forming a cobweb-like pattern; without extrafloral nectaries; petioles 513
cm long. Flowers in axillary, few-flowered racemes or panicles, the lower ones occasionally solitary;
primary peduncles 1028 cm long, secondary peduncles 413 cm long; pedicels 25 cm long;
epicalyx 79-lobed, segments coriaceous; calyx widely bell-shaped, 1.52 cm high, c. 1.5 cm wide,
coriaceous, segments usually recurved, triangular, 67 mm wide and long, stellate-tomentose outside,
at base with a ring of fairly stiff stellate hairs inside; corolla white or pinkish, base pale yellow, shiny,
petals obovate, 46.5 x 22.5 cm, apex rounded, somewhat rough by coarse minute stellate hairs and
short glandular hairs outside, at base with minute stellate hairs inside;
237
staminal tube as long as petals, 3.55.5 cm long, at base downy by slender stellate hairs, for the rest
glabrous, bearing anthers almost throughout, filaments 12 mm long, anthers 11.5 mm long; ovary
conical, c. 7 x 4 mm, hirsute, 5-locular, without false dissepiments, styles 45 mm long, soft-hairy,
stigmas capitate, 1.52 mm wide. Fruits c. 20 x 17 mm, globose to obovoid, covered with fairly long,
simple hairs and minute stellate hairs. Seeds c. 7 per locule, reniform, covered with long red-brown or
yellowish woolly hairs.
Distribution. Endemic to Borneo (Sarawak and Kalimantan).
Ecology. Primary forest, mostly in damp places near rivers, to 300 m.
Taxonomy. The species is closely related to H. decaspermus from S and E Malesia and to H.
floccosus from Peninsular Malaysia. It differs from both species in its stellate-hairy epicalyx, instead
of densely scaly epicalyx.
2. Hibiscus macrophyllus Roxb. ex Hornem.

(Greek, makros = large; phullon = leaf)
Hort. Hafn., Suppl. (1819) 149; Merrill, Philip. J. Sc. 14 (1919) 245; Ridley, FMP 1 (1922) 258; Backer & Bakhuizen f. l.c.
430; van Borssum Waalkes l.c. 47; Kochummen l.c. 31; Corner l.c. 481. Type: Wallich, s.n., Pakistan, Chittagong (holotype
CAL).
Tree to 1525 m tall, c. 20 cm diameter. Bark finely fissured, corky, grey. Twigs with annular stipular
scars, densely covered with minute, brown stellate hairs and coarse, shiny, brown stellate hairs with
long arms 810 mm long. Stipules 815 x 24 cm, initially cohering along the margins and enclosing
the terminal buds, subsequently spreading and falling off, oblong, densely stellate-tomentose outside,
sericeous inside. Leaves chartaceous, densely stellate hairy on both surfaces, lower surface on the
midrib and 26 basal veins with nectaries; more or less orbicular (5)2550 cm across; based 79veined, cordate, margin entire or crenulate, apex cuspidate; inner 26 veins with extra floral nectaries;
petioles stout, 440 cm long, densely covered with coarse, shiny stellate hairs with long arms to 810
mm long. Flowers axillary, solitary or in few-flowered, terminal raceme-like clusters; pedicels stout,
12 cm long, stellate-hairy; epicalyx cup-shaped, usually shorter than the calyx, 1014-lobed; calyx
bell-shaped, 5-lobed, 2.53 cm deep, c. 3 cm wide, segments triangular, acute, 1518 x 67 mm,
outside with stellate hairs, inside sericeous, without nectaries; corolla large and showy, yellow with
maroon eye, often turning red before falling, petals obovate, apex rounded, 67 x 45 cm, outside
stellate hairy, inisde with scattered glandular hairs; staminal tube c. 4 cm long, glabrous, bearing
anthers throughout, filaments 38 mm long, anthers c. 1.5 mm long; ovary conical c. 7 x 5 mm,
covered with stellate-sericeous hairs, 5-locular, style 56 mm long, pilose, stigmas capitate, c. 1.5 mm
across. Fruit 2235 x 1723 mm, ovoid to obovoid, acuminate, with a beak c. 5 mm long, densely
hirsute by simple and stellate hairs. Seeds many, reniform, brownish black, dorsally densely covered
with 34 mm long, ferrugineous, partly spirally twisted hairs, otherwise glabrous.
Vernacular name. Sabahrandog (Dusun Ranau).
Distribution. Peninsular India, E Pakistan, Assam, Thailand, Indo-China, Sumatra, Peninsular
Malaysia, Borneo, and Java. Cultivated as an ornamental in Palawan and the Hawaiian
238
islands. In Sabah, known from Kota Belud (SAN 76302), Kota Merudu (SAN 99493), and Ranau (SAN
61997 and SAN 123132) districts.
Ecology. Secondary forest, from lowland to 1400 m.
3. Hibiscus tiliaceus L.
Fig. 1.
(Latin, tiliaceus = resembling the lime tree, Tilia, of northern temperate forests)
Sp. Pl. (1753) 694; Merrill l.c. 375; Ridley l.c. 259; Masamune l.c. 453; Backer & Bakhuizen f. l.c. 429; van Borssum
Waalkes l.c. 29; Kochummen l.c. 312; Anderson l.c. 242; Corner l.c. 482. Type: Herb. Hermann no. 258, vol. 3, fol. 51
(holotype BM).
subsp. tiliaceus
Tree to 30 m tall, c. 40 cm diameter, without buttresses; crown low, spreading and appearing muchbranched; trunk fairly smooth, with a tough fibrous bark. Twigs terete, covered with lenticels, usually
glabrous but sometimes stellate-hairy, with ring-like stipular scars. Stipules 13.5 x 0.51 mm,
ultimately spreading, ovate to oblong, stellate-hairy outside, usually simple-sericeous inside. Leaves
59-veined at base, chartaceous to coriaceous, sparsely stellate-hairy or glabrous above, glaucous and
densely stellate-hairy beneath; usually orbicular, 7.515 cm across; base cordate, margin entire,
crenulate, or undulate, apex cuspidate, rarely rounded or obtuse; inner 15 veins with extrafloral
nectaries; petioles stout, 3.510 cm long, white hairy towards the junction with the blade. Flowers
axillary, solitary or in few-flowered, terminal raceme-like clusters; pedicels short, stout, stellate-hairy;
epicalyx cup-shaped, usually shorter than the calyx, spreading and often splitting into 811 lobes;
calyx bell-shaped, 5-lobed, after anthesis widening and often splitting, with nectaries on the veins and
stellate-hairs outside, stellate-velutinous along the margins, for the rest simple-sericeous inside;
corolla large and showy, bright yellow with maroon eye, fading dull pink after falling, petals fleshy,
obovate, 57 x 45.5 cm, base rounded, apex rounded, stellate-hairy outside, scattered glandular-hairy
inside; staminal tube 2.53 cm long, glabrous, yellow, bearing anthers throughout; ovary globose to
ovoid, 5-angular, densely sericeous, 10-locular (5 true and 5 false dissepiments), style with 5
glandular hairy arms, purple, stigmas capitate, dark purple. Fruits globose to obovoid, c. 18 mm
across, surrounded by a calyx cup, simple-sericeous or covered with stellate and simple hairs. Seeds
57 per locule, upper surface dotted with minute warts, glabrous or more or less densely stellatehairy, brownish black.
Vernacular names. Sabahkali bang-bang (Dusun Sungai). Sarawak langkubing, menoa
(Kelabit). Bruneibaru-baru (Malay).
Distribution. Widespread in the tropics and subtropics of the Old World and the Pacific Islands.
Throughout the coasts of Sabah and Sarawak; also in Brunei and Kalimantan.
Ecology. Very common along sandy seashores and tidal creeks, a common constituent of the
Barringtonia-formation, rarely in higher places in mangrove, also extending upriver to their brackish
reaches; often planted far inland.
Uses. The fibre of the bark is used by fishermen for string and cordage in making nets and as tow for
caulking boats. Grows well from cuttings and makes an excellent living fence.
239
Fig. 1. Hibiscus tiliaceus. A, flowering leafy twig; B, flower in longitudinal section; C, fruit; D, fruit in longitudinal section;
E, seed. (AB from S. 24529, CE from Balu Perumal 100.)
240
2. THESPESIA Sol. ex Corra, nom. cons.

(Greek, thespesios = divinely beautiful plant)
Ann. Mus. Herb. Paris 9 (1807) 290; Merrill, EB (1921) 375; Ridley, FMP 1 (1922) 259; Masamune, EPB (1942) 453;
Backer & Bakhuizen f., FJ 1 (1963) 435; van Borssum Waalkes, Blumea 14 (1966) 105; Kochummen, TFM 1 (1972) 313;
Corner, WSTM 3rd ed. 2 (1988) 483.
Trees or shurbs. Indumentum of scales or stellate hairs. Leaves spirally arranged, occasionally
clustered towards the ends of twigs, entire or palmately lobed, palmately veined, often with extrafloral
nectaries. Flowers solitary, axillary, often in raceme-like clusters; pedicels mostly not articulate, at
apex thickened into a hypanthium; epicalyx 38-lobed, lobes free, small, caducous; calyx usually
woody, cup-shaped, rim entire or with 5 minute teeth, persistent in fruit; corolla large and showy,
mostly yellow with a dark purple centre; staminal tube much shorter than the petals, bearing anthers
throughout; ovary 5-locular or as a result of 5 false dissepiments becoming 10-locular; style 1, short,
stigmas clavate, 5-grooved or very rarely 5-lobed. Fruit a capsule, indehiscent or loculicidally
dehiscent, 510-locular; fruit wall woody. Seeds 3many per locule, obovoid; seed coat glabrous or
short hairy.
Distribution. About 15 species, distributed throughout the tropics; 1 species in Sabah and Sarawak.
Thespesia populnea (L.) Sol. ex Corra
Fig. 2.
(Latin, populneus = resembling the poplar tree, Populus, of northern temperate forests)
l.c. 290; Merrill l.c. 375; Ridley l.c. 260; Masamune l.c. 453; Backer & Bakhuizen f. l.c. 435; van Borssum Waalkes l.c. 106;
Kochummen l.c. 313; Anderson l.c. 242; Corner l.c. 483. Basionym: Hibiscus populneus L., Sp. Pl. (1753) 694. Lectotype
(van Borssum Waalkes, 1966): Linn. n. 258, Ceylon (BM).
Shrub or small tree to 20 m tall, with rather dense crown. Bark light grey, smooth, becoming rugged
with deep fissures. Twigs terete, densely covered with brown to silvery minute scales, early
glabrescent. Stipules lanceolate to subulate, acute, 310 mm long, scaly. Leaves coriaceous, densely
covered with bronze-green minute scales, early glabrescent; ovate or oblong, triangular to cordate
when young, 723 x516 cm; base deeply cordate, 7-veined, margin entire, apex acute, acuminate, or
long-attenuate; axils of the basal veins mostly with small, saccate, extrafloral nectaries beneath;
petioles 516 cm long, scaly. Flowers axillary, solitary, large; pedicels 2.58 cm long, scaly; epicalyx
segments 3, caducous; calyx cup- shaped, almost entire; corolla bell-shaped, pale yellow with a
maroon eye, fading pink on the tree and lasting several days, petals obliquely obovate, 67 x 4.56 cm,
apex rounded; staminal tube pale yellow, bearing anthers throughout, filaments c. 4 mm long, anthers
c. 1.5 mm across; ovary globose to ovoid, c. 810 mm diameter, scaly, 10-locular, style short c. 4 mm
long, stigma clavate, 5-grooved, c. 5 x3 mm, yellow. Fruits depressed subglobose, not splitting,
faintly 5-angular, 24.5 cm diameter, containing bright yellow gum, subtended by persistent disc-like
calyx. Seeds 4 per locule, obovoid, densely rusty short-hairy or glabrous.
Vernacular names. Sabahbaru (Malay), pangiz (Dusun).
Distribution. Widely distributed in the Old World tropics. Also planted as a shade tree. In Sabah and
Sarawak, occasional in coastal areas, though not as common as H. tiliaceus.
241
Fig. 2. Thespesia populnea. A, flowering leafy twig; B, flower in longitudinal section; C, stamen; D, fruiting leafy twig. (A
C from KEP 7574, D from Balu Perumal 101.)
242
Ecology. Sandy sea shores, confined to the Barringtonia-formation.

Uses. Commonly planted as a roadside tree in some tropical countries.
Notes. Easily confused with Hibiscus tiliaceus, but the entire calyx and non-splitting gummy fruit
which is not enclosed in the calyx are obvious differences.
243
244
MYRICACEAE
A. Noorsiha
Kepong, Malaysia
Merrill, EB (1921) 210; Ridley, FMP 3 (1924) 370; Masamune, EPB (1942) 231; Backer, FM 1, 4 (1951) 277; Hutchinson,
Fam. Fl. Pl. 1 (1959) 189, Gen. Fl. Pl. 2 (1967) 120; Backer & Bakhuizen f, FJ 2 (1965) 1; Keng, OFMSP (1969) 129;
Whitmore, TFM 2 (1973) 295; Anderson, CLTS (1980) 263; Ashton, MNDTS 2 (1988) 322; Corner, WSTM 3rd ed. 2
A monogeneric family (or with 24 genera according to some botanists) with about 50
species, distributed in temperate, subtropical, and tropical regions worldwide, except in
Australia.
Ecology. Often gregarious in open scrublands, marshy places, and mountainous regions. A
number of species have been shown to form nitrogen-fixing root nodules (Bond, Ann. Rev.
Pl. Phys. 18 (1967) 108; Torrey, Bio Science 28 (1978) 586). This seems to correlate with the
occurrence of Myrica in characteristically nitrogen-deficient sites with well-drained or
swampy acids soils where available mineral nutrients are usually also in short supply. The
fully developed male spikes readily emit clouds of pollen when tapped, indicating that the
flowers are wind-pollinated. The fruits are dispersed mainly by birds and squirrels.
Uses. The small, aromatic and decorative trees or shrubs merit planting in avenues, gardens,
and courtyards.
Taxonomy. Myricaceae has been placed in different orders, viz. Juglandales (Melchior in
Englers Syll. Pflanzenfam. ed. 12, 2 (1964) 39), Rutales (Thorne, Nordic J. Bot. 3 (1983)
85), and Myricales (Hutchinson l.c.; Cronquist, Integr. Syst. Class. Fl. Pl. (1981) 59;
Dahlgren, Nordic J. Bot. 3 (1983) 119; Takhtajan, Syst. Magnol. (1987) 95).
MYRICA L.
(Greek, murike = name of an aromatic shrub)
Sp. Pl. (1753) 1024; Merrill, EB (1921) 210; Ridley, FMP 3 (1924) 370; Masamune, EPB (1942) 231; Backer, FM 1, 4
(1951) 277; Backer & Bakhuizen f., FJ 2 (1965) 2; Hutchinson, Gen. Fl. Pl. 2 (1967) 120; Whitmore, TFM 2 (1973) 295;
Anderson, CLTS (1980) 263; Ashton, MNDTS 2 (1988) 322; Whitmore, Tantra & Sutisna, CLK 2,1 (1990) 261. Synonym:
Morella Lour., Fl. Coch. (1790) 548.
Monoecious or dioecious trees or shrubs, often strongly aromatic; young parts often covered
with yellow, peltate, scale-like glands. Leaves spirally arranged, alternate, or sometimes
245
crowded at the ends of twigs, simple or sometimes deeply pinnately lobed, pinnately veined;
stipules absent or rarely present. Inflorescences axillary, simple or branched paniculate
spikes, unisexual (or bisexual, where the male flowers are located on the lower part of the
rachis; rare in Sabah and Sarawak). Flowers unisexual or rarely bisexual (New Caledonian
taxa); perianth absent, or the female flower with 2few sepaloid, whorled bracteoles; male
flowers subtended by a tiny, solitary bracteole, stamens 2many, usually 28, filaments short,
free or united at base, anthers basifixed, 2-locular, dehiscing longitudinally, pistillodes absent
or rarely present; female flowers in clusters of 38, subtended by a bract, staminodes absent,
ovary 1locular, inferior to rarely superior (non-Malesian taxa), style short, 2-branched,
stigmatose on inner side, stigmas 2, ovate-triangular, subulate or filiform, ovules one in each
locule, erect, basal, orthotropous, or pendulous-anatropous. Fruit a drupe; pericarp often
warty, warts waxy; mesocarp thin, pulpy; endocarp hard. Seed 1; testa membranous;
endosperm scanty to absent; embryo straight, cotyledons flat-convex, radicle short.
Distribution. As in the family. In Sabah and Sarawak, 2 species.
Ecology. Coastal areas to mountain tops, to 3300 m.
Key to Myrica species
Twigs usually densely hairy. Leaves of mature trees lanceolate or narrowly oblong-obovate,
thin-coriaceous; base acute to narrowly rounded, margin subentire or distantly serrulate, apex
acuminate to rounded-acute or rarely obtuse; midrib densely hairy on both sides; lateral veins
thinly hairy, arching and looping near the margin; petioles densely tomentose or sometimes
glabrous. Stigmatic arms filiform or subulate....................................................1. M. esculenta
Twigs usually glabrous. Leaves of mature trees obovate or oblong-obovate, thick-coriaceous;
base acute, margin serrate or crenate, apex rounded or slightly emarginate; midrib glabrous
on both sides; lateral veins glabrous, not arcuating nor looping but ending in marginal
toothlets; petioles glabrous. Stigmatic arms ovate-triangular..............................2. M. javanica
1. Myrica esculenta Buch.-Ham.
(Latin, esculentus = edible and nourishing; the fruit)
In D. Don, Prod. Fl. Nep. (1825) 56; Gamble, J. As. Soc. Beng. 75, 4 (1915) 404; Merrill l.c. 210; van Steenis, Bull. Jard.
Bot. Btzg. 3, 13 (1936) 229; Masamune l.c. (1942) 231; Backer l.c. 278; Backer & Bakhuizen f. l.c. 2; Keng l.c. 133;
Whitmore l.c. 295; Anderson l.c. 263; Ashton l.c. 322; Whitmore, Tantra & Sutisna l.c. 261. Type: Hamilton, s.n., Nepal
(K). Synonyms: Myrica sapida Wall., Tent. Fl. Nep. (1826) 59; M. esculenta Buch.-Ham. var. sapida (Wall.) A. Chev.,
Mon. Myric. (1901) 122, Merrill l.c. 210, Masamune l.c. 231; M. farquhariana Wall. l.c. 61; M. esculenta Buch.-Ham. var.
farquhariana (Wall.) A. Chev. l.c. 125, Merrill l.c. 210, Masamune l.c. 231; M. missionis Wall., Cat. (1832) No. 7297; M.
esculenta Buch.-Ham. var. missionis (Wall.) A. Chev. l.c. 126, Merrill l.c. 210, Masamune l.c. 231; M. lobbii Teijsm. &
Binn. ex Miq., Fl. Ind. Bat. 1 (1858) 872; M. esculenta Buch.-Ham. var. lobbii (Teijsm. & Binn. ex Miq.) A. Chev. l.c. 127,
Merrill l.c. 210, Masamune l.c. 231.
Monoecious small bushy tree, treelet or shrub to 15 m tall, 15 cm diameter; crown uneven or
oblong; bole often twisted and crooked. Bark fissured or smooth, blackish grey. Twigs and
246
MYRICACEAE (NOORSIHA)
terminal buds densely clothed with long, thin hairs and scattered yellow, peltate glands;
stipules present in young plants. Leaves spirally arranged or sometimes crowded at the ends
of twigs; thin-coriaceous, dark green and shiny above, sparsely hairy or glabrous and
sparsely glandular on both surfaces, sometimes pitted; lanceolate to narrowly oblongobovate, 311 x1.54 cm; base acute or narrowly rounded, margin subentire or distantly
serrulate, apex acuminate, rounded-acute or obtuse; midrib flat or slightly depressed above,
thick and prominent below, densely hairy on both sides; lateral veins 512 pairs, reddish
brown, slender, thinly hairy, arching and looping near the margin, prominent below;
intercoastal veins reticulate, reddish brown, conspicuous on both surfaces; petioles stout, 0.2
1.2 cm long, densely tomentose, sometimes glabrous. Inflorescences: males 16.5 cm long,
patent, branched, rather laxly flowered at the lower part, bracts ovate-triangular, 1.52.5 x1
1.5 mm, densely hairy, bracteoles ovate-triangular, c. 1.5 x1 mm, long-hairy outside; females
to 5 cm long, erect, unbranched, densely flowered on the upper part, laxly flowered on the
lower part, bracts ovate-triangular, 1.753 x12 mm, hairy and glandular outside, bracteoles
2, ovate-triangular, to 1.5 x1 mm, hairy and glandular outside. Flowers: males yellowish red,
turning to brown on drying, stamens 24, filaments 0.250.5 mm long, united at base,
pistillodes absent; females c. 2 mm across, pale green, in clusters of 36(10), ovary densely
hairy when young, styles reflexed, stigmatic arms filiform or subulate, c. 2.5 mm long. Fruits
ellipsoid-globose, 12 x1 cm, slightly flattened, dark bright red when mature; pericarp warty,
yellow, pulpy with a sweet-sourish taste when fresh.
Vernacular names. Sarawakmerbuta gunung, telur cicak (Malay).
Distribution. S and E Asia, stretching from India to Nepal, China, Myanmar, Thailand, IndoChina, and Malesia (Sumatra, Peninsular Malaysia, Java, Borneo, Phillipines, Sulawesi and
Lesser Sunda Islands). In Sabah and Sarawak, recorded from many areas.
Ecology. Uncommon in primary forest, occasional in kerangas forest, often locally abundant
in secondary forest and open scrublands on well-drained stony laterite soils, rocky beach,
sand-dunes, sandstone pavements, cliffs and screes, from lowlands to 1500 m. Flowering and
fruiting throughout the year.
Uses. Occasionally cultivated for its edible fruits, used in drinks and desserts. The bark has
been reported as a source of tanning material and dyes; its decoction is used in traditional
medicine. The waxy substance of the fruit pulp is used for making candles and soap.
Notes. Leaves of saplings and seedlings are very different from those of mature trees: blades
1923 x3.55 cm, margins deeply, pinnately incised; lateral veins to 15 pairs; stipules ovatelanceolate, 0.50.75 cm long, caducous.
2. Myrica javanica Blume
Fig. 1.
(of Java)
Bijdr. Fl. Ned. Ind. (1825) 517; Stapf, FMK (1894) 231; Merrill l.c. 210; van Steenis, Bull. Jard. Bot. Btzg. 3, 13 (1936) 229,
l.c. 3, 17 (1948) 389; Masamune l.c. 231; Backer l.c. 277; Backer & Bakhuizen f. l.c. 2; Anderson l.c. 263; Ashton l.c. 324;
Whitmore, Tantra & Sutisna l.c. 261. Type: Reinwardt, s.n., West Java, Mt. Gede (holotype L).
247
Fig. 1. Myrica javanica. A, flowering (male) leafy twig; B, side-branch of a male inflorescence; C, male flower; D, a cluster
of female flowers; E, female flower. (AC from K. M. Wong s.n., 4 June 1993, DE from Kostermans c.s. (UNESCO) 262.)
248
MYRICACEAE (NOORSIHA)
Monoecious small bushy tree or erect shrub, 212(20) m tall, 1016 cm diameter; crown
dense; bole straight or crooked. Bark brown; inner bark red; sapwood white. Twigs reddish
brown, glabrous, sparsely glandular. Leaves thick-coriaceous, glabrous, sometimes darker
below, densely glandular on both surfaces, sometimes shallowly pitted; obovate to obovateoblong, 38.5 x1.54 cm; base acute, margin serrate or crenate, apex rounded or slightly
emarginate; midrib slightly flat above, prominent below, glabrous on both sides; lateral
veins 58 pairs, slender, reddish brown, sunken above, rather prominent below, glabrous on
both sides, ending in marginal minute teeth, often forked; intercoastal veins densely
reticulate, distinct below; petioles stout, 210 mm long, glabrous, sparsely glandular.
Inflorescences: males branched, rachis angular, widely patent, to 6.5 cm long, pinkish
brown, bracts ovate-pointed, 12 mm long, shortly hairy, glandular; females 37 cm long,
erect, hardly branched, rather remotely flowered, bracts ovate-triangular, 12 mm long, acute,
hairy and glandular, bracteoles 2, adhered to the ovary, ovate, long-hairy along the margin,
glandular. Flowers: males yellow turning to red, stamens 34, filaments 11.5 mm long,
most part united into a short, thick staminal column, densely tomentose and yellowish
glandular, pistillodes absent; females in clusters of 58, ovary ovoid-ellipsoid, 0.52 mm,
hairy, style spreading, stigmatic arms ovate-triangular, c. 1.3 mm long. Fruits red-brown to
black, ovoid-globose or broadly ellipsoid, c. 0.5 cm diameter, slightly flattened, minutely
warty, glabrous, glandular; pericarp thin, yellow, pulpy; sap purplish, of acid taste.
Distribution. Sumatra, Java, Borneo, Philippines, Sulawesi, Lesser Sunda Islands, and New
Guinea. In Sabah, recorded mainly from Mt. Kinabalu; in Sarawak, known from Mt. Murut,
Mt. Api, Mt. Kanyi, Mt. Mulu, Mt. Dulit and Bt. Batanga; also in Brunei and Kalimantan.
Ecology. In lower montane to open mossy montane forest and shrubby subalpine vegetation,
at 10003300 m. Often found on precipitous ridge crests, limestone hills, volcanic and granite
rocks, and in open grassy thickets on peaty substratum. Flowering and fruiting throughout the
year.
Uses. The timber has been used mainly for making charcoal and as firewood. Ripe fruits are
consumed fresh. In Java, cultivated as roadside trees or planted in the reafforestation of
degraded lands in the mountains.
249
250
NYCTAGINACEAE
A. Latiff
Universiti Kebangsaan Malaysia,
Bangi, Selangor, Malaysia
Ridley, FMP 3 (1924) 1; Stemmerik, FM 1, 6 (1964) 450; Hsuan Keng, OFMSP (1969) 111; Whitmore, TFM 2 (1973) 297;
Anderson, CLTS (1980) 382; Corner, WSTM 3rd ed. 2 (1988) 598.
Trees, shrubs, herbs, or climbers; wood (in Pisonia) with included phloem. Leaves opposite,
alternate or rarely in whorls, simple, without stipules. Inflorescences cymose, often thyrsoid,
corymbose or umbellate, terminal, axillary or sometimes borne on the stems; bracts and
bracteoles present. Flowers radially symmetrical, bisexual or unisexual, pedicellate,
sometimes enclosed by bracts; the bracts often brightly coloured or resembling perianth
lobes; perianth tubular, bell-shaped, funnel-shaped or urceolate, shortly 510-lobed,
sometimes adnate to the pedicel, the basal part persistent, enclosing the receptacle, clavate or
funnel-shaped, the apices partly plaited or valvate in bud, green or variously coloured;
stamens 140, in 12 whorls, connate at base, free from the perianth; anthers 2-locular,
latrorse, basifixed; ovary subsessile, superior, 1-locular, ovule anatropous, style terminal,
stigma capitate or frilled to shortly lobed. Fruits indehiscent, enclosed by basal persistent
part of the perianth lobes, smooth, or with viscid ribs and glands, sometimes the glands
accrescent into prickles; pericarp thin. Seeds 1 per fruit; embryo straight or folded;
endosperm abundant.
Distribution. About 26 genera and c. 300 species, mostly American, particularly in South
America. In Malesia, 4 genera and 19 species; only Pisonia comprises small native trees. In
Sabah and Sarawak, the family is represented by 4 genera and probably 9 species, of which
only two are small trees, the rest being either shrubs, climbers or herbs.
Ecology. Occurring in lowland to montane forest, to c. 2000 m, on various types of soil.
Boerhavia is a genus of pantropical weeds; Mirabilis is naturalised and weedy, and often
cultivated as an ornamental; Bougainvillea is widely grown as an ornamental and Pisonia is a
genus of trees or treelets in various forest types especially in coastal areas.
Uses. Bougainvillea is the best known member of the family in Malesia as well as in Sabah
and Sarawak; many cultivars are very commonly cultivated for their colourful bracts. Pisonia
grandis is also cultivated for its foliage.
Key to genera
1. Herbs, unarmed. Flowers bisexual, in cymes of 210.........................................................2
Woody plants, sometimes thorny. Flowers bisexual or unisexual, in dense flowerclusters................................................................................................................................3
251
2. Leaves in a pair typically unequal in size. Inflorescence without involucre. Fruit with 5
10 prominent ribs and viscid glands..................................................................................
Boerhavia L.
Gen. Pl. ed. 5 (1754) 4; Stemmerik, FM 1, 6 (1964) 452.
Three variable species, namely B. diffusa, B. chinensis, and B. erecta; pantropical.
Annual herbs, erect, ascending or creeping, covered with minute hairs or clavate
glands or hairs. Stem often red-tinged, nodes swollen, becoming constricted on
drying. Inflorescences axillary. Flowers bisexual. Seeds with longitudinally folded
embryos.
In Sabah and Sarawak, these are introduced and have become weeds of open places.
Leaves in a pair of equal size. Inflorescence with involucre. Fruits with faint ribs, without
viscid glands..........................................................................................................................
Mirabilis L.
Gen. Pl. ed. 5 (1754) 82; Masamune, EPB (1942) 272; Stemmerik, FM 1, 6 (1964)
451.
About 60 species, mostly American.
Erect herb, c. 60 cm tall. Leaves oblong to slightly 3-angular; petioles c. 2.5 cm long.
Inflorescence a terminal corymb, 1many-flowered, each flower subtended by a
persistant involucre which is divided halfway into 5 oblong, acute lobes. Flowers
bisexual, ephemeral, trumpet-shaped. Seeds with recurved cotyledons; endosperm
mealy.
One species from the Himalayas and SW China, M. jalapa, has been introduced to
Sabah and Sarawak as an ornamental.
3. Each pedicel bearing one sessile coloured bract, 36 cm long. Shrubs or coarse climbers...
Bougainvillea Comm. ex Juss.
Gen. (1789) 91; Stemmerik, FM 1, 6 (1964) 455.
About 14 species, native of Central and Tropical South America.
Coarse climbers with supra-axillary spines. Leaves opposite, ovate to elliptic-oblong.
Inflorescence supra-axillary, peduncle bearing a single apical cluster of 3 flowers;
perianths tubular, limb 45-lobed, tube with 45 ribs. Fruits spindle-shaped,
coriaceous, 5-ribbed, not viscid. Embryo longitudinally coiled.
Three species, with many hybrids and cultivars are cultivated as ornamentals in
Malesia, including Sabah and Sarawak; particularly common are B. glabra and B.
spectabilis.
Each pedicel bearing 13 small, greenish bracteoles. Small trees, rarely climbers...Pisonia
PISONIA Plum. ex L.
(W. Piso, b. 1611, a Dutch physician)
Gen. Pl. ed. 5 (1754) 451; Ridley, FMP 3 (1924) 2; Heimerl in Engler & Prantl, Pflz. Fam. ed. 2, 16c (1934) 126; Burkill,
EPMP 1 (1935) 1755; Stemmerik, Blumea 12 (1964) 275, FM 1, 6 (1964) 457; Whitmore l.c. 298; Anderson l.c. 282; Corner
l.c. 599.
Trees to 30 m tall, unarmed, or thorny climbers (P. aculeata). Branches sympodial. Bark soft
and spongy, brittle, pale cream. Leaves opposite, entire, dull, drying chartaceous and black to
252
NYCTAGINACEAE (LATIFF)
dark brown; midrib flat above. Inflorescences axillary or terminal cymes (rarely borne on
leafless stems or branches), 28-times umbellately branched, each ultimate branch bearing 1
3 flowers; pedicels with 13, small, caducous bracteoles. Flowers bisexual or unisexual,
small, greenish white; perianths fleshy, valvate in bud, bell-shaped, tubular, urceolate, or
funnel-shaped, 5- or rarely 10-lobed, basal part tubular, coriaceous, persistent, accrescent and
elongating after anthesis, apical part coloured, often circumscissile; stamens 240, in 12
whorls, mostly exserted and sterile; ovary, elongated, smooth, style longer than the ovary,
stigma capitate, lobes frilled, radiating or unilateral. Fruits coriaceous, often crowned by a
perianth limb, smooth or with 5 ribs, viscid due to the presence of longitudinal rows of
glands, which sometimes developing into prickles; sometimes a rostrum also developed.
Seeds oblong; embryo straight, deeply longitudinally furrowed; cotyledons recurved and
surrounding the perisperm, the latter sometimes reduced to a gelatinous mass.
Distribution. About 35 species, with 20 in the Americas, only one in East Africa, and two
others in the Madagascar area, few in continental SE Asia; eight native in Malesia, including
three in Sabah and Sarawak.
Ecology. In primary rain forest, especially near the coast.
Key to Pisonia species
1. Spiny climber. Perianth limb with 5 large lobes, alternating with 5 smaller ones. Fruit 5ribbed, each rib with a row of biseriate glandular appendages..............................................
P. aculeata L.
Sp. Pl. (1953) 1026; Ridley l.c. 3; Stemmerik, FM 1, 6 (1964) 467; Whitmore l.c. 298.
Scandent climber c. 20 m high, with mostly recurved thorns. Leaves elliptic 410
x 1.25 cm. Flowers unisexual, in dense, axillary cymes. Fruits clavate, c. 15 x 2 mm,
sericeous.
Tropical and subtropical America, Africa, Madagascar, Mauritius, Seychelles, Sri
Lanka, India, the Andaman islands, Myanmar, Indo-China, throughout Malesia,
Australia, and New Caledonia. In Borneo, very rare on the northern coast of Sabah.
Tree or erect shrub, unarmed. Perianth limb with 5 rows of black glands or short brown
hairs. Fruit 5-ribbed, each rib bearing uniseriate prickles or a row of viscid
glands....................................................................................................................................2
2. Leaves with distinct dark veins contrasting with lighter coloured intercostal veins; lateral
veins and intercostal veins hairy beneath. Perianth limb glandular. Fruit, with 5 rows of
uniseriate prickles.............................................................................................1. P. grandis
Leaves without distinctly contrasting dark veins, glabrous on lower surface. Perianth limb
not glandular. Fruits with rows of viscid glands.....................2. P.umbellifera
1. Pisonia grandis R. Br.
(Latin, grandis = large; the leaves)
Prod. Fl. Nov. Holl. 1 (1810) 422; Ridley, J. Str. Br. Roy. As. Soc. 45 (1905) 215; Backer & Bakhuizen f., FJ 1 (1963) 272;
Stemmerik l.c. (1964) 464; Whitmore l.c. 298; Corner l.c. 599. Type: R. Brown, s.n., north coast of Australia (BM).
Synonym: P. alba Span., Linnaea 15 (1841) 342, Corner l.c. 599.
Shrub or tree to 30 m high, c. 100 cm diameter, usually much smaller, with large exposed
253
Fig. 1. Pisonia umbellifera. A, flowering leafy twig; B, bisexual flower with some sepals, petals and stamens removed; C, a
portion of male inflorescence; D, male flower; E, fruit. (AB from Podzorski SMHI 985, CD from Leonardo Co 3410, E
from Saw et al. FRI 40010.)
254
roots; wood soft and light. Twigs light-coloured, when dry with conspicuous furrows and
large leaf scars. Leaves membranous; elliptic, oblong or ovate, 1020 x 610 cm; base
cuneate, rounded or cordate, mostly asymmetric, apex acuminate; with red or dark coloured
veins, sericeous, early glabrescent, tardily so on the veins beneath; petioles 16 cm long.
Inflorescence a terminal, cymose, 1.73.5 x 34.5 cm; peduncles c. 1.5 cm long, light-brown
hairy; pedicels 12.5 mm long, at the upper or lower part with 24 oblong bracteoles.
Flowers bisexual; perianth white, funnel-shaped, c. 4 mm long, 5-lobed, limbs with 5 rows of
black glands; stamens 610, exserted for 2 mm long; stigma frilled, not exserted. Fruits
cylindrical to clavate, c. 12 x 2.5 mm, with 5 ribs, each bearing a row of viscid uniseriate
prickles of c. 1 mm long, hairy between the ribs; stalks 11.5 cm long. Seeds 910 x 1.52
mm.
Distribution. From Madagascar through Malesia to Australia and the Pacific. In Sabah,
known only from Pulau Sibun, Lahad Datu district.
Ecology. Wild on the northern coast of Sabah; presently widely cultivated as an ornamental
in urban areas for its beautiful yellowish green foliage.
2. Pisonia umbellifera (J. R. Forst.) v. Seemen
Fig. 1.
(Latin, umbelliferus = umbel-bearing; the inflorescences)

Bonplandia 10 (1862) 154; Kurz, For. Fl. Burm. 2 (1877) 279; Stemmerik l.c. (1964) 460; Whitmore l.c. 298; Anderson l.c.
282. Basionym: Ceodes umbellifera J. R. Forst., Char. Gen. Pl. (1776) 71. Type: Forster, s.n., Tanna (BM). Synonym: P.
excelsa Blume, Bijdr. Fl. Ned Ind. 14 (1826) 735.
Shrub or tree to 28 m high, unarmed. Bark smooth, greyish; inner bark pale green; sapwood
soft, cream. Leaves sometimes crowded at the ends of twigs, glabrous beneath; ovate to
elliptic-oblong, 923 x 411 cm; base cuneate to rounded, apex acute to rounded; without
contrasting dark veins; petioles 0.54 cm long. Inflorescence a terminal, branched, manyflowered compound umbel, 39 cm across, sericeous or glabrous; peduncles 3.56 mm long;
pedicels 1.56 mm long, with 13 small bracteoles at the base or higher. Flowers bisexual or
unisexual, yellow; perianth 2.57 mm long, bell-shaped, limbs with short brown hairs;
stamens 614, exserted to 4 mm; stigma frilled, exserted for c. 1 mm in unisexual flowers and
c. 1.5 mm in bisexual flowers. Fruits cylindrical, black, 24 x 0.3 cm, 5-ribbed, each rib
with a row of viscid glands; stalks 0.71.5 cm long. Seeds 1720 x 2 mm.
Vernacular names. Sabahtintotobokon, tutubokon (Dusun).
Distribution. South Africa, Madagascar, Indo-Malesia, Australia and the Pacific. Common
throughout Sabah but less common in Sarawak.
Ecology. In Sabah, it is found in various habitats, from coastal forest as on Banggi Is.,
limestone at Madai Caves, to higher elevation on the Crocker Range. However, in Sarawak, it
has only been collected on limestone at Niah, although it also occurs in coastal vegetation on
Talang-talang island.
255
256
SANTALACEAE
L.S.L. Chua
Kepong, Malaysia
Bentham, Fl. Austral. 6 (1873) 211; Hooker f., Fl. Br. Ind. 5 (1890) 228; King, J. As. Soc. Beng. 75, 2 (1912) 269; Merrill,
EB (1921) 241; Ridley, FMP 3 (1924) 166; Danser, Nova Guinea N.S. 4 (1940) 133; Masamune, EPB (1942) 257; Backer &
Bakhuizen f., FJ 2 (1965) 76; Stauffer, Vierteljahrsschrift Naturf. Ges. Zurich 114 (1969) 49; Whitmore, TFM 2 (1972) 341;
Anderson, CLTS (1980) 310; Hewson & George, Fl. Austral. 22 (1984) 29.
Small to medium-sized trees or hemiparasitic, climbing or epiphytic shrubs. Leaves simple,

alternate or spirally arranged, rarely opposite, often reduced to minute scales, persistent or
caducous; stipules absent. Inflorescence a raceme, spike, panicle or corymb, rarely a solitary
flower, axillary or terminal, usually bracteate. Flowers small, radially symmetrical, unisexual
or bisexual; perianth in one series, 35-lobed, lobes valvate in bud, inserted on a shallow,
cup-like receptacle, perigynous or epigynous; disc present, often thickened and lobed at
margin, rarely absent; stamens 35, opposite perianth lobes, anthers basifixed or almost so, 2locular, dehiscing longitudinally; ovary superior or inferior, 1- or incompletely 512-locular,
with 15 pendulous ovules, placentation free; style very short or none; stigma entire or 25lobed. Fruit a drupe or pyrene; pericarp often fleshy; endocarp usually hard; receptacle often
enlarged, fleshy and coloured. Seeds 1 to several per fruit, entire or lobed, without testa;
endosperm copious.
Distribution. Worldwide, with 37 genera and about 1400 species. In Sabah and Sarawak,
represented by 5 genera and 6 (9) species, of which only 2 genera with 2 species are trees.
Ecology. Occurring in a wide variety of habitats on nutrient-poor soils such as white sands, in
swamps, on limestone, and in mossy forest, from lowland to 3000 m.
Uses. The most well-known product of the family is sandalwood, derived principally from
Santalum album, native to India, Myanmar, East Java, Lesser Sunda Islands, and northern
Australia. Its aromatic heartwood or the oil distilled from it has important cosmetic and
ceremonial uses in Asia. The indigenous species have no commercial value.
Taxonomy. Santalaceae are closely related to the Olacaceae, Opiliaceae and Loranthaceae in
the order Santalales.
Key to genera
1. Small to medium-sized trees...............................................................................................2
Climbers or hemiparasitic shrubs, sometimes epiphytic.....................................................3
257
2. Trees with spiny bole. Ovary inferior. Fruit without swollen receptacle.....2. Scleropyrum
Trees without spiny bole. Ovary superior. Fruit with swollen receptacle........1. Exocarpos
3. Leaves minute, scale-like, caducous. Flowers sessile...........................................................
Phacellaria Benth.
Gen. Pl. 3, 1 (1880) 229; Danser, Blumea 3 (1940) 222.
Seven species; SE China, Indo-China, Myanmar, Thailand, Peninsular Malaysia,
Borneo.
Branches without haustoria. Male flowers with 36, 3-angular perianth lobes; anthers
cordate. Female flowers with 48 perianth lobes; stigma lobed; ovary inferior,
obovoid to cylindrical. Fruit a drupe, crowned by persistent perianth lobes and disc.
Seed 1, 5-grooved; endosperm copious; embryo small, basal.
In Sabah, 1 species (P. malayana); uncommon, known from only a single collection
(SAN 27605) from Obar Ridge, south of Sugut in Sandakan district.
Leaves large and well-developed, persistent or reduced to minute scales. Flowers
stalked.................................................................................................................................4
4. Woody climbers; stems not twining. Seeds several per fruit, manylobed.............................
Dendrotrophe Miq.
Fl. Ind. Bat. 1 (1855) 779; Backer & Bakhuizen f. l.c. 76; Stauffer l.c. 53; George in
Hewson & George l.c. 59. Synonym: Henslowia Blume, Mus. Bot. Lugd. Bat. 1
(1850) 242, Merrill l.c. 241, Masamune l.c. 257.
About 30 species; Himalayas eastwards to Malesia.
Dioecious or monoecious plants. Leaves spirally arranged. Inflorescences umbellate;
flowers unisexual or bisexual, perianth lobes 5; ovary inferior with several to many
incomplete septa, style short, stigmas 5. Fruit a pyrene with 812 longitudinal rows
of warts.
In Sabah and Sarawak, 25 species, widespread.
Hemiparasitic shrubs; stems twining. Seed one per fruit, entire............................................
Dendromyza Danser
Nova Guinea N. S. 4 (1940) 133; Backer & Bakhuizen f. l.c. 76; Stauffer l.c. 59;
George in Hewson & George l.c. 60.
Five species; Sumatra, Java, Borneo, the Philippines, New Guinea, Australia and
Solomon Islands.
Dioecious plants. Branches either leafy or with haustoria and minute scales. Perianth
lobes 45, triangular, persistent; ovary 1-locular, style short, stigmas 5. Fruit a pyrene,
smooth, 1-locular, easily splitting into 4 or 5 parts.
One species (D. reinwardtiana) in Sabah, known from Ranau, Kinabatangan and
Sandakan districts (Chew, Corner & Stainton 830, SAN 37869, SAN 95107, SAN
38577).
1. EXOCARPOS Labill.
(Greek, exo = outside, karpos = fruit; fruit borne on a swollen receptacle)
Voy. 1 (1798) 155, t. 14; Blume, Mus. Bot. Lugd. Bat. 1 (1850) 180; Bentham, Fl. Austral. 6 (1873) 228; Heyne, Nutt. Pl.
Ned. Ind. (1950) 589; George in Hewson & George, Fl. Austral. 22 (1984) 34.
258
Fig. 1. Exocarpos latifolius. A, flowering and fruiting leafy twig; B, flower; C, flower with two perianth lobes removed; D,
fruit. (All from SAN 134615.)
259
Small trees or erect shrubs; bole without spine. Leaves alternate, rarely opposite, sometimes
reduced to minute scales. Inflorescences an axillary spike or sometimes reduced to a sessile
cluster of flowers. Flowers unisexual or bisexual; perianth 45-lobed; disc glandular;
stamens 45, filament short, broad; ovary superior, stigma blunt, sessile, ovule 1, anatropous.
Fruit a drupe; receptacle swollen, fleshy.
Distribution. About 26 species; Malesia to Australia, New Caledonia, New Zealand and the
Hawaiian Islands. In Malesia, known from Java, Borneo, the Philippines, and New Guinea. In
Sabah, one species, not recorded from Sarawak or Brunei.
Exocarpos latifolius R. Br.
Fig. 1.
(Latin, latifolius = broad-leaved)

Prodr. (1810) 356; Bentham l.c. 228; Merrill, Enum. Philip. Fl. Pl. 2 (1923) 114; Heyne l.c. 589; Backer & Bakhuizen f. l.c.
76; George in Hewson & George l.c. 36. Type: R. Brown, s.n., 6 November 1802, Australia, Queensland, Coen River (BM,
BRI).
Tree to 10 m tall, 20 cm diameter. Leaves thick-coriaceous, orbicular to ovate, 38.5 x 27

cm; base broadly attenuate to rounded, margin recurved, apex rounded to emarginate, 3veined at the base; veins raised on both surfaces, intercostal veins indistinct; petioles 0.30.9
cm long. Inflorescences an axillary spike, 0.52.5 cm long, often branched, tomentose.
Flowers tiny; perianth lobes 5, broadly triangular-ovate, hairy. Fruit globose, scurfy, c. 0.8
x 0.7 cm; receptacle obovoid, 0.40.7 cm long.
Distribution. Malesia to Australia, New Caledonia, New Zealand and the Hawaiian Islands.
In Malesia, known from Java, Borneo, the Philippines, and New Guinea. In Sabah, known
only from a few collections (e.g., Berhaman & Madani SAN 134615) from Pulau Manukan
and Pulau Gaya on the west coast.
Ecology. In Sabah, known from sandstone and shale ridges fringed by coastal kerangas
vegetation.
2. SCLEROPYRUM Arn.
(Greek, skleros = hard, pyrus = pear; hard, pear-shaped fruit)
Jard. Mag. Zool. & Bot. 2 (1838) 549; Hooker f., Fl. Br. Ind. 5 (1890) 234; King, J. As. Soc. Beng. 75, 2 (1912) 275;
Lecomte, Fl. Indo-Chine 5 (1915) 220; Merrill, EB (1921) 242; Ridley, FMP 3 (1924) 171; Danser, Nova Guinea N. S. 4
(1940) 133; Masamune, EPB (1942) 258; Whitmore, TFM 2 (1972) 343; Anderson, CLTS (1980) 310.
Small to medium-sized trees; bole spiny. Leaves spiral. Inflorescence an erect or hanging
spike at leafless nodes on twigs or on the bole. Perianth 5-lobed; disc annular; stamens 5,
inserted at the base of perianth lobes, filaments short; ovary inferior, ovules 3, style short and
thick, stigma peltate. Fruit a drupe, ovoid or pear-shaped, stony, without swollen receptacle.
260
SANTALACEAE (CHUA)
Fig. 2. Scleropyrum wallichianum. A, leafy twig; B, inflorescences; C, flower with one perianth lobe removed; D,
infructescence. (AC from SAN 50943, D from SAN 122235.)
261
Distribution. Six species; India, Sri Lanka, Myanmar, Thailand, Indo-China and W Malesia
(Peninsular Malaysia, Borneo, the Philippines). In Sabah and Sarawak, one species.
Ecology. Widespread on a variety of soils, in the lowland to lower montane forest, sometimes
in swamp, kerangas and limestone forest.
Scleropyrum wallichianum (Wight & Arn.) Arn.
Fig. 2.
(N. Wallich, 17861854, superintendent of the East India Companys Botanic Garden at Calcutta)
l.c. 550; Hooker f. l.c. 234; Lecomte l.c. 221; Merrill l.c. 242; Hansen & Larsen, Stud. Fl. Thailand 50 (1969) 23; Anderson
l.c. 310; Whitmore l.c. 343. Basionym: Sphaerocarya wallichianum Wight & Arn., Edinb. Phil. J. 15 (1832) 180. Type:
Wight Cat. no. 948 (GH, M). Synonym: S. maingayi Hook. f. l.c. 235, King l.c. 276, Ridley l.c. 171, Merrill l.c. 242,
Masamune l.c. 258.
Small tree to 15 m tall, 10 cm diameter; spines on bole to 8 cm long. Bark greyish brown,
smooth to finely fissured; inner bark brown. Sapwood yellow to white, hard. Leaves spirally
arranged to alternate, often drying green, coriaceous; oblong-ovate to broadly elliptic, 926
x413 cm; base broadly rounded or cuneate, margin recurved when dry, apex acute; midrib
sunken on the upper surface, lateral veins 35 pairs, very distant, looping away from the
margin, intercostal veins forming an open network, raised on both surfaces, more prominent
on the lower surface; petioles to 1 cm long. Inflorescences 0.58 cm long, covered with
appressed velvety hairs; bracts triangular. Flowers: perianth lobes hairy outside; disc 5lobed. Infructescence axis 1530 cm long, stout, on drying finely grooved, scarred by fallen
flower stalks. Fruit a drupe, ovoid, 1.53 x0.82 cm; stalk stout, c. 2.5 cm long; perianth
lobes sometimes persistent.
Vernacular names. Sarawaklimau antu (Iban), sabong ambok (around Limbang).
Distribution. India, Sri Lanka, Thailand, Indo-China, Peninsular Malaysia, Borneo and the
Philippines. In Sabah and Sarawak, widespread; also in Brunei.
Ecology. Common on a variety of soils, from lowland to 1000 m, sometimes in swamp,
kerangas and limestone forest.
262
SAPINDACEAE
F. Adema, P.W. Leenhouts & P.C. van Welzen
With contributions by M. Davids (Paranephelium), J. van Dijk (Dictyoneura),
B. Etman (Rhysotoechia), R.W.J.M. van der Ham (Mischocarpus),
H. Turner (Arytera), and M. Vente (Harpullia)
Rijksherbarium/Hortus Botanicus,
Leiden, The Netherlands
Jussieu, Gen. Pl. (1789) 246 (Sapindi); King, J. As. Soc. Beng. 65, 3 (1896) 419; Merrill, EB (1921) 357; Ridley, FMP 1
(1922) 487; Radlkofer in Engler, Pflanzenr. 98 (19311934) 1; Masamune, EPB (1942) 424; Backer & Bakhuizen f., FJ 2
(1965) 130; Anderson, CLST (1980) 310; Whitmore & Tantra, CLS (1986) 212; Corner, WSTM 3rd ed. 2 (1988) 673; Yap,
TFM 4 (1989) 434; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 310; Adema et al., FM 1, 11 (1994) 419.
Trees, shrubs, or lianas, or rarely herbaceous climbers; monoecious, rarely dioecious or

polygamous. Indumentum usually of solitary simple hairs, sometimes also of two-branched
hairs or stellate bundles of hairs, in several genera scale-hairs present (then the young parts,
buds and inflorescences viscid). Leaves spirally arranged, rarely opposite or whorled,
simple, biternate, digitate, pinnate or bipinnate; true stipules usually absent, pseudostipules
(reduced leaflets at the base of the leaf stalk) sometimes present. Leaflets alternate to
opposite, symmetric to distinctly asymmetric, entire or dentate to serrate or crenate.
Inflorescences axillary, pseudoterminal, terminal or borne on leafless branches, corymbose,
paniculate, racemose, thyrsoid, or umbellate; bracts and bracteoles present. Flowers usually
unisexual, rarely bisexual, radially or bilaterally symmetrical; sepals 4 or 5, rarely more, free
or almost totally connate, equal to distinctly unequal, and then the outer 1 or 2 much smaller
than the inner three, herbaceous to petaloid, imbricate, valvate or apert (fringing or enclosing
a narrow calyx opening); petals absent or 26, free, usually clawed, often with 1 or 2 scales
or auricles (inrolled margins), scales crested or not; disc complete or interrupted, lobed or
annular to subannular, rarely with appendages or an erect (tubular) rim; stamens 510(74),
usually 8, inserted within the disc, often exserted in male flowers, filaments glabrous or hairy,
anthers basifixed or rarely dorsifixed, introrse or latero-introrse, in female flowers present as
staminodes with non-opening anthers; ovary superior, 13(8)-locular, lobed or not, style
usually apical, rarely inserted between the lobes, stigma entire with 13 lines or grooves of
stigmatic papillae, or 13-lobed, in male flowers rudimentary; ovules 1 or 2 per locule,
ascending, anatropous, campylotropous or amphitropous. Fruits capsular or drupaceous, or
consisting of 2 or 3 samaras, when capsular usually loculicidal, rarely septicidal or
septifragal (dehiscing with the valve surface separating from the septa or locule partitions).
Seeds globose to obovoid, sometimes compressed, often with an arillode or a sarcotesta;
endosperm absent; embryo usually thick, straight, sometimes sigmoid or convolute,
cotyledons above each other to laterally beside each other.
263
Distribution. About 150 genera with c. 2000 species, widespread in tropical and subtropical
regions of the world, especially well represented in South America. In Sabah and Sarawak,
23 genera with 62 species, most of which are trees or shrubs.
Ecology. Primary or secondary rain forest, forest edges, shrublands, coastal vegetation, often
along rivers or road sides; in everwet or seasonal climates; mainly lowlands, but also
mountains to 3600 m; on all kinds of soil.
Uses. The wood of several species is used for timber (Klaassen, FM 1, 11 (1994) 425).
Various species are used in medicine, as vegetables, as soap or fish poison. However,
Sapindaceae are more important as a source of edible fruits and seeds. The juicy arillode or
sarcotesta is particularly appreciated. Of economic importance, and hence widely cultivated,
are Dimocarpus longan (longan), Litchi chinensis (litchi, lychee) and Nephelium lappaceum
(rambutan) (PROSEA 2 (1992) 146, 191, 233; van Welzen, Lamb & Wong, Nature
Malaysiana 13 (1988) 10).
Taxonomy. The last survey of the family as a whole was made by Muller & Leenhouts (in
Ferguson & Muller (eds.), The evolutionary significance of the exine (1976) 407). The
system adopted there follows mostly the system developed by Radlkofer (l.c.). Important
changes made by Muller & Leenhouts are: (1) combination of tribes Aphanieae and
Lepisantheae; (2) discarding the division of the Eusapindaceae into Nomophyllae and
Anomophyllae by proposing a more informal grouping of three groups A, B and C; and (3)
the Dodonaeoideae (formerly Dyssapindaceae) are considered an assemblage of relicts, and
the Sapindoideae (formerly Eusapindaceae) as more homogenous and derived. An
enumeration of the subfamilies, tribes and genera occurring in Sabah and Sarawak is given
below.
Subfam. Dodonaeoideae
Tribe Dodonaeeae : Dodonaea
Tribe Doratoxyleae : Ganophyllum
Tribe Harpullieae : Harpullia
(Tribes not represented : Cossinieae, Koelreuterieae)
Subfam. Sapindoideae
Group A
Tribe Lepisantheae : Glenniea, Lepisanthes, Zollingeria
Tribe Melicocceae : Tristiropsis
(Tribe not represented
: Sapindeae)
Group B
Tribe Cupanieae
: Arytera, Dictyoneura, Guioa, Mischocarpus,

Paranephelium, Rhysotoechia, Trigonachras
Tribe Nephelieae : Alectryon, Cubilia, Dimocarpus, Litchi, Nephelium,
Pometia, Xerospermum
(Tribe not represented
: Schleichereae)
Group C
Tribe Paullinieeae : Cardiospermum
Tribe Thouinieae : Allophylus
264
SAPINDACEAE (ADEMA, LEENHOUTS & VAN WELZEN)
Key to genera
(based on flowering material and vegetative characters)
1. Herbaceous or woody climbers. Leaves biternate. Inflorescences with basal tendrils...........

Cardiospermum L.
Sp. Pl. (1753) 366; Radlkofer in Engler, Pflanzenr. 98 (1932) 370; Leenhouts, FM 1,
11 (1994) 483.
Inflorescences axillary, thyrsoid. Flowers unisexual, obliquely bilaterally
symmetrical; sepals 4, free, imbricate, outer 2 smaller; petals 4; disc a gland at the
base of every petal; ovary 3-locular, with a short style and a 3-lobed stigma; ovule 1
per locule. Fruits capsular, 3-lobed, inflated, 3-locular, septicidal, papyraceous. Seeds
with a cordate hilum.
Twelve species, tropical and subtropical America, one species extending to Africa;
one species in Sabah and Sarawak (C. halicacabum), which is a worldwide tropical
and subtropical weed.
Trees or shrubs. Leaves simple, unifoliolate, pinnate, bipinnate or digitate. Inflorescences
without basal tendrils...........................................................................................................2
2. Leaves bipinnate..........................................................................................19. Tristiropsis
Leaves simple, unifoliolate, paripinnate, imparipinnate, or digitate....................................3
3. Leaves imparipinnate...........................................................................................................4
Leaves simple, unifoliolate, paripinnate or digitate.............................................................5
4. Pseudostipules present. Petals shorter than the sepals..........11. Lepisanthes (in part)
Pseudostipules absent. Petals longer than the sepals.............................15. Paranephelium
5. Scale-hairs present (rarely only visible in inflorescences), often also with solitary simple
hairs. Young parts viscid (inflorescence of Lepisanthes fruticosa, race glandulosa may
be sticky). Petals absent.......................................................................................................6
Hairs simple, solitary, or two-branched (T-hairs in Litchi chinensis), or in stellate tufts
(stellate hairs). Young parts not viscid. Petals absent or present......................................8
6. Leaves simple. Ovules 2 per locule.......................................................................................
Dodonaea Miller
Gard. Dict. ed. 4, 1 (1754); King l.c. 450; Merrill l.c. (1921) 362; Ridley l.c. (1922)
509; Radlkofer in Engler, Pflanzenr. 98 (1933) 1350; Masamune l.c. 426; Backer &
Bakhuizen f. l.c. 141; Anderson l.c. 310; Corner l.c. 678; Yap l.c. 440; Leenhouts l.c.
(1994) 522.
Sixty five species, mainly in Australia, three in Malesia.
Inflorescences axillary and terminal, thyrsoid. Flowers bisexual or unisexual, radially
symmetrical; sepals 4 or 5, free, valvate , all equal; petals absent; disc absent in male
flowers, in the form of a slight swelling around the gynophore in female and bisexual
flowers; stamens 515; ovary 23-locular, style and stigma longer than the ovary,
hardly to deeply divided at apex; ovules 2 per locule. Fruit a septifragal capsule, with
3 wings. Seeds subglobose, attached to a swollen funicle.
One species, a pantropical shrub (D. viscosa), present in Sabah and Sarawak; in
coastal vegetation, along the beach or on limestone.
265
Leaves paripinnate. Ovules 1 or 2 per locule.......................................................................7

7. Leaflets subentire to crenate. Sepals free, petaloid. Stigma grooved. Ovules 1 per
locule.............................................................................................................5. Dictyoneura
Leaflets entire. Sepals connate up to halfway, not petaloid. Stigma lobed. Ovules 2 per
locule..........................................................................................................7. Ganophyllum
8. Leaves digitate, 15-foliolate. Flowers bilaterally symmetrical, with 4 petals......................
.........................................................................................................................2. Allophylus
Leaves paripinnate, if unifoliolate then flowers radially symmetrical, usually with 5 petals
(Lepisanthes senegalensis) or without petals (Glenniea thorellii).......................................9
9. Lower side of leaflets with exposed glands.......................................................................10
Lower side of leaflets without exposed glands..................................................................14
10. Leaves with pseudostipules.................................................................16. Pometia (in part)
Leaves without pseudostipules..........................................................................................11
11. Sepals almost totally connate, apert. Stamens 5, filaments glabrous or with few
hairs.........................................................................................................4. Cubilia (in part)
Sepals free or basally connate, usually slightly imbricate. Stamens usually (6)8(10),
filaments mostly hairy.......................................................................................................12
12. Ovary 3-locular, smooth..............................................................18. Trigonachras (in part)
Ovary 2-, rarely 3-locular, warty........................................................................................13
13. Leaf with 12 pairs of leaflets, exceptionally unifoliolate or with 3 pairs of leaflets. Hairs
if present mainly in stellate tufts. Petals absent or present. Disc
hairy...................................
........................................................................................................6. Dimocarpus (in part)
Leaf with 17 pairs of leaflets. Hairs if present solitary, simple. Petals present. Disc
glabrous or (exceptionally) hairy...............................................20. Xerospermum (in part)
14. Hairs often mainly in stellate tufts, mixed with solitary simple ones.................................15
Hairs solitary, simple, exceptionally two-branched (Litchi chinense), if few small tufts of
hairs present then leaves with pseudostipules (Pometia)...................................................17
15. Sepals free. Petals always 5............................................................................10. Harpullia
Sepals connate at base. Petals absent or 15(6)................................................................16
16. Sepals imbricate in bud. Petals absent or present. Stamens (6)8(10), filaments usually
hairy. Ovary tuberculate..................................................................6. Dimocarpus (in part)
Sepals valvate to slightly imbricate in bud. Petals absent. Stamens 6 or 7, filaments
glabrous. Ovary smooth.....................................................................................8. Glenniea
17. Leaves with pseudostipules...............................................................................................18
Leaves without pseudostipules..........................................................................................19
18. Sepals free, imbricate in bud, outer 1 or 2 distinctly smaller than inner 3. Stamens (4) 8
(10)..............................................................................................11. Lepisanthes (in part)
266
Sepals connate up to halfway, valvate in bud, slightly unequal. Stamens 5 (or 6)..............................
............................................................................................................................16. Pometia (in part)
19. Sepals free, imbricate in bud, outer 1 or 2 distinctly smaller than the inner three...........................20
Sepals free or to almost completely connate, apert, valvate or slightly imbricate in bud, usually all
equal, rarely slightly unequal...........................................................................................................22
20. Leaflets concolorous, yellowish green (or green) when dry. Petal scales absent................................
.................................................................................................................................17. Rhysotoechia
Leaflets usually discolorous, brown or greyish (green) when dry, rarely concolorous, yellowish
green
(Lepisanthes
falcata
subsp.
borneensis).
Petal
scales
or
auricles
present..............................................................................................................................................21
21. Petals with 2 scales. Stigma pyramidal, grooved...................................................................9. Guioa
Petals with 1 scale, or with auricles, rarely with a hairy ridge. Stigma usually capitate, globose or
dome-shaped, rarely elongated, grooved.....................................................11. Lepisanthes (in part)
22. Petals usually absent, rarely up to 3..................................................................................................23
Petals present, usually 5, sometimes reduced...................................................................................26
23. Ovary smooth, 3-locular, not lobed, rather long-stalked...........................13. Mischocarpus (in part)
Ovary warty or smooth, 12(rarely 3)-locular, lobed, sessile or short-stalked.................................24
24. Two-branched hairs often present. Leaflets mostly opposite or subopposite. Sepals connate,
apert.....................................................................................................................................12. Litchi
All hairs simple. Leaflets alternate to opposite. Sepals free or connate, valvate or slightly
imbricate in bud...............................................................................................................................25
25. Leaflets opposite to rarely alternate. Ovary smooth.......................................................1. Alectryon
Leaflets alternate to rarely opposite. Ovary warty........................................14. Nephelium (in part)
26. Petals without scales or auricles. Ovary warty................................................................................27
Petals with scales or auricles. Ovary smooth or warty....................................................................29
27. Sepals almost totally connate, calyx urceolate with a narrow mouth and minute lobes. Stamens
5.............................................................................................................................4. Cubilia (in part)
Sepals free or connate to halfway, calyx flat or cup-shaped with a wide mouth and distinct lobes.
Stamens 410..................................................................................................................................28
28. Leaflets opposite, lower surface not glaucous, with flat orbicular glands, without domatia.
Stamens 8, exceptionally 7 or 9. Stigma grooved..................................20. Xerospermum (in part)
Leaflets alternate to rarely opposite, lower surface mostly glaucous, without glands, with domatia.
Stamens 410. Stigma lobed........................................................................14. Nephelium (in part)
29. Petals with 1 large, sometimes deeply cleft scale. Ovary warty...................14. Nephelium (in part)
Petals with 2 scales or auricles. Ovary smooth...............................................................................30
30. Twigs white or silvery grey. Ovary 1-locular, septa incomplete................................21. Zollingeria
Twigs light to dark brown or grey to black. Ovary 25-locular, septa complete............................31
267
31. Intercostal veins rather densely reticulate. Petals usually with auricles. Ovary short- to
long-stalked. Stigma distinctly lobed......................................................13. Mischocarpus (in part)
Intercostal veins more-or-less laxly reticulate. Petals with 2 scales. Ovary sessile. Stigma
grooved or slightly lobed................................................................................................................32
32. Lower surface of leaflets without small, round glands, but with domatia. Petals shorter than, rarely
as long as, the sepals; petal scales shorter than the petals. Ovary 2(3)locular.................................................................................................................................3. Arytera
Lower surface of leaflets with small, round glands, without domatia. Petals as long as to longer
than the sepals; petal scales nearly as long as the petals. Ovary 3-locular..........................................
..................................................................................................................18. Trigonachras (in part)
Key to genera
(based on fruiting material and vegetative characters)
1. Herbaceous or woody climbers. Leaves biternate. Inflorescences with basal
tendrils........... .......................................................................Cardiospermum (see notes in
previous key)
Trees, shrubs, or occasionally woody climbers. Leaves simple, unifoliolate, digitate,
pinnate, or bipinnate. Inflorescence without tendrils...........................................................2
2. Leaves bipinnate...........................................................................................19. Tristiropsis
Leaves simple, unifoliolate, digitate or pinnate...................................................................3
3. Leaves all simple, unifoliolate, digitate, or imparipinnate...................................................4
Leaves paripinnate or pseudo-imparipinnate (with one of the final two leaflets reduced
and
vestigial);
occasionally
also
some
leaves
unifoliolate............................................................................................................................7
4. Fruits winged. Leaves simple....................................Dodonaea (see notes in previous key)
Fruits
not
winged.
Leaves
unifoliolate,
digitate
or
imparipinnate........................................................................................................................5
5. Leaves (1)3(5)-digitate; rachis not winged. Fruits globose to obovoid, coriaceous, at
most 0.41.3 x 0.3-0.8 cm...............................................................................2.
Allophylus
Leaves unifoliolate or imparipinnate. Fruits either slightly lobed and coriaceous, or
subglobose and very woody, larger than 1 1.5 cm; if about 1 1.5 cm then rachis winged
.............................................................................................................................................6
6. Pseudostipules present or absent; rachis winged or not. Fruits slightly lobed, smooth,
coriaceous; fruit wall glabrous inside............................................11. Lepisanthes (in part)
Pseudostipules absent; rachis not winged. Fruits subglobose, spiny to warty, woody; fruit
wall pilose inside.....................................................................15. Paranephelium (in part)
7. Fruits winged................................................................................................21. Zollingeria
Fruits without wings............................................................................................................8
8. Fruits warty to densely spiny or densely scaly.....................................................................9
Fruits smooth to rugose when dry......................................................................................14
268
9. Fruit wall hairy inside. Seeds without sarcotesta or arillode....15. Paranephelium (in part)
Fruit wall glabrous inside. Seeds with sarcotesta or arillode..............................................10
10. Seeds with sarcotesta.........................................................................................................11
Seeds with arillode.............................................................................................................12
11. Leaf with 15(18) pairs of leaflets, usually papillose below (dull), often minutely
sericeous; without glands, often with domatia. Sepals free to connate more than halfway.
Spines and other forms of appendages on fruit usually longer than
broad.............................
.........................................................................................................14. Nephelium (in part)
Leaf with 12(3) pairs of leaflets, smooth below (more or less shiny), glabrous or at
most hairy on midrib and lateral veins, not sericeous; with scattered glands, without
domatia. Sepals free or only basally connate. Spines and other forms of appendages on
fruit broader than long.................................................................................................20.
Xerospermum
12. Arillode covering the lower half of seed only. Fruit dehiscent. Indumentum of simple
hairs
only.......................................................................................................................4.
Cubilia
Arillode almost completely covering the seed. Fruit indehiscent. Indumentum of tufted, 2branched or simple hairs....................................................................................................13
13. Indumentum often partly or mainly of dense stellately tufted hairs. Glands present on
lower side of leaflets near axils of lateral veins (seldom absent in all leaflets). Seeds about
as long as broad...............................................................................6. Dimocarpus (in part)
Indumentum of solitary, simple or 2-branched hairs. Glands absent. Seeds longer than
broad......................................................................................................................12. Litchi
14. Seeds without sarcotesta or arillode. Placenta neither thickened nor cup-shaped..............15
Seeds with sarcotesta (sometimes as a narrow basal ring around hilum) and/or arillode.
Placenta thickened and cup-shaped...................................................................................20
15. Leaves with hairs and glandular scales (distinct under the microscope)....7. Ganophyllum
Leaves
without
glandular
scales,
at
most
with
hairs
only.....................................................................................................................................16
16. Fruit wall hairy inside........................................................................................................17
Fruit wall glabrous inside...................................................................................................19
17. Fruits stalked, obovoid, dehiscing loculicidally into 3 equal valves; 3-locular; fruit wall
thick, fleshy; hilum covering less than lower one-third of the seed..........18. Trigonachras
Fruits sessile, ellipsoid, shortly obovoid to subglobose, not dehiscing to dehiscing into 3
or 4 usually unequal valves or tearing apart at random; 2- or 3-locular; either fruit wall
thick and hilum covering up to lower three-quarters of the seed, or fruit wall thin and
hilum covering less than lower one-third of the seed.........................................................18
18. Indumentum at most of short hairs. Leaflets entire to serrate. Fruits capsular; fruit wall
2.512 mm thick; hilum covering up to lower three-quarters of the seed..............................
.................................................................................................15. Paranephelium (in part)
Indumentum often of hairs of more than 5 mm long. Leaflets entire. Fruits drupaceous;
fruit wall less than 2 mm thick; hilum covering less than lower one fifth of the
seed................................................................................................11. Lepisanthes (in part)
269
19. Leaf with 1 to more than 40 pairs of leaflets; pseudostipules present or absent; rachis
winged or not. Outer 1 or 2 sepals smaller than inner ones. Fruits 23(4)-locular,
glabrous or pilose, less than 5 cm long; fruit wall usually thin, sometimes thick and
fleshy.............................................................................................11. Lepisanthes (in part)
Leaf with 16 pairs of leaflets; pseudostipules absent; rachis not winged. Sepals all equal.
Fruits 2-locular, glabrous, either less than 2 cm long and fruit wall rather thin or more
than 6 cm long and fruit wall very thick and fleshy............................................8. Glenniea
20. Indumentum of stellately tufted hairs besides solitary hairs..............................................21
Indumentum of solitary hairs or plant glabrous.................................................................22
21. Fruits lobed, loculicidally dehiscent into valves; hilum small, covering less than one sixth
of the seed.......................................................................................................10. Harpullia
Fruits not lobed, breaking up irregularly; hilum very large, covering up to half of the
seed.................................................................................................6. Dimocarpus (in part)
22. Pseudostipules present (sometimes reduced or early caducous, but then leaving scars).
Fruits lobed, but seemingly 1-locular and not lobed by abortion; exocarp thin, hard,
mesocarp thick, juicy when fresh......................................................................16. Pometia
Pseudostipules absent. Fruits lobed or not; carpels usually thin to thick, coriaceous to
woody................................................................................................................................23
23. Fruit wall completely hairy or papillose inside, or only hairy along the suture.................24
Fruit wall glabrous inside except sometimes for the lower part of the axis.......................27
24. Fruit lobes each dehiscing by an irregular transverse break..............................1. Alectryon
Fruits dehiscing loculicidally.............................................................................................25
25. Fruit wall only hairy along the suture inside.........................................................3. Arytera
Fruit wall nearly completely hairy or papillose inside.......................................................26
26. Fruit on a slender stalk, inside without an extra fleshy layer. Aril with pseudofunicle
....................................................................................................13. Mischocarpus (in part)
Fruit hardly stalked, inside with an extra fleshy layer. Aril without pseudofunicle...............
......................................................................................................................5. Dictyoneura
27. Placenta thickened below the seed, more-or-less cup-shaped. Seeds not covered with
sarcotesta or arillode.................................................................................17. Rhysotoechia
Placenta seldom thickened below the seed. Seeds (partly) covered with sarcotesta and/or
arillode..............................................................................................................................28
28. Fruits 1-locular. Seeds partly to completely covered with a sarcotesta..................................
.........................................................................................................14. Nephelium (in part)
Fruits 3-locular. Seeds partly to completely covered with an arillode...............................29
29. Fruits lobed. Stigma a pyramidal remnant without recurved lobes on top of the fruit.
Sepals dimorphous, free..........................................................................................9. Guioa
Fruits not lobed. Stigma with recurved lobes. Sepals all equal, usually connate at the base
....................................................................................................13. Mischocarpus (in part)
270
1. ALECTRYON Gaertn.
(Greek, alektruon = rooster; the crested fruits)
P.W. Leenhouts
Fruct. Sem. Pl. 1 (1788) 216, pl. 46; Radlkofer in Engler, Pflanzenr. 98 (1933) 983; Backer & Bakhuizen f., FJ 2 (1965) 139;
Leenhouts, FM 1, 11 (1994) 450.
Trees or shrubs, dioecious or sometimes monoecious. Indumentum of solitary simple hairs

only. Leaves paripinnate, each with 18 pairs of leaflets; petiole and rachis not winged; true
stipules absent, lowermost pair of leaflets sometimes stipule-like. Leaflets variably hairy to
glabrous, lower surface not papillose; base often oblique, margin entire or serrate, dentate or
crenate; lateral veins ending free or the upper ones looping and joining near the margin.
Inflorescences axillary, pseudoterminal, or borne on leafless branches, thyrses, panicles or
racemes. Flowers unisexual or sometimes bisexual, radially symmetrical; sepals 4, 5 (or 6),
about halfway to nearly completely connate, all equal, hardly or not petaloid; petals absent or
4 or 5, about as long as or shorter than the sepals, short-clawed, with 2 scales, without a crest;
disc complete, annular or more-or-less lobed, without appendages, glabrous; stamens (5)8,
exserted in male flowers, anthers basifixed, the base usually deeply cleft, latrorse or laterointrorse; ovary sessile or short-stalked, (1)24(5)-locular, style apical, columnar, mostly
shorter than the ovary, stigma grooved or with reflexed lobes; ovules 1 per locule. Fruits
sessile or short-stalked, 1(or more)-lobed, capsular, dehiscing by an irregular transverse
break on each lobe or along the septum; fruit wall smooth or slightly warty, hairy or late
glabrescent outside, glabrous or hairy inside. Seeds black, partly covered with a red
sarcotesta.
Distribution. About 25 species; in Malesia, Australia, New Zealand, New Caledonia, the
New Hebrides, the Solomon Is., Fiji, Samoa, and the Sandwich Is. One species in Sabah.
Ecology. Often at forest edges, on river banks, in coastal vegetation, on limestone; in the
lowlands but sometimes also on mountains. The seeds, with contrasting light yellowish
greenish capsules, red sarcotesta, and shiny black testa, are probably mainly dispersed by
birds.
Uses. Some species are sources of good timber.
Alectryon glaber (Blume) Radlk.
Fig. 1.
(Latin, glaber = smooth; the fruits lacking in crests)
Sapind. Holl.-Ind. (1879) 14, in Engler l.c. (1933) 993; Leenhouts l.c. (1994) Sapind. Holl.-Ind. (1879) 14, in Engler l.c.
(1933) 993; Leenhouts l.c. (1994) 456, fig. 5f. Basionym: Spanogheaglabra Blume l.c. (1847) 174. Type: Spanoghe 52,
Lesser Sunda Is. (L).
Shrub or small tree to 30 cm diameter; stem slightly fluted, with rather inconspicuous thick
buttresses. Bark smooth, pale brown to ash-grey. Leaves each with 25(7) pairs of leaflets;
petioles 18.5 cm long, (0.5)12 mm thick. Leaflets opposite to alternate, parchment-like to
271
Fig. 1. Alectryon glaber. A, fruiting leafy twig; B, fruit. (From Alston 16075.)
272
chartaceous, glabrous to slightly hairy on the midrib and lateral veins beneath, exceptionally
subtomentose; ovate to elliptic, c, 414 x 15.5 cm; base symmetric to asymmetric, acute to
rounded, somewhat attenuate, margin slightly serrate, dentate, or crenate in the upper half,
or rarely entire, apex rounded to acute; midrib slightly raised above, lateral veins 0.52 cm
apart, ending free or in marginal teeth, or looping and joining near the margin, slightly raised
on both sides or flat above, intercostal veins and veinlets laxly to minutely reticulate, slightly
raised on both sides, or indistinct beneath; petiolules 18 mm long. Inflorescences to 9 cm
long; unbranched or with branches to 4 cm long, few-flowered, hairy; peduncles 0.752.5 cm
long; pedicels 11.5 mm long. Flowers: sepals c. 1 mm high, slightly connate, hairy inside;
petals absent; stamen filaments short, anthers c. 1.2 mm long; ovary 2(3)-locular with a
lobed stigma. Fruits 2-lobed and then often cordate (or only 1 lobe developed), keeled and
shouldered to slightly grooved, densely to sparsely fulvous or ferruginous tomentose or
tomentulose, the lobes almost globose, 0.910 x 750.9 cm; fruit wall 0.51 mm thick.
Distribution. E Java, the Lesser Sunda Is., N Borneo (Sabah, one specimen), the Philippines,
Sulawesi, Ceram, and the Kai Is.
Notes. In Sabah, once recorded (Berhaman AB21) on coral limestone on Balambangan Is.
2. ALLOPHYLUS L.
(Greek, allos = another, phulon = tribe; its distinction from other known tribes)
P.W. Leenhouts
Sp. Pl. (1753) 348; Merrill, EB (1921) 357; Ridley, FMP 1 (1922) 488; Radlkofer in Engler, Pflanzenr. 98 (1932) 455;
Masamune, EPB (1942) 424; Backer & Bakhuizen f., FJ 2 (1965) 133; Leenhouts, Blumea 15 (1968) 301, FM 1, 11 (1994)
459; Anderson, CLST (1980) 310; Corner, WSTM 3rd ed. 2 (1988) 670; Yap, TFM 4 (1989) 436.
Shrubs, small trees, or sometimes woody climbers, monoecious or dioecious. Indumentum

sometimes of stellately tufted hairs. Leaves digitate, (1)3(5)-foliolate. Inflorescences
axillary, usually either simple or composed of a few long and slender, racemose thyrses,
sometimes paniculate. Flowers unisexual, obliquely bilaterally symmetrical; sepals 4, free,
imbricate in bud, outer 2 distinctly narrower than the inner ones; petals 4, nail-shaped to
spathulate, inside above the claw with a 2-lobed, usually bearded scale; disc more-or-less
interrupted abaxially, mostly lobed or modified into glands in front of the petals, in female
flowers sometimes saucer-shaped; stamens 8 (rarely fewer), all of about the same length,
exserted in male flowers; ovary deeply 2(3)-lobed, 2(3)-locular, ovule 1 per locule, style 1,
inserted between the lobes, more or less deeply 2(3)-branched, the branches exceptionally
also forked at the apex. Fruits drupaceous, mostly of 1 mericarp only, globose to obovoid,
thin-walled, mostly almost glabrous when ripe. Seeds without arillode.
Distribution. Probably monotypic, though up to 250 species may be accepted.
Circumtropical, in some regions penetrating into the subtropics.
273
Allophylus cobbe (L.) Raeuschel
Fig. 2.
(after the Sri Lankan plant-name kobbae)
Nomencl. ed. 3 (1779) 108; Merrill l.c. (1921) 357; Radlkofer in Engler l.c (1932) 594; Masamune l.c. 424, 425; Anderson
l.c. 310; Corner l.c. 677; Backer & Bakhuizen f. l.c. 133. Basionym: Rhus cobbe L., Sp. Pl. (1753) 267. Type: Herb.
Hermann vol. 2, fol. 46 (Kobbae), Sri Lanka (BM). Synonyms: A. javensis (Blume) Blume l.c. (1847) 126, Merrill l.c.
(1921) 357, Masamune l.c. 424; A. sumatranus Blume l.c. (1847) 132, Merrill l.c. (1921) 357, Masamune l.c. 425; A.
timorensis (DC.) Blume l.c. (1847)
130, Merrill l.c. (1921) 357, Masamune l.c. 425; A. scandens Ridl., J. Str. Br. Roy. As. Soc. 75 (1917) 26, Merrill l.c. (1921)
357, Masamune l.c. 425; A. obliquus Radlk. in Merrill, PEB (1929) 173, Masamune l.c. 424; A. glaber (Roxb.) Radlk. in
Engler l.c. (1932) 566, Masamune l.c. 424; A. racemosus (L.) Radlk. in Engler & Prantl, Nat. Pflz. Fam. 3, 5 (1895) 312, in
Engler l.c. (1932) 568, Merrill l.c. (1921) 357, Masamune l.c. 425; A. ternatus (Forst.) Radlk. in Engler l.c. (1932) 572,
Masamune l.c. 425.
Shrub or treelet, rarely a tree to 25 m high, 30 cm diameter, or a climber (not in Sabah and
Sarawak). Twigs glabrous except for the sparsely appressed hairy terminal bud or more or
less persistently thinly to densely covered with often stellately tufted, appressed to patent,
short, fulvous hairs. Leaves: petioles terete, rarely quadrangular, mostly flattened to grooved
above especially at apex and base, 4.520 cm long, indumentum usually similar to that of the
twig at earlier stage. Leaflets membranous, fleshy or coriaceous, on drying greenish to
greyish or dark-brown and dull to shiny above, light green to reddish brown and dull
(sometimes even glaucous) to shiny beneath, glabrous (often with a distinct gland) to densely
velvety (especially beneath) and often bearded in the axils of lateral veins and sometimes also
of the intercostal veins; elliptic or oblong, rarely lanceolate, lateral ones often ovate,
terminal ones sometimes obovate, 2.535 x 1.522 cm (terminal ones slightly to distinctly
larger than the lateral ones); base cuneate to rounded, often slightly decurrent, margin entire
to serrate, crenate or dentate, apex more or less attenuate-acuminate; midrib hardly
prominent to keeled above, lateral veins 615 per side, looping and joining near the margin
or not, intercostal veins usually rather inconspicuous above, hardly conspicuous to prominent
beneath; petiolules absent to 25 mm long, indumentum like that of petioles. Inflorescences to
40 cm long, few- to many-flowered, with umbel-like dichasia or sometimes comb-like
cincinni, glabrous to densely pubescent; peduncles usually about one third to one fifth the
length of rachis, laxly to densely flowered; pedicels absent or short; bracts mostly minute,
sometimes longer than the pedicels, subulate. Flowers often solitary in the upper part of an
inflorescence; sepals 12.5 x 0.82 mm, green to whitish, entire to denticulate, mostly ciliate,
glabrous to sparsely appressed short-hairy (mainly in the central part) outside; petals 12.2
mm long, white, almost glabrous to densely woolly along the entire margin, blade entire to
bilobed, claw about two third of the petal length, scale very small to nearly equalling the
blade, glabrous to densely woolly along the entire margin, often bearded; disc 0.20.8 mm
high, glabrous or puberulous, orange; anthers c. 0.5 mm long; ovary smooth or rough,
glabrous or minutely stellately tufted-hairy, or sparsely to densely pilose by appressed, long,
stiff hairs, style 11.5 mm long, glabrous or pilose up to the stigmatic lobes. Fruits globose
(smaller ones) to obovoid and narrowed at the base (larger ones), 4.512.5 x 3.58 mm,
smooth to slightly rugose, red turning to brown or black when dry, somewhat pulpy and
almost glabrous when ripe.
Vernacular names. Sabahbonbongan (Murut), embuakat, gamperut (Dusun), kalasi
(Bajau), kerimbau (Murut), mambahino, sasah (Bajau East Coast), sisipen pelandok (Dusun),
tukil-tukil (Brunei Malay). Sarawakbalaro (Kayan), buah penancang (Malay), ising (Iban,
Paloh), kelampu (Iban).
274
Fig. 2. Allophylus cobbe. A, fruiting leafy twig; B, male flower; C, petal from inside; D, female flower; E, fruit. (A and E
from Forbes 2557, BD from Maxwell 77-375.)
275
Distribution. Throughout the range of the genus. Common in Sabah and Sarawak; also in
Ecology. In different types of primary and secondary forests, from lowland to c. 1300 m (Mt.
Kinabalu), on various soil types. Flowering and fruiting throughout the year. The flowers are
visited by bees; the fruits are dispersed by birds (Docters van Leeuwen, Trop. Natuur 21
(1932) 142).
Uses. The wood is reported to be very hard, but not very durable. It is mainly used for
temporary and indoor structures, for making canes, hilts, and as firewood. The pulped leaves,
or an extraction or decoction of them, as well as a decoction of the roots and bark, are used in
medicine against stomach-ache and fever. The fruits, though a bit sour, are edible.
Taxonomy. A. cobbe is a very variable species, with wide distribution and broad ecological
amplitude. The species probably comprises a large number of local races. Leenhouts (l.c.
1967, 1994) concluded that all Malesian specimens examined belong to a single species.
However, on a local scale it may be possible to distinguish the more important races
(varieties or forms). For Borneo, and especially Sabah and Sarawak, no attempt has been
made to subdivide the species into infraspecific taxa.
3. ARYTERA Blume
(Greek, aruter = ladle; the shape of the fruit-lobes)
H. Turner
Rumphia 3 (1847) 169; Merrill, EB (1921) 361; Ridley, FMP 1 (1922) 507; Radlkofer in Engler, Pflanzenr. 98 (1933) 1268;
Masamune, EPB (1942) 425; Backer & Bakhuizen f., FJ 2 (1965) 140; van der Ham, Blumea 23 (1977) 289; Anderson,
CLTS (1980) 310; Reynolds, Austrobaileya 2 (1985) 158; Yap, TFM 4 (1989) 438; Turner, Blumea 38 (1993) 144, FM 1, 11
(1994) 467.
Trees, rarely shrubs. Indumentum of solitary, simple hairs. Leaves paripinnate; petioles
subterete, pulvinate; rachis subterete. Leaflets subopposite; petiolules usually consisting of
pulvinus only, usually 1-grooved; domatia usually present. Inflorescences axillary to
pseudoterminal or borne on leafless twigs, thyrsoid; bracts and bracteoles triangular, hairy
outside, usually glabrous inside; pedicels hairy. Flowers seemingly hermaphrodite; calyx 5dentate to 5-partite, teeth all equal to slightly unequal, apert to narrowly imbricate, margin
entire; petals (2)5(6), slightly longer or shorter than the calyx, scales present or rarely
absent, free to basally adnate to the margins, not crested; disc complete; stamens (6)8(10),
filaments at least basally pilose, anthers pilose or glabrous; ovary sessile, 23-locular,
smooth, hairy, ovule 1 per locule, stigmas apical, with 2 or 3 stigmatic lines, or with 2 or 3
lobes, the lobes distinctly recurved in fruit. Fruits with 13 well-developed lobes, loculicidal,
short- or long-stalked, lobes laterally not or slightly flattened; endocarp hairy at least on
sutures of carpels. Seeds partly covered by an arillode, consisting of 1 or 2 layers, the outer
or only one soft, yellow, the inner one firm, dark brown when dry.
276
Distribution. About 28 species, from NE India and SE Asia, throughout Malesia, to

Australia, the Solomon Islands and the Pacific Islands. In Sabah and Sarawak, one species.
Arytera litoralis Blume
Fig. 3.
(Latin, litoris = coastal region; its natural habitat)
l.c. (1847) 170; Merrill l.c. (1921) 361; Ridley l.c. (1922) 507; Masamune l.c. 425; Backer & Bakhuizen f. l.c. 140;
Anderson l.c. 310; Turner l.c. (1994) 473. Lectotype (Turner, 1993): Blume 1314, Java, Nusa Kambangan (L).
Tree, rarely shrub, to 40 m tall, 90 cm diameter. Bark smooth, greyish green to reddish to
almost black. Leves each with 14 pairs of leaflets; petioles 1.39.5 cm long; rachis 0.811.5
cm long. Leaflets thin-coriaceous to chartaceous, not to densely punctate; ovate to elliptic or
obovate, 431 x 1.512 cm; base symmetrical or asymmetrical, rounded to slightly attenuate,
apex acuminate to cuspidate, rarely retuse or rounded, acumen retuse, obtuse to rounded;
glabrous above, glabrous to hairy especially on major veins beneath, domatia often pustular,
in the form of large to small pockets or sacs, opening in front (on top); midrib slightly raised
above, concolorous with lamina to reddish brown or yellowish, lateral veins raised below,
indistinctly looping near margin, intercostal veins scalariform to almost reticulate, obscure;
petiolules 214 mm long. Inflorescences basally branched or unbranched; rachis terete to
slightly flattened, 1.517 cm long, hairy when young; first-order branches to 10 cm long;
cymules dichasial or sometimes monochasial, 17-flowered; bracts 0.31.2 mm long;
bracteoles 0.10.6 mm long; pedicels 15 mm long, hairy. Flowers 13.5 mm diameter;
calyx 0.82 mm high, deeply incised, hairy outside, glabrous inside, apex acute to acuminate;
petals triangular to rhomboidal to obovate, 0.52.2 x 0.31.9 mm, usually gradually
decurrent into the claw, margin entire, subpilose, apex obtuse to acute to acuminate, glabrous
to pilose outside, subglabrous to subpilose inside, claw 0.10.4 mm long, scales 0.21.2 mm
long, apex irregulaly broadened, slightly to densely pilose; disc glabrous to hairy; stamens 6
8(10), filaments 24 mm long, pilose, anthers 0.71.1 mm long, straight, pilose, connective
not protruding; ovary 2(3)-locular, style and stigma elongating up to 3 mm in fruit, not to
slightly 2(3)-lobed. Fruits 12(3)-lobed, 736 x 523 mm, smooth to slightly rugose to
verrucose, glabrous to sparsely hairy, lobes 823 x 521 mm, dorsally rounded to slightly
angular; stalk absent or to 3 mm long, slender to broadly cuneate. Seeds ellipsoid to
orbicular, not to slightly flattened laterally, 624 x 519 mm; arillode covering the seed
halfway to completely, dentate to lobed, not to slightly folded towards the base inside, thick
towards the base, coriaceous, 2-layered; cotyledons obliquely superposed to almost
collateral, equal to slightly unequal, upper or lower one larger; hypocotyl and radicle 0.53
mm long, glabrous.
Vernacular names. Sabahampungit (Murut), anging manuk (Kadazan), nunuk-nunuk,
petinag (Sungei Kinabatangan).
Distribution. From NE India (Bay of Bengal) to SE Asia, S China (Hainan), Malesia, and the
Solomon Islands. Common in Sabah and Sarawak; also in Brunei and Kalimantan.
Ecology. In primary and secondary forests on all kinds of soil, lowland to 1500 m. Flowering
and fruiting throughout the year.
277
Fig. 3. Arytera litoralis. A, fruiting twig; B, female flower; C, petal from inside; D, fruits. (A from SAN 95571, BC from
Gibbs 2664, D from van Balgooy 6099.)
278
4. CUBILIA Blume
(from the Philippine plant-name kubili)
P.W. Leenhouts
Rumphia 3 (1847) 100; Radlkofer in Engler, Pflanzenr. 98 (1932) 921; Backer & Bakhuizen f., FJ 2 (1965) 143; Leenhouts,
Blumea 24 (1978) 297, 298, FM 1, 11 (1994) 490; Whitmore & Tantra, CLS
Medium-sized, monoecious trees. Indumentum of solitary, simple hairs. Leaves paripinnate.

Leaflets opposite to alternate, base equal-sided to oblique, margin entire; venation mainly
open but somewhat irregular. Inflorescences terminal or pseudoterminal, thyrsoid or
corymbose; cymules many-flowered; bracts inconspicuous. Flowers unisexual, radially
symmetrical; calyx urceolate, the narrow mouth surrounded by 5 minute lobes, densely
tomentulose on both surfaces, not petaloid; petals 5, included in the calyx, hardly clawed,
margin sometimes slightly inflexed just above the base but without a clear scale; disc
annular, broad, adnate to the receptacle, glabrous; stamens 5 (or 6), hardly exserted,
filaments flattened, glabrous or with a few hairs, anthers adnate, glabrous, introrse; ovary
sessile, 2-locular, divided to near the base, warty and densely hairy, stigma sessile, 2-lobed,
inserted between the ovary lobes, ovules 1 per locule, attached at the base. Fruits 2-parted,
the parts obovoid, loculicidal, glabrous, densely aculeate; fruit wall coriaceous, glabrous
inside. Seeds basally attached, up to about halfway enveloped by a thin-fleshy, more-or-less
entire arillode, hilum large, nearly orbicular.
Distribution. Monotypic; Borneo (Sabah, Sarawak, E Kalimantan), the Philippines,
Sulawesi, and the western Maluku.
Cubilia cubili (Blanco) Adelbert
Fig. 4.
Blumea 6 (1948) 325; Backer & Bakhuizen f. l.c. 143; Leenhouts l.c. (1978) 397, l.c. (1994) 491. Basionym: Euphoria
cubili Blanco, Fl. Philip. (1837) 287, nom. illeg. Neotype (Leenhouts, 1978): Merrill Spec. Blanc. 705, Philipines, Luzon (L;
isoneotypes BO, P).
Tree to 25(50) m tall, 75 cm diameter; buttresses to 6 m high. Bark usually smooth and
reddish, sometimes greyish brown. Leaves each with 35(7) pairs of leaflets; petioles terete
or sometimes slightly flattened above, smooth or thicker ones canaliculate, strongly swollen
at base, 525 cm long, 1.53 mm thick; rachis slightly flattened above, otherwise terete,
sometimes carinate above towards the apex. Leaflets elliptic to ovate, (5)1015(40) x(2.5
)35(10) cm, the upper ones often slightly falcate, chartaceous, often with an exposed gland
in or above the axils of some lateral veins beneath; base rounded and attenuate to acute, apex
obtuse- to acute-acuminate, mucronate or not; midrib slender, slightly raised to sunken above,
mostly sharp-triangular, sometimes more or less rounded beneath, lateral veins 12(4) cm
apart, sometimes a few looping and joining towards the margin, slightly raised above, more
so beneath; intercostal veins mostly feeble, reticulate, mostly faint above, slightly raised
beneath; petiolules 37.5 mm long. Inflorescences to 30 cm long, densely and minutely
appressed brown-hairy, the branches sparsely branched; cymules to 25-flowered; peduncles to
1 cm long; pedicels slender, 34 mm long, slightly thickened towards the apex; bracts
triangular and hardly 0.5 mm long to subulate and 3 mm long. Flowers copper-coloured;
279
Fig. 4. Cubilia cubili. A, flowering leafy twig; B, male flower in longitudinal section; C, female flower; D, fruit; E, open
fruit with seed; F, seed in longitudinal section. (A from FB 1996, BC from PNH 22872, D from NIFS Cel. V/236 =
Waturandang 221, E from Radlkofer in Engler, Pflanzenr. (19321933) f. 22, F from Kostermans 7011.)
280
calyx in male flowers c. 3 x 2 mm, opening c. 1 mm diameter, lobes minute; in female

flowers 1.52 x 34 mm, opening wider than in male, lobes broad-triangular, c. 0.8 mm high;
petals in male flowers elliptic, c. 1 x0.7 mm, in female flower subovate, c. 1.5 x 1.5 mm;
stamen filaments c. 1 x 0.2 mm, anthers c. 0.3 mm long, in female flowers hardly reduced;
ovary c. 1.5 mm high, stigma lobes triangular, dorsoventrally flattened, c. 1 mm long. Fruit
parts 34(5) x 22.5 cm, green to brown, warty with the warts pyramidal to triangular, to 2
mm long; fruit wall c. 0.8 mm thick; endocarp smooth, white. Seeds ovoid-ellipsoid, c. 2.5 x
1.7 cm; testa smooth, shiny dark brown; arillode to 610 mm high.
Distribution. As the genus. Uncommon in Sabah and Sarawak.
Ecology. Primary and secondary forests, apparently mostly on poorly aerated, often basic
soil, often along rivers; mostly at low altitudes, to 600 m. Flowering and fruiting throughout
the year.
Uses. The whitish to light reddish brown timber is used in the Philippines for indoor
construction. The arillode is edible; the seeds, when cooked or roasted, are are said to be
comparable with, but more delicious, than chestnuts.
5. DICTYONEURA Blume
(Greek, dictuon = net, neuron = nerve; the leaflet venation)
J. van Dijk
Rumphia 3 (1847) 163; Merrill, EB (1921) 361; Radlkofer in Engler, Pflanzenr. 98 (1933) 1219; Masamune, EPB (1942)
426; Reynolds, Austrobaileya 2 (1985) 153; J. van Dijk, Blumea 31 (1986) 437, FM 1, 11 (1994) 507.
Shrubs to medium-sized trees, monoecious. Indumentum of simple, solitary hairs and small,
round scales. Leaves paripinnate. Leaflets alternate to opposite, variably hairy or scaly,
domatia absent or pocket-like. Inflorescences axillary, often pseudoterminal, simple and
raceme-like or more-or-less strongly branched; cymules 1few-flowered; bracts and
bracteoles densely hairy, mostly caducous. Flowers unisexual, radially symmetrical; sepals
5(or 6), imbricate in bud, slightly connate at the base, about equal or the inner- and/or
outermost one distinctly smaller, persistent in fruit; petals absent; disc entire, swollen,
tomentose; stamens (4)5(6), exserted in male flowers, filaments usually patently long-hairy
in the basal half, anthers basifixed to halfway dorsifixed; ovary subsessile, 2(3)-locular,
more-or-less hairy, sparsely to densely scaly, style apical, shorter to a little longer than the
ovary, with 2 (or 3) stigmatic lines, ovules 1 per locule, axillary. Infructescences with
thickened axes. Fruits capsular, subsessile, loculicidal; fruit wall granular, inner side more
or less fleshy with a thicker fleshy layer in a narrow to broad strip along the dissepiment and
covering the inside of the valves for 30100%. Seeds ovoid to ellipsoid, 12(3) per fruit;
testa parchment-like or a more-or-less hard shell; sarcotesta cupular on the ventral side and
covering 25100% of the seed, dorsally interrupted by a very narrow to very broad cuneate
cleft; embryo straight, cotyledons superposed, the upper slightly to very much larger than the
lower one.
Distribution. Two to three species; Borneo, Philippines, Sulawesi, Maluku and New Guinea.
In Sabah and Sarawak, one species, represented by one of its two subspecies.
281
Dictyoneura acuminata Blume
(Latin, acuminatus = gradually attenuated; the leaflet apex)
Fig. 5.
l.c. (1847) 163; Merrill l.c. (1921) 361; Radlkofer in Engler l.c. (1933) 1221; Masamune l.c. 426; J. van Dijk l.c. (1986) 441,
l.c. (1994) 508. Lectotype (J. van Dijk, 1986): Muller, s.n., Borneo (L).
subsp. acuminata
Tree, sometimes treelet or shrub to 12(15) m tall, 20(30) cm diameter. Bark greenish or
greyish, smooth; inner bark pale brown. Wood white. Leaves each with 27(11) pairs of
leaflets; petioles 1.57.5 cm long; rachis 418 cm long, terete to marginate. Leaflets slightly
oblique, parchment-like, sometimes coriaceous, glabrous or rarely very sparsely hairy above,
hairy beneath; narrowly ovate or elliptic, (2.5)3.510.5(18) x(1)24(5) cm; base
attenuate-acute, margin entire at base, distantly serrate towards apex, apex acute or
acuminate, acumen obtuse or slightly retuse; lateral veins 58 on each side, 516(22) mm
apart; domatia mostly absent. Inflorescences mostly much-branched, sometimes unbranched;
bracts and bracteoles to 3 mm long; pedicels to 2 mm long. Flowers greenish, whitish or
yellowish brown, c. 5 mm diameter; sepals suborbicular to subelliptic, 1.52.5 x 12 mm,
glabrous or sparsely to sometimes densely hairy outside, subglabrous inside; disc (1.2)1.5
1.8 mm diameter; stamen filaments with long patent hairs, in male flowers (2.5)3.23.5 mm
long, in female flowers (0.5)1.52 mm long, anthers narrowly cordate to ovate, 0.60.8x
0.40.7 mm, glabrous or sometimes sparsely hairy; ovary ellipsoid, 2-locular, (1.5)1.82x
(1)1.31.5 mm, sparsely hairy, often densely hairy along the sutures, mostly densely scaly,
style 11.2(1.5) mm long; pistillodes 0.51.2 mm long. Fruits brownish, obovoid to
globose, rarely transversely ellipsoid, 1(2)-seeded, 914(20) x814(20) mm; fruit wall 1
1.5(3) mm thick, sparsely to densely hairy or more or less densely scaly outside, densely
hairy inside. Seeds subellipsoid, 812 x5.58 mm; sarcotesta covering (25)4560% of the
seed, dorsally interrupted by a wide cuneate cleft; testa parchment-like.
Distribution. Borneo, Philippines, Sulawesi, Maluku, New Guinea. In Sabah, very
uncommon, known only from Silumpat Is. in the Lahad Datu district; not recorded from
Sarawak.
Ecology. Primary and secondary forests, often at forest edges, along rivers, on the banks of
lakes, on ridges, or in open vegetation; mostly in well-drained, sometimes in swampy habitat;
on limestone rock, sandy, or loamy soil; to 1700 m.
6. DIMOCARPUS Lour.
(Greek, didumos = double, karpos = fruit; the deeply 2-lobed fruits)
P.W. Leenhouts
Fl. Coch. (1790) 233; Leenhouts, Blumea 19 (1971) 113, FM 1, 11 (1994) 511; Anderson, CLTS (1980) 310; Whitmore &
Tantra, CLS (1986) 213; Yap, TFM 4 (1989) 438; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 310.
Trees or shrubs, mostly monoecious. Indumentum often partly or mainly of dense tufts of
hairs. Leaves paripinnate, rarely unifoliolate. Leaflets opposite or alternate, not or hardly
282
Fig. 5. Dictyoneura acuminata subsp. acuminata. A, flowering leafy twig; B, fruit valve from inside; C, seed, ventral view;
D, seed, dorsal view; E, embryo. (A from SAN 37193), BE from Kostermans (6892.)
283
papillose and often with exposed glands beneath, margin entire to dentate. Inflorescence a
panicle, terminal and sometimes in the axils of upper leaves. Flowers unisexual, radially
symmetrical; sepals 56, confluent at base, imbricate, equal, not petaloid, densely tomentose
outside, short-hairy at least in the upper part inside, not ciliate, entire; petals absent or 56,
mostly longer than sepals, more-or-less oblanceolate, without scales; disc entire, 5-lobed,
hairy; stamens (6)8(10), equal or more-or-less distinctly alternately long and short,
exserted or not, filaments usually hairy, often tufted, anthers glabrous; ovary sessile, 23carpellate, broadly cordate, tuberculate, hairy, usually each wart crowned by a hair-tuft,
style slender, slightly shorter to longer than the ovary, variably hairy at least in the lower
half, hairs mostly tufted, stigma with spreading lobes, ovules 1 per locule, axillary near the
base; pistillodes small, densely pilose. Infructescences with thickened and sometimes
elongated pedicels; persistent calyx slightly or not accrescent. Fruits nearly always with only
1 lobe developing, globose or broad-ellipsoid, indehiscent or sometimes loculicidal; fruit wall
warty, sometimes smooth, rarely spiny, mostly glabrescent outside, smooth and glabrous
inside. Seeds more or less globose; hilum subbasal, suborbicular, large; testa shiny, blackish
brown; arillode thin, translucent-white, fleshy.
Distribution. Six species; in S and SE Asia from Sri Lanka and India to eastern Malesia and
Australia. In Sabah and Sarawak, three species.
Key to Dimocarpus species
1. Leaflet margin serrate-dentate. Leaf with 47 pairs of leaflets......................1. D. dentatus
Leaflet margin entire or repand, rarely sinuous or distantly dentate. Leaf with 14(6)
pairs of leaflets....................................................................................................................2
2. Glands on lower surface of leaflets present in the axils of lateral veins and elsewhere,
often near or along the margin. Petals absent or 1, reduced. Disc woolly......2. D. fumatus
Glands on lower surface of leaflets usually only present in the axils of lateral veins,
sometimes absent. Petals 5. Disc velutinous.....................................................3. D. longan
1. Dimocarpus dentatus Meyer ex Leenh.
Fig. 6AD.
(Latin, dentatus = toothed; the leaflet margin)
l.c. (1971) 116, l.c. (1994) 513. Type: Ambulah SAN 37193, Borneo, Sabah (holotype L; isotype K).
Tree to 15(24) m tall, 40 cm diameter, sometimes with buttresses. Leaves each with 47
pairs of leaflets; petiole and rachis densely hairy; petioles 618 cm long. Leaflets thincoriaceous to stiff-chartaceous, hairy on midrib and lateral veins above, glabrescent, rather
densely to sparsely tufted-hairy on midrib and lateral veins beneath, in between lateral veins
often with scattered tufted, paired or solitary hairs, with exposed glands in the axils of lateral
veins and along the margin; oblong-obovate to lanceolate, 5.524 x3.29 cm; base acute and
symmetric or slightly asymmetric and cordate in the lower leaflets, margin distantly serrate
or dentate, apex acute to rounded, sometimes attenuate-acuminate; midrib flat above, lateral
veins 815 mm apart, alternately ending in and between the marginal teeth, sunken above,
intercostal veins scalariform, rather dense, hardly visible above, raised beneath; petiolules 1
2 mm long.
284
Fig. 6. Dimocarpus dentatus (AD); D. fumatus (E); D. longan var. echinatus (F); D. longan var. longan (G); D. longan var.
malesianus (H). A, fruiting leafy twig; B, detail of lower leaflet surface; C, fruit; D, young seed with partly developed
arillode; EH, fruits. (AB from SAN 37193, CD from Kostermans 12654, E from Kostermans 13245, F from SAN 61671,
G from Tilley, s.n., H from S. 32681.)
285
Inflorescences 2555 cm long, densely ferruginous-tomentose, hairs tufted; branches few,

erecto-patent, to 20 cm long, bearing rather numerous sessile, few-flowered, glomerulous
cymules; pedicels to 1.5 mm long; bracts narrowly triangular, to 4 mm long, patent to
recurved; sepals 2.53 x 1.82 mm, sparsely hairy in the upper part inside; petals 5,
oblanceolate, 3.54.2 x 1 mm, sericeous outside except at the base and sometimes in the
upper half, densely long-hairy along the margin and inside except at the base; disc velutinous;
stamen filaments 2.24 mm long, patently tufted-hairy mainly towards the apex, anthers to
0.8 mm long. Fruit lobes subglobose, c. 16 x 15 mm; fruit wall with rather large, hardly
raised, flat warts, granular, glabrous.
Vernacular name. Sabahrambutan kunyit (Kedayan).
Distribution. Endemic to Borneo. In Sabah, locally common; so far no record from Sarawak;
also in E Kalimantan.
Ecology. Primary, sometimes secondary, forest, in well-drained as well as in periodically
inundated or marshy localities, on various types of soil, from sea level to 750 m. Flowering in
May, July, October to November; fruiting in July to August.
Uses. Arillode sweet, edible.
2. Dimocarpus fumatus (Blume) Leenh.
Fig. 6E.
(Latin, fumatus = smoke-coloured; the twigs)

l.c. (1971) 119, l.c. (1994) 515; Anderson l.c. 310; Yap l.c. 439. Basionym: Nephelium fumatum Blume, Rumphia 3 (1847)
111. Type: Korthals, s.n. (or Mller s.n.?), SE Borneo (L). Synonym: Pseudonephelium fumatum (Blume) Radlk., Sapind.
Holl.-Ind. (1879) 71, in Engler, Pflanzenr. 98 (1932) 329, Merrill l.c. 359, Masamune l.c. 432.
subsp. fumatus
Tree, rarely shrub, to 27 m tall, 1 m diameter, sometimes with buttresses. Leaves each with
(2)3(4) pairs of leaflets; petiole and rachis thin-hairy, mostly early glabrescent; petioles
1.512 cm long, flat to sometimes slightly hollowed above, exceptionally terete. Leaflets
alternate to subopposite, thin-coriaceous to sometimes chartaceous, glabrous and minutely
punctate above, glabrous to sparsely hairy on midrib, vein axils and veins beneath; exposed
glands present in the axils of lateral veins and along or in the incisions of the margin
beneath; elliptic to oblong, 6.528 x 2.510.5 cm; base symmetric to asymmetric, cuneate to
rounded, decurrent or not, margin repand to sinuous (mainly in the apical part) or distantly
dentate, sometimes entire, apex attenuate to abruptly acuminate; midrib slightly raised to
hardly sunken above, lateral veins 12.8 cm apart, not joined, slightly raised or rarely
grooved above, intercostal veins and veinlets finely tessellate-reticulate, mostly
inconspicuous above, distinct beneath; petiolules 2.515 mm long, grooved above, often with
a median rib. Inflorescences lax, to 50 cm long, with a few branches; cymules sessile, mostly
many-flowered, sparsely hairy; pedicels 24 mm long, slender; bracts subulate, to 3 mm long.
Flowers: sepals connate to one third of their length, 23 x 1.52.5 mm, hairy inside; petals
absent or rarely 1, much reduced; disc woolly; stamen filaments 1.52.5 mm long, glabrous
or nearly so, anthers 0.60.8 mm long. Fruit lobes 23.5 cm diameter, hardly warty to shortspiny, granular, glabrous; dehiscent by valves.
286
Vernacular names. Sabahmambuakat (Dusun Kinabatangan). Sarawakmarakiang,

ribau balabau (Iban).
Distribution. Peninsular Thailand, Peninsular Malaysia and Borneo. Common in Sabah,
uncommon but widespread in Sarawak; also in Brunei and Kalimantan.
Ecology. In primary mixed dipterocarp forest, on flat country, slopes, hill tops, or along
riverbanks, mainly on yellow/red sandy loam soils, from lowlands to 1350 m. Flowering in
April to July, September, November; fruiting in July, September, December.
3. Dimocarpus longan Lour.
(from the Chinese fruit-name long-yen = dragons eye; the dark seed subtended by a pale arillode)
Fig. 6FH.
Fl. Coch. (1790) 233; Leenhouts l.c. (1971) 122, l.c. (1994) 517; Anderson l.c. 310; Yap l.c. 439; Wong & Ketso, PROSEA
2 (1991) 146. Neotype (Leenhouts, 1971): Liao & Kuo 1598, Taiwan (L; isoneotype TAI). Synonyms: Nephelium longan
(Lour.) Hook., Curtis Bot. Mag (1844) t. 4096; N. long-yan Blume, Rumphia 3 (1849)108.
Tree, very rarely shrub, to 30(40) m tall, 30(80) cm diameter; buttresses to 2 m high.
Leaves each with 24(6) pairs of leaflets; rachis mostly densely hairy; petioles 320 cm
long, terete to flattened above. Leaflets stiff-chartaceous to coriaceous, with or without
exposed glands or hair-tufts in the axils of lateral veins beneath, often hairy in the basal part
of the midrib above; elliptic to ovate, 345 x 1.820 cm; base symmetric to asymmetric, acute
to rounded, rarely decurrent, margin entire, apex mostly attenuate-acute to obtuse-acuminate,
sometimes obtuse to acute; midrib sunken or rarely flat to raised above, lateral veins 0.43
cm apart, at least in the lower half of leaflet not joined, slightly raised to faintly grooved
above, intercostal veins mostly more-or-less scalariform, rather dense, often inconspicuous
above, more-or-less raised beneath; petiolules 0.235 mm long, mostly grooved above.
Inflorescences 840 cm long, densely tufted tomentose; branches few to several; cymules
subsessile to distinctly stalked, (1)35-flowered; pedicels 13 mm long, rather slender;
bracts patent, oblong-ovate to narrowly lanceolate, 1.55 x 0.61.5 mm. Flowers: sepals 25
13 mm, partly short-hairy inside; petals 5, broadly to narrowly spathulate, 1.56 x0.82
mm, for the greater part densely woolly (large ones) to subglabrous (small ones) on both
sides, apex woolly or with sessile glands inside; disc velutinous; stamen filaments 16 mm
long, mostly woolly (hairs often tufted) except at the base and less often at the apex, anthers
0.61.5 mm long. Fruit lobes broad-ellipsoid to globose, 13 x 13 cm; fruit wall smooth to
warty or with spines to 1 cm long, sometimes granular, glabrescent.
Vernacular names. Sabahbambo (Idahan), mabo (Dusun), mahau (Kedayan), mambu
(Tenggara), mumboh (Dusun Kinabatangan), rafag (Banggi), sabon-sabon (Bajau).
Sarawakapong keliau (Berawan), buah keregan (Punan Tutoh), chiru (Malay-Lundu),
enkiong, gurin (Iban), kayo bilong (Kayan), kayu soman (Kenyah), mata kucing (Malay),
merkiang (Iban).
Distribution. Continental Asia from Sri Lanka and India to S China, Hainan, and Taiwan,
and Malesia (in Java and New Guinea probably only naturalized; not seen from the Lesser
Sunda Islands). In Sabah and Sarawak, 2 subspecies with 3 varieties are recognised.
287
Key to subspecies and varieties

1. Leaves mostly distinctly asymmetric at base; midrib and lateral veins nearly always flat
or slightly raised above; petiolules rarely grooved................................................................
subsp. longan var. longan
Synonyms: Sapindus longifoloius Vahl, Symb. 3 (1794) 53; Nephelium bengalense G.
Don, Gen. Hist. 1 (1831) 670; N. pupillum Wight, Illustr. 1 (1840) 141; Euphoria
echinulata Radlk., Pflanzenr. 98 (1932) 903; N. echinulatum (Radlk.) Ridl., FMP 1
(1922) 503.
Leaves each with (2)45 pairs of leaflets; petioles 38 cm long. Leaflets usually
opposite, stiff-chartaceous, mostly glabrous above, subglabrous beneath; domatia rare
and never hairy; oblong-ovate to oblanceolate, 319 x 1.86.5 cm; base at least in the
upper leaflets distinctly asymmetric, acute, apex obtuse to shortly, broadly, and
obtusely acuminate; midrib flat to slightly raised above, lateral veins 0.51.5 cm
apart, usually slightly raised on both sides; petiolules 210 mm long. Inflorescences
with distinctly stalked, 13-flowered cymules, Fruit subglobose, c. 1.2 cm across,
mostly finely pustulate, granular or nearly smooth.
Continental S and SE Asia and Malesia (probably indigenous only in Peninsular
Malaysia, and naturalized in some parts of Java, Borneo, Philippines and New
Guinea). In Sabah, uncommon and of uncertain origin.
Leaves nearly always symmetric at base; midrib and lateral veins nearly always sunken
above; petiolules often grooved...........................................................................................2
2. Fruits densely covered with 0.81 cm long, flattened spines.................................................
subsp. malesianus var. echinatus Leenh.
l.c. (1971) 128, l.c. (1994) 519. Based on Euphoria nephelioides Radlk., Philip. J. Sc.
8 (1914) Bot. 457. Type: Kleme EB 15218, Philippines, Basilan (M).
Leaves each with 14 pairs of leaflets; petioles 69 cm long. Leaflets 422 x 1.59
cm, hairy or not; domatia present or absent; base more-or-less symmetric, apex
acuminate; lateral veins 11.8 cm apart, intercostal veins slightly sunken beneath,
invisible above; petiolules 315 mm long.
Borneo (Sabah and Kalimantan) and the Philippines (Mindanao and Basilan).
Fruits smooth to warty...........................................................................................................
subsp. malesianus var. malesianus Leenh.
l.c. (1971) 126, l.c. (1994) 519. Based on Sapindus cinereus Turc., Bull. Soc. Nat.
Mosc. 31 (1858) 402. Neotype (Leenhouts 1971): Cuming 1131, Philippines, Luzon
(isoneotypes BM, FI, K, L). Synonyms: Nephelium malaiensis Griff., Notul. 4 (1854)
549, Masamune l.c. 428; Euphoria cinerea Radlk., Pflanzenr. 98 (1932) 905; E.
malaiensis (Griff.) Radlk. l.c. (1932) 909; Sapindus stellatus Turz. l.c. 403; E. stellata
(Turz.) Radl. l.c. (1932) 908, Merrill l.c. (1921) 359, Masamune l.c. 426, Anderson
l.c. 311; E. elongata Radlk., Sapind. Holl.-Ind. (1879) 7, 25, Merrill l.c. (1921) 359,
Masamune l.c. 426; E. sclerocarpa Radlk. in Merrill, PEB (1929) 174, Masamune l.c.
426; E. succulenta Radlk. in Merrill l.c. (1929) 174, Masamune l.c. 426; E.
microcarpa Radlk. l.c. (1932) 907, Anderson l.c. 311.
Leaves each with 24(6) pairs of leaflets; petioles (3)610(20) cm long. Leaflets
opposite to alternate, mostly slightly hairy, often with hairy domatia; base mostly
symmetric, apex often acuminate. Inflorescences often with subsessile cymules.
Myanmar, Laos, Cambodia, S Vietnam and Malesia (Sumatra, Peninsular Malaysia,
Borneo, Philippines, Sulawesi and Maluku). Common in Sabah and Sarawak; also in
Brunei.
288
7. GANOPHYLLUM Blume
(Greek, ganos = shiny, phullon = leaves)
P.W. Leenhouts
Mus. Bot. Lugd. Bat. 1 (1850) 230; Radlkofer in Engler, Pflanzenr. 98 (1933) 1423; Backer & Bakhuizen f., FJ 2 (1965)
141; Whitmore & Tantra, CLS (1986) 213; Yap, TFM 4 (1989) 440; Leenhouts, FM 1, 11 (1994) 538.
Tall trees, possibly monoecious. Indumentum of only simple, solitary hairs on the
inflorescences; glandular scales common on twigs, leaves, inflorescences, and on outside of
sepals; young parts sticky resinous. Leaves paripinnate, each with 410 pairs of leaflets;
pseudostipules absent; neither petiole nor rachis winged. Leaflets herbaceous to stiffly
parchment-like; base slightly to strongly asymmetric, margin entire. Inflorescences axillary
thyrses, flowers solitary or in small cymules on racemose branches; bracts caducous. Flowers
unisexual, radially symmetrical; sepals 46, nearly free to about halfway connate, valvate, all
about equal, not petaloid, entire; petals absent; disc complete, consisting of short-hairy or
glabrous lobes opposite the sepals; stamens 57, alternating with the disc lobes, in male
flowers far exserted, glabrous, anthers basifixed, laterally dehiscent; ovary 2(3)-locular,
sessile, sparsely short-hairy, style apical, about as long as the ovary, stigma indistinctly lobed,
ovules 2 per locule, one above the other, pendulous. Fruit a sessile drupe, not winged,
smooth, glabrous, with 1 or 2 locules. Seed 1 per locule; arillode absent; embryo curved,
radicle in a pocket of the testa, directed towards the hilum, outer cotyledon more or less
enveloping the smaller inner one; germination epigeal, seedling leaves from the start
paripinnate, first leaves with the petiole and rachis marginate.
Distribution. Two species, one in W and C Africa, the other from the Andamans and
Nicobars to NE Australia and the Solomon Is. One species in Sabah.
Ecology. Canopy tree of evergreen and deciduous forest.
Ganophyllum falcatum Blume
Fig. 7.
(Latin, falcatus = sickle-shaped; the leaflets)
l.c. (1850) 230; Radlkofer in Engler l.c. 1424; Backer & Bakhuizen f. l.c. 142; Whitmore & Tantra l.c. 213; Yap l.c. 440;
Leenhouts l.c. 538. Type: Sine coll., s.n. (= L. sheet no. 902.3569), New Guinea (L).
Tree to 42 m tall, 150 cm diameter, with buttresses. Bark dark to grey-brown, peeling off as
papery flakes. Leaves each with (4)58(10) pairs of leaflets; petioles 3.69 cm long,
subterete with an upwards widening groove above, grading into the rachis. Leaflets:
petiolules 25 mm long, lowermost leaflets ovate, c. 2.5 x 1 cm, base strongly asymmetric;
upper leaflets elliptic, often more or less sickle-shaped, to 23 x 8 cm, base hardly asymmetric;
apex hardly to distinctly acuminate, acumen short, broad, and rounded; midrib flat to slightly
raised above, distinct below, lateral veins rather distant, oblique-patent, mostly looping and
joining near the margin, intercostal veins and veinlets many, distinct, reticulations rather lax,
in thick leaflets indistinct above. Inflorescences to 20 cm long, widely and (especially in the
female ones) sparsely branched; bracts and bracteoles broad-triangular scales; pedicels c. 2
mm long.
289
TREE FLORA OF SABAH AND SARAWAK VOL. 2 (199
Fig. 7. Ganophyllum falcatum. A, flowering leafy twig; B, fruit; C, seed; D, embryo. (A from Brass 2440, BD from SAN
64306.)
290
Flowers: sepals 1.251.5 mm long, green, hairy inside; disc orange, short-hairy; stamen
filaments 34 mm long, white, anthers c. 0.75 mm long, yellow, staminodes in female
flowers strongly reduced; ovary c. 2.25 mm high, yellow-green, ellipsoid, tapering into a
columnar style, pistillodes in male flowers well-developed. Fruits ovoid to subglobose, 10
20 x 710 mm, red; persistent calyx reflexed.
Vernacular name. Sabahpanapok ayer (Dusun).
Distribution. Andaman and Nicobar Is. to Australia (Western parts, the Northern Territory,
and Queensland) and the Solomon Is. In Borneo, uncommon, so far reported from Sabah only
(Tenom and Tambunan districts).
Ecology. Primary and secondary forests; along forest edges, on riverbanks, inner edge of
mangroves, on well-drained or periodically inundated level lands, on slopes, clay, and
limestone, to 1200 m. Flowering mainly in AugustNovember; fruiting in NovemberMarch.
The fruits are eaten and dispersed by different kinds of birds (Japing & Oey Djoen Sen,
Tectona 29 (1936) 421, fig. 29; Meyer Drees, Commun. Forest. Res. Inst. 33 (1951), 109).
Uses. The timber is of a good quality; also used for making matches and match-boxes. The
bark is used in preparing soap and as a fish poison (Japing & Oey Djoeng Sen, Tectona, 29
(1936) 421, fig. 29; Quisumbing, Philip. J. Sci. 77 (1948) 161; Brown, Useful Pl. Philipp. 2
(1950) 361, fig. 176).
8. GLENNIEA Hook. f.
(Probably after Lady Glennie)
P.W. Leenhouts
In Bentham & Hooker f., Gen. Pl. 1 (1862) 404; Radlkofer in Engler, Pflanzenr. 98 (1932) 858; Leenhouts, Blumea 22
(1975) 411, FM 1, 11 (1994) 540; Whitmore & Tantra, CLS (1986) 213; Yap, TFM 4 (1989) 440; Whitmore, Tantra &
Sutisna, CLK 2, 1 (1990) 311. Synonyms: Crossonephelis Baill., Adansonia 11 (1874) 245; Hedyachras Radlk., Bot. Jahrb.
56 (1920) 258.
Trees, monoecious or dioecious. Indumentum mainly of solitary hairs or small tufts of hairs.
Leaves spirally arranged or partly decussate, unifoliolate or paripinnate, each with 16
pairs of leaflets, without pseudostipules, neither petiole nor rachis winged. Leaflets opposite
to alternate, smooth, glabrous or variably hairy beneath; ovate to elliptic; base symmetric or
slightly oblique, margin subentire, apex rounded-acuminate, acumen rounded; lateral veins
looping and joining in the upper part only, intercostal veins and veinlets finely reticulate,
slightly raised on both sides. Inflorescences terminal or in the axils of upper leaves, thyrsoid
or paniculate, with few spreading branches. Flowers radially symmetrical, unisexual, if plant
monoecious the male and female flowers in the same inflorescence; sepals 4 or 5, connate at
base, valvate to narrowly imbricate in bud, spreading during anthesis, persistent and recoiled
291
after anthesis, equal, deltoid, not petaloid, densely hairy outside, variably hairy inside; petals
absent; disc for the greater part adnate to the base of the calyx, complete, broad and flat, more
or less distinctly lobed, purplish black when dry, glabrous or rarely variably hairy; stamens 6
or 7, equal, exserted, glabrous, filaments filiform, anthers basifixed, dehiscing lengthwise,
latrorse to introrse, connective broad; staminodes short; ovary sessile, 2-locular, hairy, style
apical, conical, with 2 stigmatic lobes or grooves, ovules 1 per locule; pistillodes small, white
woolly. Fruits indehiscent; pericarp thick, endocarp membranous to thin-crustaceous, wings
absent. Seeds without arillode; testa thin-crustaceous, closely adhering to the endocarp.
Distribution. Eight species, three occurring in tropical Africa, one in Madagascar, one in Sri
Lanka, and three in Malesia (of which two are found in Borneo).
Ecology. In lowland rain forest.
Key to Glenniea species
Fruits 6 cm long or more. Inflorescences densely hairy, bearing densely arranged, manyflowered cymules........................................................................................1. G. philippinensis
Fruits less than 2 cm long. Inflorescences sparsely hairy, with scattered, few-flowered
cymules or solitary flowers....................................................................................2. G. thorelii
1. Glenniea philippinensis (Radlk.) Leenh.
l.c. (1975) 412, l.c. (1994) 542. Basionym: Hedyachras philippinensis Radlk., Bot. Jahrb. 56
(1920) 258. Type: Villamil FB 20635, Philippines (holotype M; isotypes BM, K, L).
Synonym: Crossonephelis philippinensis Leenh., Blumea 21 (1973) 100.
Monoecious tree to 18 m tall, 30 cm diameter. Hairs mainly solitary. Leaves spirally
arranged, each with (1)46 pairs of leaflets; petioles 39 cm long, flat to grooved at base,
with marginal ribs, more or less flattened at the upper part. Leaflets opposite to alternate,
stiff-chartaceous, with hairy domatia in the axils of lateral veins beneath; 522 x2.759 cm;
base more or less symmetric, rounded in the lower, acute and attenuate in the upper leaflets,
apex obtuse to broadly and obtusely acuminate; midrib slightly raised above, lateral veins 1
3.5 cm apart, slightly to distinctly curved, those in the upper half of the leaflets more or less
distinctly looping and joining at some distance from the margin, slightly raised above, more
so beneath, intercostal veins more-or-less strongly developed; petiolules 210 mm long, with
a broad and flat to narrow and deep groove above. Inflorescences thyrsoid, 2025 cm long,
fulvous-velutinous; cymules crowded, sessile, glomerulous, many-flowered; pedicels 1.52
mm long; bracts and bracteoles lanceolate, 22.5 mm long. Flowers: sepals 4, valvate to
narrowly imbricate, 23 x1.52.5 mm, hairy inside; disc glabrous; stamens 6 or 7, filaments
c. 5 mm long, anthers broadly ellipsoid, c. 1 mm long, introrse; ovary obcordate, style very
short, stigma grooved. Fruits pear-shaped, 67 x 4.56 cm, yellow when fresh, glabrous;
292
pericarp thick, fleshy, endocarp tough. Seeds more or less ovoid, 33.5 x 1.752.25 cm; testa
brown, smooth, glabrous.
Distribution. SE Thailand, Vietnam, Borneo, Philippines. Very uncommon in Sabah (Tenom
district); no record from Sarawak.
Ecology. In thickets and forests along streams at low altitude. Flowering in June, October;
fruiting in MayDecember.
Use. The fruits are edible (Madulid, Nat. Mus. Papers 2, 1 (1991) 56).
2. Glenniea thorelii (Pierre) Leenh.
(C. Thorel, 18331911, French physician, botanist, and plant collector)
Fig. 8.
l.c. (1975) 412, l.c. (1994) 544; Whitmore & Tantra l.c. 213; Whitmore, Tantra & Sutisna l.c. 311. Basionym: Cnemidiscus
thorelii Pierre, Fl. Coch. (1894) t. 320a, text. Type: Pierre 4089, S Vietnam (P). Synonyms: Crossonephelis thorelii (Pierre)
Leenh. l.c. (1973) 101; Lepisanthes palawanica Radlk. in Elmer, Leafl. Philip. Bot. 5 (1913) 1604; Crossonephelis
palawanicus (Radlk.) Leenh. l.c. (1973) 97.
Tree to 30 m tall, 50 cm diameter, buttressed; probably dioecious. Indumentum mainly of

small stellate tufts of hairs. Leaves spirally arranged to subdecussate, each with 12 pairs of
leaflets; petioles 2.510 cm long, 3-angular to terete. Leaflets subopposite, chartaceous,
glabrous except for occasional hair-tufts in the axils of some lateral veins beneath; 626 x3
13.5 cm; base symmetric to slightly asymmetric, cuneate to rounded, attenuate, apex obtuse
or gradually to abruptly, broadly, and obtusely acuminate; midrib slightly raised above,
sometimes sunken towards the base, lateral veins 1.256 cm apart, usually strongly curved,
about equally slightly raised on both sides, intercostal veins faintly to sometimes strongly
developed; petiolules 0.31.5 cm long, with a shallow broad groove above. Inflorescences
thyrsoid to paniculate, to 25 cm long, slightly hairy; cymules sometimes distant, few-flowered
or flowers solitary; pedicels to 5 mm long; bracts and bracteoles 3-angular, to 1 mm long.
Flowers: sepals 4 or 5, valvate, 22.5 x1.52.5 mm, tomentose, with glabrous longitudinal
strips inside; disc glabrous to velutinous; stamens 6 or 7, filaments c. 5 mm long, anthers
ovoid, c. 1 mm long, latero-introrse; ovary slightly 2-lobed, style short, stigma 2-lobed with
short, thick lobes curved outwards, to knobby. Fruits 2-lobed, c. 1.25 x2 1 cm, often 1 lobe
suppressed, then transversely ovoid and c. 1.25 x1.5 1.25 cm, smooth and glabrous; pericarp
thin, fleshy. Seeds: testa closely adhered to the endocarp, thin-coriaceous.
Distribution. S Vietnam, Sumatra, Borneo, Philippines, and New Guinea. Uncommon in
Sabah; also in Kalimantan.
Ecology. Primary forest on alluvial plains, slopes, or ridges, also on riverbanks, to 200 m.
Flowering April, October, December; fruiting March, September.
Uses. Produces a good quality timber (Desch, Mal. For. Rec. 15, (1954) 529). The bark is
easily inflammable and is used for kindling fire.
293
Fig. 8. Glenniea thorelii. A, flowering leafy twig; B, fruit. (A from Lambach 1336, B from Boschproefstation T. 973.)
294
9. GUIOA Cav.
(J. Guio, Spanish botanical artist)

lokud-lokud (Dusun, Sabah)
P.C. van Welzen

Icon. 4 (1798) 49, t. 373; Merrill, EB (1921) 361; Radlkofer in Engler, Pflanzenr. 98 (1933) 1157; Masamune, EPB (1942)
426; Backer & Bakhuizen f., FJ 2 (1965) 139; Anderson, CLST (1980) 311; Corner, WSTM 3rd ed. 2 (1988) 680; Yap, TFM
4 (1989) 441; van Welzen, Leiden Bot. Ser. 12 (1989) 146, FM 1, 11 (1994) 548.
Shrubs or trees. Indumentum of simple solitary hairs and indistinct glandular hairs. Leaves
paripinnate, each with 19 pairs of leaflets, without pseudostipules; petioles pulvinate; rachis
terete to distinctly winged. Leaflets opposite to alternate, often punctate; glandular hairs
indistinct, white; lower surface smooth to whitish papillose, domatia usually present as sacs
or pockets; petiolules very short, pulvinate. Inflorescence a thyrse, axillary, pseudoterminal
or borne on leafless twigs, usually few-flowered; cymules in cincinnate to dichasial clusters;
pedicels articulate. Flowers bilaterally symmetrical, structurally hermaphrodite, but
presumably functionally unisexual; sepals 5, margin usually with glands, inner 3 sepals with
a petaloid margin; petals 5, clawed, scales present and usually crested, petals between two
adjacent large sepals usually with reduced blade and scales; disc interrupted or complete,
lobed, smooth, glabrous; stamens 8, anthers basifixed in cleft, latrorse; ovary 3-lobed, 3locular, smooth, ovules 1 per locule, stigma sessile, pyramidal, longitudinally grooved, style
elongating in fruit. Fruit a subcordate loculicidal capsule with 3 lobes, of which one to all
develop, red; stalk narrow and high to broadly obconical and indistinguishable; lobes laterally
flattened. Seeds globose to obovoid, black, in open fruits dangling from the pseudofunicle;
arillode completely enveloping the seed, apically open and lobed, at base with a rim from
which a pseudofunicle is attached to the basal corner of the seed; hilum ovoid; cotyledons
superposed to subcollateral, usually unequal, with the upper one larger.
Distribution. About 65 species, in SE Asia (Thailand to S Vietnam as the northern limit),
throughout Malesia (43 species) to E Australia and into the Pacific up to Samoa and New
Caledonia. In Sabah and Sarawak, 6 species.
Ecology. In secondary and primary forests, often locally common; along road sides,
riverbanks, forest edges, beaches, plantation edges. Several species are partly confined to
ultramafic soil; from sea level to1800 m.
Key to Guioa species
1. Rachis of leaves slightly to distinctly winged......................................................................2
Rachis of leaves not winged................................................................................................4
2. Leaves each with 14 pairs of leaflets; upper leaflets elliptic; lower surface of leaflets
smooth to somewhat papillose, rather shiny; domatia absent or a single small sac...............
........................................................................................................1. G. bijuga (in part)
295
Leaves each with (1)27 pairs of leaflets; upper leaflets elliptic to obovate; lower
surface of leaflets densely greyish papillose, dull; domatia many......................................3
3. Wings of rachis narrow, to 3 mm broad. Leaflet apex obtuse, abruptly acuminate to
cuspidate; midrib convex and raised below; domatia pocket- to sac-shaped.........................
...................................................................................................4. G. pleuropteris (in part)
Wings of rachis to 4 mm broad. Leaflet apex gradually acuminate to cuspidate; midrib
hardly raised or flattish below; domatia sac-shaped................................5. G. pterorhachis
4. Domatia absent or a single sac.............................................................................................5
Domatia of two to many sacs or pockets.............................................................................8
5. Petal scales very much broadened at apex; crest usually absent. Fruit lobes usually
broader than high, glabrous.......................................................................2. G. diplopetala
(in part)
Petal scales hardly broadened at apex; crest always present. Fruit lobes broader than high
and sparsely sericeous or about as broad as high and glabrous............................................6
6. Leaves each with 14 pairs of leaflets. Leaflets elliptic, usually symmetric, apex abruptly
acuminate to cuspidate, acumen usually obtuse to acute, not mucronulate...........................
.............................................................................................................1. G. bijuga (in part)
Leaves each with 16 pairs of leaflets. Leaflets ovate to elliptic, asymmetric, apex usually
gradually acuminate to caudate, acumen acute to usually mucronulate...............................7
7. Leaflets smooth, shiny, glabrous below. Ovary subhirsute. Fruits glabrous..........................
...................................................................................................3. G. koelreuteria (in part)
Leaflets papillose, dull, and slightly short-sericeous below. Ovary densely hirsute. Fruits
sparsely sericeous, glabrescent......................................................6. G. pubescens (in part)
8. Petal scales very much broadened at apex; crest usually absent............................................
.....................................................................................................2. G. diplopetala (in part)
Petal scales not to hardly broadened at apex; crest present..................................................9
9. Lower leaflet surface smooth, glabrous; ovary subhirsute.........3. G. koelreuteria (in part)
Lower leaflet surface slightly short-sericeous to silky or rough hairy. Ovary densely
hirsute or slightly rough-hairy...........................................................................................10
10. Upper leaflets elliptic to obovate, apex obtuse, abruptly acuminate to cuspidate; lower
surface somewhat silky hairy to rough-hairy, domatia pocket- or sac-shaped. Ovary
slightly rough-hairy. Fruits glabrous..........................................4. G. pleuropteris (in part)
Upper leaflets ovate to elliptic; apex gradually acuminate to cuspidate; lower surface
slightly short-sericeous, domatia sac-shaped. Ovary densely hirsute. Fruits sparsely
sericeous, glabrescent....................................................................6. G. pubescens (in part)
1. Guioa bijuga (Hiern) Radlk.
(Latin, bijugus = with two pairs of leaflets; the leaf)

Sapind. Holl.-Ind. (1879) 38; Anderson l.c. 311; Corner l.c. 680; Yap, l.c. 441; van Welzen l.c. (1989) 183, l.c. (1994) 567.
Basionym: Cupania pleuropteris Blume var. bijuga Hiern in Hooker f., Fl. Br. Ind. 1 (1875) 677. Type: Wallich KD 8094,
Malaya (holotype K; isotypes BM, K). Synonyms: Guioa pleuropteris (Blume) Radlk. var. bijuga (Hiern) King, J. As. Soc.
Beng. 65, 2 (1896) 444; Guioa rubrofusca Radlk. ex Merr., EPB (1929) 175, nom. nud., Masamune l.c. 427.
296
Tree to 30 m tall, 35 cm diameter. Bark smooth to somewhat finely fissured, usually hard,
whitish to dark grey with dark patches, or red-brown; inner bark yellow to red to brown,
fibrous; cambium white to yellow to red. Sapwood white to brown; heartwood pinkish
brown. Leaves each with 14 pairs of leaflets; rachis 0.816.8 cm long, winged or not, wings
to 2 mm broad; petioles 0.711 cm long. Leaflets usually symmetric, thin- to thickcoriaceous, punctate; upper surface glabrous to slightly sericeous, lower surface duller,
smooth to slightly papillose, glabrous to sparsely sericeous especially on the midrib; domatia
absent or in the form of a single small sac; elliptic, 2.120 x1.18.4 cm; base sharply
attenuate, margin entire, flat to revolute, apex abruptly acuminate to cuspidate, acumen
obtuse to acute; lateral veins 0.34.2 cm apart, marginally looped and joined, less so in lower
third of leaflets, intercostal veins laxly reticulate, indistinct. Inflorescences axillary to
pseudoterminal, unbranched to basally branched; rachis 0.716.5 cm long, terete to flattened,
glabrous to pilose; first-order branches to 7.4 cm long; cymules cincinnate, 13(6)-flowered;
bracts and bracteoles deltate to triangular, bracts 0.51.9 mm long, bracteoles 0.21 mm long;
pedicels 2.210 mm long. Flowers 3.54.2 mm diameter; sepals 5, ovate, 2 outer smaller
ones 12.8 x12.3 mm, 3 inner larger ones 1.43.4 x1.23.8 mm, margin petaloid; petals 5,
white to yellow, slightly hairy, obovate, 1.83.8 x0.71.7 mm, blade obovate, subglabrous,
gradually decurrent into the 0.41.2 mm high claw, margin pilose, apex emarginate to acute,
scales (0.8)1.22 mm long, not to hardly broadened at apex, crest a pilose flat part of the
bifid scale apex; disc interrupted; stamens 8, filaments 1.65 mm long, anthers 0.20.4 mm
long; ovary 0.31.8 mm high, sparsely hirsute, style and stigma 0.22.5 mm long. Fruits 1.2
2.3 x1.32.6 cm, glabrous; stalks 25 mm long; lobes 8.514 x816 mm. Seeds globose to
obovoid, 810 x 78 mm; embryo 79 x 57 mm.
Vernacular names. Sabahbuah sungket (Kedayan), lapak lapak (Murut), rupag (Sungei
Kinabatangan), tinggal asam (Murut Tenom). Sarawakilat (Iban).
Distribution. Thailand, Peninsular Malaysia, Sumatra, Borneo, Philippines. In Sabah and
Sarawak, locally common; also in Brunei and Kalimantan.
Ecology. In primary and secondary kerangas, mixed dipterocarp and mixed peat swamp
forests, along road sides, riverbanks, edges of forest, mangroves, and sometimes in cultivated
fields; on different types of soil, including sandstone, podsolized white sand, ultramafic and
red-yellow loam. From sea level to 1500 m. Flowering mainly in November to April, less so
in August to September; fruiting mainly in March to June, less so in September and October.
Notes. In Borneo, it is often difficult to distinguish G. bijuga from G. pubescens. In most
cases, G. bijuga has elliptic leaflets which are smooth and glabrous below, and glabrous
fruits. G. pubescens, on the other hand, usually has ovate leaflets, which are papillose and
sericeous below and glabrescent fruit. In specimens collected from Sarawak and Mt.
Kinabalu, leaflets of G. bijuga may be papillose and subsericeous, while those of G.
pubescens are subelliptic and more glabrous. In these specimens the more ovate leaflets and
the few hairs remaining on the fruit are suggestive of G. pubescens, and the flattened to
winged rachis are reminiscent of G. bijuga. Notwithstanding these somewhat overlapping
characters, there are, in addition to those shown in the key, additional features which can be
used to separate G. bijuga from G. pubescens, viz., the former always has secretory idioblasts
in the leaf tissue, whereas the latter is without them.
297
2. Guioa diplopetala (Hassk.) Radlk.
(Greek, diplos = double, petalon = petal; the broad scales on the inside of the petals)
Fig. 9.
Sapind. Holl.-Ind. (1879) 88, in Engler l.c. (1933) 1162; Masamune l.c. 426; Backer & Bakhuizen f. l.c. 140; van Welzen l.c.
(1989) 197, fig. 76, 77, l.c. (1994) 570. Basionym: Cupania diplopetala Hassk., Flora 25, 2, Beibl. (1842) 39. Type: (Not
indicated, probably Hasskarls collection from Java; BO). Synonyms: Guioa regularis (Blume) Radlk. l.c. (1879) 12, 41,
Masamune l.c. 427; Guioa diplopetala (Hassk.) Radlk. forma borneensis Radlk., Sitzungsber. Math.-Phys. Cl. Knigl.
Bayer. Akad. Wiss. Mnchen 9 (1879) 610; Guioa diplopetala (Hassk.) Radlk. var. borneensis (Radlk.) Radlk., Bot. Jahrb.
49 (1913) 370; Guioa bullata Radlk., in Fedd. Repert. 18 (1922) 342, in Engler l.c. (1933) 1164, Masamune l.c. 426,
Anderson l.c. 311.
Shrub or tree to 20 m tall, 45 cm diameter. Bark smooth, grey-brown to grey-white, inner

bark pink to pale brown. Sapwood white to light yellow. Leaves each with 19 pairs of
leaflets; rachis 2.333.5 cm long, not winged; petioles 1.413.5 cm long. Leaflets coriaceous,
usually punctate; ovate to elliptic, 2.324.3 x0.87.8 cm; base attenuate, slightly asymmetric,
margin entire, flat to revolute, apex obtuse to caudate, usually not mucronulate; upper surface
usually glabrous to slightly sericeous, lower surface smooth to papillose, glabrous to slightly
sericeous or subvillose, domatia absent or in the form of 1 to many small sacs (or pockets);
lateral veins 0.23.7 cm apart, marginally looped and joined (less distinctly so in lower part
of leaflets), intercostal veins laxly reticulate, usually indistinct; petiolules absent to 0.9 cm
long. Inflorescences borne on leafless twigs, or axillary to pseudoterminal, unbranched to
basally branched; rachis 0.418 cm long, terete to slightly flattened, subsericeous to
subhirsute; first-order branches to 9 cm long; cymules in cincinnate or dichasial clusters, 26flowered; bracts and bracteoles triangular, bracts 0.52 mm long, bracteoles 0.20.9 mm
long; pedicels 1.87.3 mm long. Flowers 34.5 mm diameter, without scent; sepals 5, ovate,
2 outer smaller ones 0.92.8 x0.82.1 mm, 3 inner larger ones 1.43.4 x1.23.6 mm, margin
petaloid, white; petals 5, elliptic to obovate, 0.54 x0.32.2 mm, white, claw 0.21 mm high,
margin pilose, subglabrous inside, apex rounded to acute, scales 0.32 mm long, very much
broadened at apex, crest usually absent or present as a hairy flat part of the bifid scale apex;
disc complete, yellow; stamens 8, filaments 1.25 mm long, white, anthers 0.30.8 mm long,
pink; ovary 0.22 mm long, subhirsute, light green to white, style and stigma 0.12 mm long.
Fruits 0.71.5 x0.71.8 cm, glabrous; stalks 25 mm long; lobes 510 x49.5 mm. Seeds
obovoid, 59 x 4.17.3 mm; embryo 47.8 x 36 mm.
Vernacular names. Sabahbelimbing talun (Bajau), gulambir ayam, kayu lulus (Tidong).
Distribution. Myanmar, Thailand, Cambodia, Vietnam, Peninsular Malaysia, Sumatra, Java,
Borneo, Sulawesi. Rather common in Sabah and Sarawak; also in Kalimantan.
Ecology. In primary and secondary kerangas, mixed dipterocarp, and lower montane forests,
edges of forest, along riverbanks, road sides, and seashore, in deserted cultivated fields; on
different soil types including granitic sand, basalt, clay, loam on sandstone, limestone, and
water-logged sand. From sea level to 1700 m. Flowering in September to April; fruiting in
December to April.
Notes. Two forms of G. diplopetala are found in Borneo. In Sabah and northern parts of
Sarawak the leaflets are slender, rather symmetric, and glabrous; in southern parts of Sarawak
(Kuching area) the leaflets are very broad, asymmetric, bullate and are slightly villose below.
Both forms are united by transitional forms found in Peninsular Malaysia, Sumatra, and Java,
and therefore cannot be designated as two separate infraspecific taxa.
298
Fig. 9. Guioa diplopetala. A, fruiting leafy twig; B, petal; C, fruit. (A and C from Rahayu & Maskura 540, B from Beume A
765.)
299
3. Guioa koelreuteria (Blanco) Merr.
(J.G. Koelreuter, 17331806, German botanist)

Sp. Blanc. (1918) 241; Radlkofer in Engler l.c. (1933) 1172 (footnote); van Welzen l.c. (1989) 219, fig. 7b, 86, l.c. (1994)
376. Basionym: Sapindus koelreuteria Blanco, Fl. Filip. (1837) 289 (kolreuteria). Neotype (Merrill, 1918): Merrill Spec.
Blanc. 644, Philippines, Luzon, Rizal Province (PNH, destroyed; isoneotypes BM, F, K, L, NY, P, W).
Shrub or tree to 16 m tall, 20 cm diameter. Bark brown to dark grey, smooth to rough; inner
bark pink to red-brown, fibrous. Sapwood white to brownish, rings slightly visible. Leaves
each with 16 pairs of leaflets; rachis 2.420.5 cm long, not winged; petioles 18 cm long.
Leaflets usually asymmetric, thick-coriaceous, usually punctate, upper surface glabrous,
lower surface smooth, glabrous; domatia absent to many in the form of small sacs or
pockets; ovate, 2.817.8 x0.95.4 cm; base attenuate, margin entire, flat to revolute, apex
usually gradually acuminate to caudate, acumen acute to mucronulate; lateral veins 0.22.5
cm apart, marginally looped and joined; intercostal veins laxly reticulate, usually distinct;
petiolules absent to 0.7 cm long. Inflorescences axillary to pseudoterminal, unbranched to
basally branched; rachis 0.820.5 cm long, slightly flattened, brown-subsericeous; first-order
branches to 9.8 cm long; cymules cincinnate, 25(8)-flowered; bracts and bracteoles
triangular, bracts 0.51.5 mm long, bracteoles 0.21 mm long; pedicels 1.47 mm long.
Flowers 3.24 mm diameter; sepals 5, ovate, pink, 2 outer smaller ones 13.1 x13.3 mm, 3
inner larger ones 1.53.8 x1.24 mm, margin petaloid; petals 5, elliptic to obovate, 1.83.8x
12.5 mm, creamy white to reddish, gradually decurrent into the 0.41.3-mm-high claw,
pilose along margin and outside, subglabrous inside, apex obtuse to acute, scales 0.92.1 mm
long, not to hardly broadened at apex, crest present as a hairy, flat part of the bifid scale
apex; disc interrupted; stamens 8 (or 9), filaments 1.25 mm long, anthers 0.30.6 mm long;
ovary 0.22.4 mm long, subhirsute, style and stigma 0.12.7 mm long. Fruits 12.2 x12.3
cm, glabrous; stalks 25.5 mm long; lobes 6.513 x712 mm. Seeds globose to obovoid, 5.3
8.5 x 57.5 mm; arillode edible; embryo 57.8 x 3.86 mm.
Vernacular names. Sabahsaasa, saksah, sasah (east-coast Bajau).
Distribution. Borneo and the Philippines. Common on the islands northeast of Sabah.
Ecology. In primary and especially in secondary forest, ridge forest, in kerangas forest, along
the seashore, road sides, streams, edges of plantations. Soil: sand, gravel, limestone,
ultramafic. From sea level to 1350 m. Flowering in August to March; fruiting in March to
October.
Uses. The wood is used for agricultural implements and tool handles (Reyes in Desch, Mal.
For. Rec. 15 (1954) 526). Oil extracted from the seeds can be used to cure certain skin
diseases (Guerrero in Brown, Useful Pl. Philip. 2 (1950) 363.
4. Guioa pleuropteris (Blume) Radlk.
(Greek, pleuron = lateral outgrowth, pteron = wing; the winged petioles)
Fig. 10.
l.c. (1879) 10, in Engler l.c. (1933) 1164; Merrill l.c. (1921) 361, l.c. (1929) 175; Masamune l.c. 426; Anderson l.c. 311;
Corner l.c. 680; Yap l.c. 442; van Welzen l.c. (1989) 257, fig. 109, 110, l.c. (1994) 584. Basionym: Cupania pleuropteris
Blume, Rumphia 3 (1847) 158. Type: Korthals, s.n., Borneo (holotype L; isotypes L, W).
300
Fig. 10. Guioa pleuropteris. A, fruiting leafy twig; B, petal; C, fruit. (A and C from Davidson 1325; B from Maxwell 81-34.)
301
Shrub or tree to 30 m, 2070 cm diameter, without buttresses. Bark smooth to sometimes

irregularly fluted, usually dark brown, often mottled with white or grey spots; inner bark
white to dark brown, finely fibrous; cambium yellow to brown. Sapwood finely granular, soft
to hard, white to yellow, without odour or sap. Leaves each with (1)25(7) pairs of
leaflets; rachis 0.825.3 cm long, usually slightly winged, wings to 3 mm broad; petioles 0.6
9.3 cm long. Leaflets asymmetric at base and apex, subcoriaceous, usually punctate; upper
surface glabrous to subsericeous or hirsute, lower surface dull, papillose, subsericeous to
usually hirsute, domatia many, in the form of pockets or sacs; ovate (lower ones) to elliptic or
obovate (upper ones), 0.918.7 x0.58.3 cm; base acute, cuneate to attenuate, margin entire,
flat to revolute, apex obtuse to abruptly acuminate or cuspidate, often mucronulate; midrib
convex and raised below; lateral veins 0.24.9 cm apart, marginally looped and joined, less
distinctly so in the lower half of leaflets; intercostal veins laxly reticulate, usually distinct.
Inflorescences axillary to pseudoterminal, unbranched to basally branched; rachis 0.521.6
cm long, terete, usually brown hirsute, first-order branches to 13.8 cm long; cymules
cincinnate, 25-flowered; bracts and bracteoles triangular, bracts 0.73.8 mm long, bracteoles
0.21.5 mm long; pedicels 1.17 mm long. Flowers 34.2 mm diameter, fragrant; sepals 5,
ovate, green to tinged reddish or whitish, 2 outer smaller ones 0.72.8 x0.62 mm, 3 inner
larger ones 1.53.5 x0.93.3 mm, margin petaloid; petals 5, elliptic to obovate, 1.33.5 x0.7
2.2 mm, white, gradually decurrent into the 0.21 mm high claw, pilose at the margin and
outside, glabrous inside, apex rounded to acute, scales 0.82.2 mm long, apex not to hardly
broadened, crest yellow, usually developed as a pilose flat part of the bifid scale apex; disc
complete to interrupted by small gaps; stamens 8, filaments 1.35.1 mm long, white, anthers
0.20.7 mm long, pink; ovary green, 0.32.8 mm long, slightly rough-hairy, style and stigma
0.12.2 mm long. Fruits 11.9 x12.5 cm, glabrous; stalks 25 mm long; lobes 7.513 x6.5
13 mm. Seeds globose to obovoid, 5.59.7 x58 mm; arillode yellow to orange, edible, sour;
embryo 3.39 x 3.37 mm.
Vernacular names. Sabahandipatan, gulambir ayam (Dusun), gurujod, kanawit (Dusun
Kinabatangan), kangi runok (Dusun Tambunan), kejangan (Dusun Kundasan), mata pait
(Dusun Banggi), munggulan ayam (Dusun), pengkul (Dusun Kinabatangan), piri manok,
saasa (Bajau), tanggianuk, tanjsanuk (Dusun Kinarut), tangking manok (Dusun), tingir
manok (Bajau), tongianum (Dusun Kinarut).
Distribution. Myanmar, Cambodia, Vietnam, Thailand, east coast of Peninsular Malaysia,
Sumatra, Borneo, Philippines. Uncommon to rather common in Sabah and Sarawak; also in
Ecology. In primary and secondary mixed dipterocarp forest; edges of forests and
plantations; in open landscapes like cultivated land and belukar; in mossy forest; along
riverbanks, road sides, and on seashores; on different types of soil, including sandstone,
yellow sandy loam, black sand, ultramafic, and alluvial deposit. From sea level to 1800 m.
Flowering in August to May; fruiting throughout the year, but mainly in February to April.
Uses. The wood is used for torches (Radlkofer, 1913). The timber is durable and elastic, but
thin and in Indonesia, it is used for making handles of axes and shafts of wagons and plows
(Heyne, Nutt. Pl. ed. 2, 1, 1927, 1000). A decoction of the roots is used medicinally in NE
Pahang (Peninsular Malaysia) against fever and stomach ache. The Malay name pokok
seriawan burung probably refers to its medicinal use against sprue (Burkill, EPMP, ed. 2, 1,
1966, 1134). Also used to exterminate intestinal worms. The arillode is edible, but due to its
thinness offers no real food value.
302
5. Guioa pterorhachis Welzen
(Greek, pteron = wing, rhachis = main leaf axis)

Blumea 33 (1988) 419, pl. 12a, b, l.c. (1989) 269, fig. 114, l.c. (1994) 590. Type: Elmer 20268, Borneo, Sabah (holotype L;
isotypes BM, F, K, M, NSW, P, U).
Shrub or tree to 16 m tall, 1015 cm diameter. Bark smooth, flaky, white to brown, soft;
inner bark fibrous, pink to brown; cambium yellow. Sapwood white to brown. Leaves each
with 37 pairs of leaflets; rachis winged, wings to 4 mm broad; petioles 1.112.2 cm long.
Leaflets subcoriaceous, punctate; upper surface glabrous to puberulous on the midrib and
veins, lower surface dull, papillose, sparsely short-serriceous, domatia many, in the form of
sunken sacs; elliptic to obovate, 2.414.6 x14.6 cm; base often slightly asymmetric, acute to
attenuate or cuneate, margin entire, flat, apex gradually acuminate to cuspidate, sometimes
mucronulate; midrib hardly raised below, lateral veins 0.23.6 cm apart, marginally looped
and joined, less so in the lower half of leaflets, intercostal veins laxly reticulate, rather
indistinct. Inflorescences axillary to pseudoterminal, unbranched to basally branched; rachis
120.8 cm long, terete, subsericeous, first-order branches to 11.5 cm long; cymules
cincinnate, 25-flowered; bracts and bracteoles triangular, bracts 0.71.2 mm long; bracteoles
0.30.8 mm long; pedicels 27 mm long. Flowers 3.54 mm diameter; sepals 5, ovate, 2
outer smaller ones 11.9 x 0.81.5 mm, 3 inner larger ones 1.73 x1.22.7 mm, margin
petaloid; petals 5, elliptic to obovate, 1.12.8 x0.81.6 mm, white, claw 0.20.5 mm high,
margin pilose, glabrous on both sides, apex more or less acute, scales 1.12 mm long, crest
present as a pilose flat part of bifid scale apex; disc interrupted to complete; stamens 8,
filaments 25 mm long, anthers c. 0.3 mm long; ovary 0.31 mm long, sparsely rough-hairy,
style and stigma 0.11 mm long. Fruits 1.41.9 x 1.22.2 cm, glabrous; stalks 47 mm long;
lobes 8.511 x 812 mm. Seeds obovoid, 7.510.5 x69 mm; arillode edible, with yellow
exudate; embryo 6.89.2 x 57.5 mm.
Vernacular names. Sabahangil manuk (Malay), galid (Dusun Kinabatangan), garong
(Sungei), guruyod (Dusun Kinabatangan), lipang lipang (Sungei), sikip sikip (Dusun
Kinabatangan), tanggianggi, tengaranuk (Dusun Kinarut), tingir manuk (Bajau).
Distribution. Endemic to E Sabah (Sandakan and Tawau districts).
Ecology. In primary and secondary forests, along rivers and road sides, and on flat to
undulating country, on different soil types, from sea level to 500 m. Flowering in July to
November to January; fruiting in November to May.
Uses. Firewood.
6. Guioa pubescens (Zoll. & Moritzi) Radlk.
(Latin, pubescens = soft-hairy; the fruits)
l.c. (1878) 302, in Engler l.c. (1933) 1169; Merrill l.c. (1921) 361; Masamune l.c. 427; Backer & Bakhuizen f. l.c. 140;
Anderson l.c. 311; Corner l.c. 681; Yap l.c. 442; van Welzen l.c. (1989) 272, fig. 115, 116, l.c. (1994) 591. Basionym:
Sapindus pubescens Zoll. & Moritzi in Moritzi, Syst. Verz. (1846) 22, p.p. Type: Zollinger 1105, Java (holotype L; isotypes
A, BM, FI, P).
303
Tree to 25 m tall, 30 cm diameter; without buttresses. Bark smooth, hard, sometimes deeply
fissured to flaky, greyish white or greyish brown to dark brown; inner bark yellowish white to
reddish brown; cambium white. Sapwood soft to hard, white to ochre; heartwood red. Leaves
each with (1)26 pairs of leaflets; rachis 2.129.5 cm long, not winged; petioles 1.410.8
cm long. Leaflets coriaceous, not punctate; upper surface glabrous to sparsely shortsericeous, lower surface dull, papillose, slightly short-sericeous, domatia absent or present
as 1many small sacs; ovate to elliptic, often slightly sickle-shaped, 2.919.2 x0.87.2 cm;
base asymmetric, attenuate, margin entire, flat to revolute, apex gradually acuminate to
cuspidate or caudate, usually mucronulate; lateral veins 0.33.3 cm apart, marginally looped
and joined (less so in the lower third of leaflets), intercostal veins laxly reticulate, rather
indistinct; petiolules to 1 cm long. Inflorescences axillary to pseudoterminal, unbranched to
basally branched; rachis 1.424.2 cm long, terete, brown-sericeous; first-order branches to
9.3 cm long; cymules cincinnate, 24-flowered; bracts and bracteoles triangular, bracts 0.6
1.8 mm long, bracteoles 0.21.2 mm long; pedicels 1.88 mm long, sericeous. Flowers 3.5
4.5 mm diameter; sepals 5, ovate, 2 outer smaller ones 12.8 x0.82.1 mm, 3 inner larger
ones 1.43.3 x1.33.1 mm, margin petaloid; petals 5, obovate, 1.93.4 x0.71.8 mm, white to
yellow, gradually decurrent into the 0.31.2 mm-long claw, pilose at the margin and outside,
glabrous inside, rounded to acute at apex, scales 1.12 mm long, apex not to hardly
broadened, crest present as a pilose flat part of the bifid scale apex; disc interrupted; stamens
8, filaments 1.95.2 mm long, white, anthers 0.20.4 mm long, pink; ovary 0.32 mm long,
densely rough-hairy, yellowish green, style and stigma 0.12.5 mm long. Fruits 11.5 x1
1.9 cm, sparsely sericeous, glabrescent; stalks 1.55 mm long; lobes 710 x 6.510 mm.
Seeds globose to obovoid, 6.810 x 5.87.5 mm; embryo 5.98.3 x4.96 mm.
Vernacular names. Sabahangir manuk (Kedayan), geronok (Dusun Tambunan), saasa,
saksah (Bajau). Sarawakrutan berangat (Kelabit).
Distribution. Peninsular Malaysia, Singapore, Sumatra (including Banka), W Java, KarimunDjawa Is., Borneo (above equator), Philippines (Palawan). Uncommon to rather common in
Sabah and Sarawak; also in Brunei.
Ecology. In primary and secondary mixed dipterocarp forests, kerangas, peat swamp forests,
beach forest, on different soil types including sand, sandy loam, sandstone, limestone,
ultramafic, and well-drained peat; from sea level to 1800 m. Flowering in August to
February; fruiting in January to April.
Uses. The wood is used as a construction timber, although it is brittle and the stem diameter
is too small for planks (Heyne, Nutt. Pl. Ned. Indie ed. 2 (1927) 1001).
10. HARPULLIA Roxb.
(Latin form of the Bengal plant-name harpulli)
P.W. Leenhouts & M. Vente

Fl. Ind. 2 (1824) 441; Merrill, EB (1921) 362; Radlkofer in Engler, Pflanzenr. 98 (19331934) 1433; Masamune, EPB
(1942) 427; Backer & Bakhuizen f., FJ 2 (1965) 142; Anderson, CLST (1980) 311;
Leenhouts & Vente, Blumea 28 (1982) 1, FM 1, 11 (1994) 598; Whitmore & Tantra, CLS (1986) 214; Yap, TFM 4 (1989) 443;
Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 311.
304
Shrubs or medium-sized trees, dioecious. Indumentum of solitary and stellate tufts of simple
hairs and glandular hairs; glandular scales absent. Leaves paripinnate, each with 19 pairs
of leaflets, without pseudostipules; petiole and rachis winged. Leaflets alternate or opposite,
not papillose beneath; margin entire (Malesian species). Inflorescences usually thyrses,
axillary, pseudoterminal, or terminal, solitary or borne in tufts on leafless twigs or stems;
bracts and bracteoles usually caducous. Flowers unisexual, radially symmetrical; sepals 5,
free, imbricate, equal or the outer two sometimes slightly smaller, not petaloid, not ciliate,
glandular hairs mainly present along the entire margin; petals 5, longer than the sepals,
distinctly clawed with a pair of auricles above the claw, or sessile with a broad or narrow
base and without auricles and scales, entire; disc complete to divided into 5 lobes, without
appendages; stamens 58, in male flowers exserted, filaments glabrous, anthers basifixed,
base cleft for up to one-fifth the length, latero-introrse; ovary 23(4)-locular, sessile or
short-stalked, hairy, ovules 1 or 2 per locule, style apical, shorter to much longer than the
ovary, slender, often hooked and twisted at the upper part, lower part hairy, stigmatic lines
usually extended to slightly above the base. Fruits loculicidal capsules, usually short-stalked,
not winged, 23-lobed, the lobes erect to spreading, inflated, rounded; fruit wall parchmentlike to woody. Seeds with a thin-crustaceous testa; arillode entire and without appendages,
restricted to a narrow annular sarcotesta around the hilum, or composed of a basal
sarcotestal part covering half the seed, and an upper free arillodal part, reaching to near the
apex; hilum covering less than one-sixth of the seed; embryo straight; cotyledons superposed,
about equal.
Distribution. About 26 species; from Sri Lanka and India through SE China and Malesia to
Australia, New Caledonia, and Tonga (Av in van Balgooy, Pacific Plant Areas 4 (1984) 238,
239). Two species are known from Sabah and Sarawak.
Ecology. In primary and sometimes secondary rain forests, sometimes growing in low or
open forest and in coastal shrubby vegetation; from sea level to 2000 m. The seeds are
probably mainly dispersed by birds, possibly also by mammals and lizards.
Key to Harpullia species
Petals clawed and auricled, membranous. Arillode restricted to a sarcotesta ring around the
hilum......................................................................................................................1. H. arborea
Petals not clawed, without auricle, thin-fleshy. Arillode completely or nearly completely
enveloping the seed.........................................................................................2. H. cupanioides
1. Harpullia arborea (Blanco) Radlk.
Fig. 11AC.
(Latin, arbor = tree)
Sitzungsber. Math.-Phys. Cl. Knigl. Bayer. Akad. Wiss. Mnchen 16 (1887) 404, in Engler, Pflanzenr. 98 (1934) 1456;
Merrill, Sp. Blanc. (1918) 243, l.c. (1921) 362; Masamune l.c. 427; Backer & Bakhuizen f. l.c. 142; Anderson l.c. 311;
Leenhouts & Vente l.c. (1982) 11, l.c. (1994) 601, fig. 45, 46af.; Whitmore
305
Fig. 11. Harpullia arborea (AC); H. cupanioides (DE). A, flowering and fruiting leafy twig: B, petal: C, ovary; D, petal;
E, ovary. (A from PNH 12482, BC from van Beusekom et al. 2862, DE from Waterhouse 36 B.)
306
& Tantra l.c. 214; Yap l.c. 444; Whitmore, Tantra & Sutisna l.c. 311. Basionym: Ptelea arborea Blanco, Fl. Filip. (1837)
63. Neotype (Merrill, 1918): Merrill Sp. Blanc. 339, Philippines, Luzon, Bulacan Province, Angat (A; isoneotypes BM, BO,
K, L, NSW, P, US, W). Synonym: Harpullia tomentosa Ridley, Kew Bull. (1933) 192, Masamune l.c. 427.
Shrub or tree to 33 m tall, 60 cm diameter, but usually much smaller. Young parts more or
less densely hirsute. Leaves each with 26 pairs of leaflets; petioles 4.515 cm long; all axes
hairy, glabrescent. Leaflets herbaceous, glabrous except for the midrib above, glabrous or
sparsely hairy on midrib and lateral veins beneath; ovate to elliptic, 5.530 x210 cm, base
asymmetric with the upper half cordate or both sides acute, or symmetric and acute to
rounded, apex acute to rounded or acuminate, acumen usually short, acute; midrib flat to
slightly raised above, lateral veins 0.752.25 cm apart, flat above, intercostal veins
inconspicuous; petiolules 38 mm long. Inflorescences axillary or borne on leafless twigs or
stems, hairy, to 17 cm long and branched only near the base into several axes of about the
same length, or to 35 cm (in fruit to 60 cm) long with short branches. Flowers: sepals all
equal, ovate to obovate, 510.5 x35 mm, sparsely glandular hairy especially along the
margin; petals membranous, clawed, obovate-oblong, 817 x310 mm, white, glabrous
outside or sparsely hairy on the claw and central lower half of the blade, margin often ciliate
mainly in the basal part, often sparsely hairy inside, claw 37 mm long; disc sparsely to
densely hairy; stamens 5(7), filaments 1017 mm long, anthers 22.5 mm long; ovary 2(4)locular, style 1417 mm long. Fruits 931 x2765 mm; stalks to 4.5(7) mm long; lobes
spreading, slender ellipsoid to globose; fruit wall thin, chartaceous to woody, prominently
veined to smooth, red, fairly densely to sparsely hairy outside, reddish, sparsely hairy to
glabrous inside. Seeds 1 or 2 per locule, black, mahagony-brown, or dark purple; arillode
restricted to a ring around the hilum, to 2.5 mm wide, orange.
Vernacular names. Sabahbambuakat, tambuakat (Dusun, Kinabatantan). Sarawakapoh
(Iban), arip, ensiru (Bidayuh), sakubong (Melanau).
Distribution. Sri Lanka, SE India, Assam, Thailand, Vietnam, Malesia, the Solomon Is.,
Samoa, Tonga and N Queensland. In Sabah and Sarawak, locally common; also in Brunei
and Kalimantan.
Ecology. Usually in well-drained primary and secondary lowland and hill mixed dipterocarp
forest and lower montane forest, from sea level to 1200 m. On various types of soil, including
clay, loam, sand, limestone and volcanic. Flowering and fruiting throughout the year.
Uses. The bark is used as a fish poison. A watery exudate of the bark and sometimes the
fruits is used for washing, to keep away leeches, or is drunk to allay pain. The timber is of
good quality. The oil pressed out of the seeds is used as an anti-rheumatic (Brown, Useful Pl.
Philip. 2 (1950) 363; Desch, Malayan Forest Rec. 15 (1954) 528).
2. Harpullia cupanioides Roxb.
Fig. 11DE.
(Greek, cupanioides = resembling Cupania)
Fl. Ind. 2 (1824) 442; Merrill l.c. (1921) 362; Radlkofer in Engler l.c. (1934) 1444; Masamune l.c. 427; Backer & Bakhuizen
f. l.c. 142; Leenhouts & Vente l.c. (1982) 26, l.c. (1994) 606; Whitmore & Tantra l.c. 214; Yap l.c. 444; Whitmore, Tantra &
Sutisna l.c. 311. Type: Roxburgh, s.n., 1813, Bangladesh (holotype K; isotypes A, BO). Synonym: Harpullia confusa
Blume, Rumphia 3 (1847) 176, Masamune l.c. 427.
307
Shrub or tree to 20(40) m tall, 40 (100) cm diameter; buttresses to 2 m high and 2 m wide.
Young parts densely appressed short-hairy, glabrescent. Leaves each with (1)36(7) pairs
of leaflets; petioles to 20 cm long; rachis glabrous or sparsely puberulous. Leaflets
herbaceous to chartaceous, glabrous or very sparsely hairy on midrib and on lateral veins
below; ovate (lower ones) or elliptic to obovate (upper ones), 536 x215 cm; base
symmetric to asymmetric, acute or the broader half (rarely both halves) rounded, not or only
slightly decurrent, apex acute, rounded or gradually to abruptly acuminate, acumen short to
rather long, rounded, obtuse or acute; midrib usually slightly raised or sunken, lateral veins
0.753 cm apart, slightly sunken above, intercostal veins feeble or absent; petiolules 212
mm long. Inflorescences axillary, pseudoterminal or terminal, erect or pendulous, 585 cm
long, unbranched or variously branched, often few-flowered, usually tardily hairy on the
upper parts; bracts small, solitary or in clusters of threes. Flowers fragrant; sepals elliptic to
suborbicular, 36 x2.54.5 mm, persistent in fruit; petals not clawed, thin-fleshy, oblongobovate to oblanceolate, 510 x23 mm, white to creamy, yellow, greenish, pink, glabrous;
disc complete, low, short-velvety; stamens 5 (or 6), filaments 2.53.5 mm long, white,
anthers 1.53.5 mm long, yellowish white, greyish yellow or dark mauve; ovary 2-locular,
light green, yellow or reddish brown, style 1.756.5 mm long, light green, stigma whitish.
Fruits subreniform, transversely ellipsoid, broadly ovoid, obovoid, or globose, 1220 x12
32.5 mm; base rounded to truncate to slightly concave, apex slightly concave to obtuseangular, apiculate; fruit wall coriaceous to woody, red, variably hairy outside, early to late
glabrescent, glabrous to laxly long-hairy or glandular hairy inside; stalks 13 mm long. Seeds
shiny brown to black; arillode bright glossy red, completely or nearly completely enveloping
the seed.
Vernacular names. Sabahlokud (Dusun Ranau), mampongoh (Murut).
Distribution. S China, Assam, the Andaman Is., Bangladesh, Myanmar, Thailand, IndoChina, throughout Malesia, and the Northern Territory of Australia. Locally common in
Ecology. In primary and secondary mixed dipterocarp forest, tidal forest, scrubland, and open
places; from sea level to 1800 m. On different types of soil, including rocky, sandy, clay,
loam, limestone, and volcanic. Flowering in (January)AprilJuly(December); fruiting in
(January)AprilOctober(December).
Uses. The wood is used for making charcoal and as firewood; the bark is reportedly used as a
fish poison.
11. LEPISANTHES Blume

(Greek, lepis = scale, anthos = flower;
the presence of scales on the petals)
P.W. Leenhouts
Bijdr. Fl. Ned. Ind. (1825) 237; Merrill, EB (1921) 358; Ridley, FMP 1 (1922) 492; Radlkofer in Engler,
Pflanzenr. 98 (1932) 726; Masamune, EPB (1942) 427; Backer & Bakhuizen f., FJ 2 (1965) 134; Leenhouts,
Blumea 17 (1969) 33, FM 1, 11 (1994) 627; Anderson, CLTS (1980) 311; Whitmore &
308
Tantra, CLS (1986) 214; Yap, TFM 4 (1989) 444; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 312.
Synonyms: Erioglossum Blume l.c. (1825) 229, Merrill l.c. (1921) 358, Ridley l.c. 491, Radlkofer in Engler l.c.
(1932) 692, Masamune l.c. 429, Corner, WSTM 3rd ed. 2 (1988) 679; Aphania Blume l.c. (1825) 236, Merrill
l.c. (1921) 358, Ridley l.c. 492, Radlkofer in Engler l.c. (1932) 699, Masamune l.c. 425, Backer & Bakhuizen f.
l.c. 134; Otophora Blume, Rumphia 3 (1847) 142, Merrill l.c. (1921) 358, Ridley l.c. 492, Radlkofer in Engler
l.c. (1932) 753, Masamune l.c. 429.
Trees or shrubs, mostly monoecious. Indumentum of solitary, simple hairs. Leaves pari- or
imparipinnate, sometimes simple, with 1 to more than 40 pairs of leaflets; petiole and/or
rachis winged or not; with or without pseudostipules. Leaflets opposite or alternate, not
papillose beneath, margin entire. Inflorescences terminal, axillary, or borne on leafless twigs
or stems. Flowers unisexual, radially symmetrical; sepals 4 or 5 (rarely 3 or 6), free,
imbricate, outer 2 (or 1) mostly distinctly smaller, mostly at least inner ones partly petaloid,
entire or partly denticulate; petals 4 or 5 (rarely 3, 6, or 7), shorter to longer than sepals,
mostly distinctly clawed, scale mostly well-developed, sometimes only represented by a hairy
rim or a pair of small auricles, crested or not; disc interrupted or not, mostly slightly lobed;
stamens (4)8(18), in male flowers not to distinctly exserted, filaments nearly always hairy
except for the base or the apex, anthers hairy or glabrous; ovary sessile to short-stalked, lobed
or not, 23(4)-locular, style apical, about as long as the ovary or stigma sessile, stigma
globose or dome-shaped, slightly lobed, ovules 1 per locule, sub-basal to median, placenta
with an obturator. Fruits sessile to short-stalked, not to distinctly lobed, drupaceous, smooth
or slightly warty outside, hairy to glabrous; fruit wall thin- or thick- fleshy, hairy or glabrous
inside; septa mostly complete, sometimes interrupted or represented merely by a rib. Seeds
with a shiny brown to black, glabrous or sometimes hairy testa; without arillode or
sarcotesta.
Distribution. Twenty four species; tropical Africa, Madagascar, S and SE Asia from Sri
Lanka to Hainan, Malesia, and NW Australia. In Sabah and Sarawak, 11 species.
Ecology. In Borneo, most species are found in primary and secondary forests, especially in
open places, escarpments, steep slopes, riverbanks, forest edges, clearings, and other types of
open secondary vegetation; from sea level to c. 2000 m. The aggregated and sweet-scented
flowers are probably pollinated by insects. The fruits, of at least those species with a fleshy
fruit wall, are dispersed by animals.
Taxonomy. Leenhouts (1969) combined several genera were into Lepisanthes. He
subdivided the genus as follows:
Subgenus Lepisanthes
Section Lepisanthes (L. tetraphylla)
Section Hebecoccus (L. falcata subsp. borneensis)
Subgenus Otophora
Section Otophora (L. amoena, divaricata, kinabaluensis, multijuga)
Section Pseudotophora (L. fruticosa)
Section Anomotophora (L. alata, ramiflora)
Subgenus Erioglossum (L. rubiginosa)
Subgenus Aphania (L. senegalensis)
Uses. A few species have some value as ornamental trees. The wood of several species is
used, but only a few reach a sufficient size to have some value as timber trees. Different parts
309
of several species are of medicinal value. The fruits of some species are edible, but only L.
fruticosa and L. alata are planted for their fruits.
Key to Lepisanthes species
1. Pseudostipules (a pair of leaflets attached at the base of the petiole) absent.......................2
Pseudostipules present.........................................................................................................5
2. Leaf venation closed (lateral veins distinctly looping and joining). Sepals subglabrous, as
long as the petals. Petals usually dark red. Fruits glabrous, lobed..........10. L. senegalensis
Leaf venation open (lateral veins not looping and joining) (if fruits subglabrous) or open
to closed (if fruits densely hairy). Sepals sericeous, shorter than the petals. Petals greenish
white to yellowish or pink. Fruits densely hairy or subglabrous, not or slightly (rarely
distinctly) lobed...................................................................................................................3
3. Leaf venation open. Fruits dark purple to black, distinctly lobed, lobes spreading,
subglabrous.................................................................................................9. L. rubiginosa
Leaf venation open to closed. Fruits bright orange, yellowish or greyish pink, not or only
slightly lobed, or if distinctly lobed then the lobes erect and densely hairy........................4
4. Leaflets mostly with glandular-pitted warts. Petal scales not crested. Dry fruit coarsely
rugose................................................................................................................4. L. falcata
Leaflets without glandular-pitted warts. Petal scales mostly crested. Dry fruit smooth........
..................................................................................................................11. L. tetraphylla
5. Petiole and rachis winged....................................................................................................6
Petiole not winged, rachis very rarely winged.....................................................................7
6. Twigs glabrous. Leaflets glabrous, mostly lanceolate, smaller and narrower rarely more
than 20 x 4 cm, base acute or sometimes rounded. Ovary 3(4)-locular. Fruits 3-locular,
trigonous-obovoid................................................................................................1. L. alata
Twigs sparsely hairy. Leaflets sparsely hairy, oblong or sometimes slightly obovate,
larger and broader to 33 x 10.5 cm, base cordate. Ovary 2-locular. Fruits incompletely 2locular, transversely ellipsoid........................................................................8. L. ramiflora
7. Fruits distinctly lobed, lobes spreading...............................................................................8
Fruits not or slightly lobed, if distinctly lobed then the lobes more or less erect.................9
8. Leaves each with 714 pairs of leaflets. Twigs up to 1 cm diameter, tomentose..................
...............................................................................................................6. L. kinabaluensis
Leaves each with (15)3040 pairs of leaflets. Twigs 1.52 cm diameter, glabrous............
......................................................................................................................7. L. multijuga
9. Leaves paripinnate or with a reduced terminal leaflet, each with 18(14) pairs of leaflets.
Leaflets densely finely-pitted underneath. Inflorescences rarely terminal.....5. L. fruticosa
Leaves mostly imparipinnate, each with (3)742 pairs of leaflets. Leaflets with pitted
warts resembling small white scales. Inflorescences terminal and axillary.......................10
310
10. Leaf venation open..........................................................................................2. L. amoena

Leaf venation closed....................................................................................3. L. divaricata
1. Lepisanthes alata (Blume) Leenh.
Fig. 12G.
(Latin, alatus = winged; the petiole and rachis)
l.c. (1969) 80, l.c. (1994) 646; Anderson l.c. 311. Basionym: Otophora alata Blume l.c. (1847) 145, Merrill l.c. (1921) 358,
Radlkofer in Engler l.c. (1932) 768, Masamune l.c. 429, Backer & Bakhuizen f. l.c. 135. Lectotype (Leenhouts, 1969):
Korthals, s.n., S Borneo (L). Synonym: Otophora edulis C. E. C. Fischer, Kew Bull. (1932) 178, Radlkofer in Engler l.c.
(1934) 1493, Masamune l.c. 430.
Tree or shrub to 15 tall m, 30 cm diameter. Twigs 0.81.2(1.5) cm diameter, glabrous.

Leaves paripinnate, each with 35(13) pairs of leaflets; petioles 1.57.5(25) cm long;
petiole and rachis winged, wings 38 mm wide; pseudostipules oblique-ovate, 13(8.5)
x0.82.2(5) cm, base deeply cordate, apex obtuse, sometimes acuminate, pinnately veined.
Leaflets opposite to alternate, sessile or petiolules to 2 mm long, glabrous, thin-chartaceous,
greyish to blackish brown above, brown to greyish green beneath; lanceolate (exceptionally
linear-lanceolate or oblong to obovate-oblong, lower ones sometimes ovate-lanceolate), 10
20(45) x 1.84(8) cm; base mostly hardly asymmetric, acute and sometimes attenuate, in
asymmetric leaflets one half or both halves rounded, apex attenuate long-acuminate, acute;
midrib mostly sharply raised beneath; lateral veins 11.5(2) cm apart, curved, at least those
in the upper half of the leaflet looping and joining near the margin. Inflorescences often
drooping, purple to reddish brown, apparently mostly unisexual; male inflorescences usually
axillary, narrowly thyrsoid, unbranched, 2025(40) cm long; flowers in scattered, sessile
fascicles of 35(7) along the rachis; pedicels 27 mm long; female inflorescences axillary in
the axils of lower leaves or on leafless twigs or stems, strongly branched at least near the
base of the rachis, to 45 cm long; flowers scattered, and mostly solitary along the rachis;
pedicels 613 mm long. Flowers dark wine-red to purple; sepals sparsely glandular-ciliolate,
obovate-orbicular, 2.54 x23 mm, outer sepals slightly smaller than inner ones, the inner
sepals partly petaloid with crenulate margin; petals sessile or with a claw of c. 0.5 mm,
sparsely, partly glandular, ciliolate, glabrous, or rarely the basal half sparsely appressed shorthairy outside, blade suborbicular, 24 x 2.53 mm, scale erect, slightly hood-shaped, 0.20.3x
as long as the blade, glabrous or densely ciliolate; disc glabrous, pink; stamens 8, anthers c.
1.8 mm long, subglabrous, yellow; ovary ellipsoid, 3(4)-locular, pale mauve, style very
short, stigma dome-shaped to flat, slightly 3-lobed, white. Fruits shortly stalked, trigonousobovoid, 2.54 x2.23 cm, apiculate, dark brownish purple to nearly black when ripe; pulp
rather thick, fleshy, white; pedicels patent, slender, to 2 cm long. Seeds ellipsoid, to 2.5 x1.5
cm; hilum rhomboid, c. 6 x 5 mm.
Vernacular names. Sabahsintatanod (Dusun Kinabatangan). Sarawakenkelili (Iban),
peraju (Bidayuh Bau), sipaju (Bidayuh Padawan), sokungu (Bidayuh Sadang).
Distribution. Peninsular Malaysia, Java (probably only naturalized), and Borneo.
Uncommon in Sabah and Sarawak; also in Brunei.
Ecology. In lowland mixed dipterocarp forest, on riverbanks on clay soil, to 450 m.
Flowering in August to May; fruiting in August to April. Ants may live under the stipules.
311
Fig. 12. Lepisanthes amoena (AC); L. senegalensis (D); L. fruticosa (E); L. rubiginosa (F); L. alata (G); L. falcata subsp.
borneensis (H); L. multijuga (I); L. tetraphylla (J). A, leaf; B, detail of lower surface of leaflet; CJ, fruits. (AB from
Danser 5474, C from Wirawan 63, D from Kostermans 60, E from Nooteboom 1323, F from Kartawinata 683, G from S.
40142, H from Podzorski SMHI 618, I from Kokawa & Hotta 2329, J from SAN 25591.)
312
Uses. The fruits and possibly also the seeds are edible. In Sumatra, Java, and Borneo
sometimes grown as a fruit tree (Heyne, Nutt. Pl. Ned.-Indi, ed. 2 (1927) 990; Jansen et al.
PROSEA 2 (1991) 343).
Notes. L. alata is on the whole a rather uniform species. Only a few specimens from Borneo
show some characters which deviate considerably from the mean. Among these are Endert
1702 and Jaheri 1693 from Kalimantan, characterized by their rather large (1523 x57.5
cm), relatively wide (length c. 3 times the width), and distinctly obovate leaflets that have
rather distant lateral veins (1.52 cm apart); Hotta 12660 from Brunei, has a leaf with 13
pairs of very long and narrow leaflets (33 x2.2 cm, length c. 14 times the width), a petiole and
rachis that is rounded or flat above, and very large stipules (8.5 x5 cm); and Orolfo 1319
from Sabah, (type of Otophora edulis), which has rather narrow leaflets (length 8 times the
width) with nearly perpendicular lateral veins, distinctly clawed petals which are hairy on the
outside of the scale, thick ovary wall, and large fruits (c. 4 x 3 cm) with a thick, fleshy pulp.
2. Lepisanthes amoena (Hassk.) Leenh.
Fig. 12AC.
(Latin, amoenus = lovely, charming; the habit)
l.c. (1969) 71, l.c. (1994) 636; Anderson l.c. 311; Whitmore & Tantra l.c. 214; Yap l.c. 444; Whitmore, Tantra & Sutisna l.c.
312. Basionym: Melicoccus amoenus Hassk., Flora 25, 2 (1842) Beibl. 39. Type: Reinwardt, s.n., Java (L). Synonyms:
Otophora amoena (Hassk.) Blume l.c. (1847) 142, Merrill l.c. (1921) 358, Radlkofer in Engler l.c. (1932) 771, Masamune
l.c. 429, Backer & Bakhuizen f. l.c. 135; Otophora spectabilis Blume l.c. (1847) 142, Merrill l.c. (1921) 359, Masamune l.c.
431; Otophora imbricata Blume l.c. (1847) 144, Merrill l.c. (1921) 358, Ridley l.c. 495, Radlkofer in Engler l.c. (1932) 773,
Masamune l.c. 430; Otophora pubescens Blume l.c. (1847) 145, Merrill l.c. (1921) 359, Radlkofer in Engler l.c. (1932) 770,
Masamune l.c. 430; Otophora cordigera Radlk., Sapind. Holl.-Ind. (1879) 85, Merrill l.c. (1921) 358, Radlkofer in Engler
l.c. (1932) 770, Masamune l.c. 430.
Tree to 10 m tall, 15 cm diameter, or shrub to 6 m. Twigs 0.81(1.2) cm diameter, sparsely

to rather densely, fulvous short-hairy when young, mostly early glabrescent. Leaves
imparipinnate, each with 742 pairs of leaflets, axial parts usually glabrous to thinly
tomentose or sometimes densely fulvous-tomentose or rarely hirsute, glabrescent; flush white
over pink to salmon or coral-red; petioles 19 cm long; pseudostipules orbicular, ovate, or
transversely elliptic, usually asymmetric, 16 x 0.86 cm, base truncate to deeply cordate,
apex rounded, obtuse, acute, or shortly and broadly acuminate, in extremely asymmetric ones
sometimes with an additional lateral apex, pinnately or reticulately veined, usually connected
with the normal leaflets by a series of intermediates. Leaflets opposite to alternate, subsessile,
thin-chartaceous or parchment-like, greenish grey to dark brown above, yellowish to redbrown beneath; glabrous or the midrib densely fulvous- to ferrugineous-tomentose above,
sparsely hirsute or, rarely thinly hairy beneath, both surfaces or the upper surface only often
with scattered, minute, glandular-pitted warts; linear or sometimes ovate-lanceolate, 722.5x
1.25 cm; base oblique or not, obtuse to subcordate, apex obtuse to attenuate-acuminate,
acumen short to long, obtuse to acute; lateral veins 0.52 cm apart, curved, only the upper
ones loopping and joining at some distance from the margin. Inflorescences terminal and
axillary, pyramidal, up to 60 cm long, sparsely to densely fulvous short-hairy; axial parts
often reddish; rachis and main branches sharply triangular; branches nearly transverse to
ascending, to 30 cm long, the lower ones often with some short, spreading branches; cymes
sessile to short-stalked, 1several-flowered; pedicels 24 mm long. Flowers mostly white,
313
sometimes creamy to yellow or pink to red, not scented; sepals red, outer two ovate to
oblong, 1.53.5 x1.22 mm, inner 3 more or less orbicular, 2.54 x 23.5 mm; thinly
appressed short-hairy to glabrous outside, glabrous to sparsely appressed short-hairy in the
basal half inside, towards the base usually sparsely partly glandular ciliolate; petals white or
red, sparsely to densely appressed long-hairy outside, variably hairy inside, blade subdeltoid
to suborbicular, to 1.5 x 1.2 mm, densely to very sparsely ciliate at base, with 2 incurved,
more or less joined basal lobes, claws to c. 0.6 mm long, densely to very sparsely ciliate; disc
glabrous, orange or yellow; stamens 79, filaments white or reddish, anthers 1.51.8 mm,
densely hairy, yellowish; ovary 3-locular, densely velvety to nearly glabrous, cream to
reddish, style 1.5 mm long. Fruits slightly 3-lobed, widest in or above the middle, 22.5x
2.22.8 cm (fresh c. 2.5 x3 cm), apiculate by the style base, scurfy or sometimes hardly
scurfy, thinly short-hairy to glabrous, light green to spotted brown when unripe, brownish or
purple when ripe; pulp yellowish to white. Seeds obliquely ellipsoid, hilum orbicular to
lanceolate, small.
Distribution. Sumatra, Peninsular Malaysia, Borneo, W Java, Timor (one collection).
Common in Sabah, less so in Sarawak.
Ecology. In primary and secondary mixed dipterocarp forests, scrub- or bamboo-forest, often
along rivers, in well-drained as well as in swampy or periodically inundated localities; on
acid, basic loam, clay, or sandy soils; from sea level to 1650 m. Flowering and fruiting
throughout the year. The often large pseudostipules, appressed to the twigs, are often used as
shelter by ants.
Uses. In Java, sometimes planted as an ornamental tree. The bark as well as the young leaves
contain saponin and are applied as a poultice for treating ulcers (Borneo). The wood is
reported to be very hard and has been used for making hooks to catch crocodiles. The fruit
pulp is sweet and edible (Heyne, Nutt. Pl. Ned.-Indi, ed. 2, 1927, 990; Burkill, EPMP (1935)
1613; Jansen et al., PROSEA 2 (1991) 343).
Taxonomy. L. amoena is a variable species but, apart from one race in the east coast of
Sumatra, no clearly delimited infraspecific taxa can be distinguished. In Borneo, three rather
distinct forms or races can be distinguished, but outside Borneo, the differences often
become less clear. These races are:
amoena leaves with rather many pairs of leaflets; leaflets not very narrow, linear, often
greyish green above, reddish brown beneath. Throughout the area of distribution.
imbricata leaves with a few pairs of leaflets; leaflets large, relatively wide, brown,
margin not parallel, base obtuse. The presence of a series of intermediates between
pseudostipules and leaflets in this race is unique. E Borneo and Java.
pubescens leaves with many pairs of leaflets; leaflets small, narrow-linear, leaflet base
cordate, mostly dark brown above, midrib usually hirsute beneath. W Borneo and Sumatra.
3. Lepisanthes divaricata (Radlk.) Leenh.
(Latin, divaricatus = spreading; the inflorescence branches)
Fig. 13.
l.c. (1969) 72, l.c. (1994) 638; Anderson l.c. 312; Whitmore, Tantra & Sutisna l.c. 312. Basionym: Otophora divaricata
Radlk., Fedd. Repert. 18 (1922) 338, in Engler l.c. (1932) 758, Masamune l.c. 430.
314
Fig. 13. Lepisanthes divaricata forma divaricata. A, flowering leafy twig: B, female flower; C, ovary in longitudinal section;
D, petal from inside, E, sepal from inside; F, staminode from outside; G, fruit. (AF from S. 17527, G from Rehal 13015.)
315
Type: Anon., s.n., Borneo, Sarawak (holotype M; isotype SAR). Synonym: Otophora
pyramidalis Radlk. l.c. (1922) 338, in Engler l.c. (1932) 759, Masamune l.c. 431.
Small tree to 7 m tall, or shrub. Twigs 313 mm diameter, glabrous to rather densely
ferrugineous short-hairy. Leaves pari- or imparipinnate, each with (3)711(16) pairs of
leaflets, 2540 cm long; axial parts ferrugineous or fulvous short-hairy, glabrescent; petioles
26(17) cm long; pseudostipules suborbicular to elliptic, often slightly asymmetric, (0.4)1
1.8 x (0.3)1.22.2 cm, base cordate to cuneate, apex rounded to acute, reticulately,
palmately, or pinnately veined. Leaflets opposite to alternate, sessile or with petiolules up to 2
mm long, stiff-chartaceous, mostly greenish grey to blackish above, light to dark brown
beneath, glabrous or ferrugineous-tomentose on the midrib on both sides, with pitted warts
beneath; elliptic to lanceolate, 720 x27 cm; base slightly cordate or rounded to cuneate,
asymmetric, long-attenuate in terminal leaflet, apex attenuate, acuminate, acute to obtuse;
lateral veins 12.2 cm apart, nearly straight, distinctly looping and joining at some distance
from the margin. Inflorescences terminal or in the axils of upper leaves, pyramidal, to more
than 40 cm long, glabrous to fulvous short-hairy, repeatedly branched, main branches patent,
up to 25 cm long; cymes short-stalked to sessile, few- to many-flowered; pedicels slender, 1
1.2 mm long. Flowers orange-yellow; sepals sparsely (outer) to densely (inner) partly
glandular ciliolate, outer two broad-ovate to obovate, 1.82.2 x1.2 mm, inner ones
suborbicular to ovate, 1.83.5 x1.82.2 mm; petals distinctly clawed, long-hairy, blade
transversely subelliptic, 0.71.5 x1.22 mm, auricled at base; disc glabrous or fairly densely
short-hairy; stamens 58, anthers c. 1 mm long, rather densely woolly; ovary 23-locular,
glabrous to thinly appressed hairy; stigma sessile or on a short style. Fruits slightly 23lobed, 23 x23 cm, hardly pointed, smooth and glabrous; yellow to pale brown. Seeds
obovoid, hilum elliptic, c. 3 mm long.
Distribution. Endemic to Sarawak and W Kalimantan.
Ecology. Disturbed kerangas or swamp forest behind the mangrove, to 400 m. Flowering in
October to July; fruiting in October to January.
Key to forms
Leaves imparipinnate, each with 6 or more pairs of leaflets. Pseudostipules orbicular, never
pinnately veined, at least 8 x 8 mm, sessile; base rounded to cordate. Leaflets sessile, the
length 34 times the width, to 4 cm wide; margins parallel. Ovary 2locular...............................
forma divaricata
Synonym: Otophora macrocarpa Ridl., Kew Bull. (1933) 190, Radlkofer in Engler
l.c. (1934) 1494, Masamune l.c. 430.
Known from a few collections from Kuching and Samarahan areas.
Leaves paripinnate, each with up to 6 pairs of leaflets. Pseudostipules elliptic, mostly
pinnately veined, up to 8 x 4 mm, subsessile; base cuneate to subcordate. Leaflets shortstalked, the length 2.53 times the width, to 7 cm wide; margins not parallel. Ovary 3locular......................
forma lunduensis (Radlk.) Leenh.
l.c. (1969) 72, l.c. (1994) 639. Basionym: Otophora lunduensis Radlk. l.c. (1922) 339,
in Engler l.c. (1932) 769, Masamune l.c. 430. Type: Foxworthy 17, Borneo Sarawak
(holotype M).
Known from 3 collections from Lundu area, 1st Div., Sarawak.
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Notes. A few specimens, including Native collector 375 (Type of Otophora pyramidalis
Radlk.), collected from Kuching and its surrounding area are intermediate between forma
divaricata and forma lunduensis.
4. Lepisanthes falcata (Radlk.) Leenh.
Fig. 12H.
(Latin, falcatus = sickle-shaped; the leaflets)
l.c. (1969) 69, l.c. (1994) 633; Whitmore, Tantra & Sutisna l.c. 312. Basionym: Hebecoccus falcatus Radlk., Philipp. J. Sc. 8
(1914) Bot. 453, in Engler l.c. (1932) 721. Lectotype (Leenhouts, 1969): Rosenbluth FB 12631, Philippines, Leyte (M;
isolectotype L).
subsp. borneensis (Leenh.) Leenh.

Blumea 18 (1970) 429, l.c. (1994) 634. Basionym: Lepisanthes borneensis Leenh. l.c. (1969) 68, Anderson l.c. 311. Type:
Chew, Corner & Stainton RSNB 2936, Borneo, Sabah (holotype L; isotypes BO, SAR).
Tree to 22 m tall, 40 cm diameter. Twigs 11.5 cm diameter, when young sparsely yellowish
brown tomentose to glabrous. Leaves each with 410 leaflets; petioles swollen at base, 725
cm long, light yellowish green. Leaflets subopposite to alternate, thin- to stiff-chartaceous,
shiny bright green above, dull green with scattered, glandular-pitted warts beneath; elliptic to
ovate or oblong to lanceolate, 1428 x410 cm, not or slightly sickle-shaped; base symmetric
to asymmetric, acute to rounded and attenuate or subcordate, apex acute to attenuateacuminate; lateral veins c. 15 per side, 1.53 cm apart, spreading, slightly to strongly curved,
some intermediate lateral veins sometimes as strongly developed as the lateral veins, venation
mostly open, only the lateral veins in the upper one third to one fourth distinctly looping and
joining at several mm from the margin; petiolules swollen, 415 mm long. Inflorescences
2070 cm long, minutely brown-tomentose, branches up to 40 cm long, patent to obliqueerect; cymes condensed, several-flowered, stalks c. 5 mm long; pedicels c. 1 mm long; bracts
narrowly triangular, c. 3 mm long. Flowers: sepals ovate to obovate to suborbicular, 34.5x
24.5 mm; petals 47 mm long, blade elliptic to suborbicular, 2.35 mm long, base glabrous
or sericeous, basal one fifth densely long-ciliate, glabrous inside, claw 0.752.5 mm long,
sparsely hairy inside, glabrous or sericeous outside, densely long-ciliate, scales short and
broad, entire, bilobed, or 3-dentate, densely bearded; stamen filaments 23 mm long, anthers
ovoid, 11.5 mm long. Fruits slightly 3-lobed, often only 1 lobe developed, with the sterile
ones not visible, 1.82.5 x 1.83.5 cm, densely brown short-velvety and corrugated outside.
Seeds obovoid-globose, 15 x 810 mm, hilum basal, circular, small.
Vernacular name. Sarawaksegera (Iban).
Distribution. Borneo and the Philippines (Palawan). Probably not uncommon in Sabah,
uncommon in Sarawak; also in Brunei and Kalimantan.
Ecology. Primary lowland to lower montane forest, on rocky banks of streams, at 601500 m.
Flowering in March, June, September, November; fruiting in March, May.
Taxonomy. Two other subspecies are found in Sulawesi (subsp. celebica (Radlk.) Leenh.)
and the Philippines (subsp. falcata). They differ mainly in the venation of the leaflets: closed
317
in subsp. celebica and subsp. falcata; mostly open or closed in the upper one-third to onefourth only in subsp. borneensis.
5. Lepisanthes fruticosa (Roxb.) Leenh.
Fig. 12E.
(Latin, fruticosus = shrubby)
l.c. (1969) 76, l.c. (1994) 643; Anderson l.c. 312; Yap l.c. 446; Whitmore, Tantra & Sutisna l.c. 312. Basionym: Sapindus
fruticosa Roxb., Fl. Ind. ed. 2, 2 (1832) 283. Type: Roxburgh, s.n., Hort. Bot. Calcutta (introduced from the Moluccas)
(CAL, n.v.). Synonyms: Otophora fruticosa (Roxb.) Blume l.c. (1847) 142, Merrill l.c. (1921) 358, Radlkofer in Engler l.c.
(1932) 759, Masamune l.c. 430, Backer & Bakhuizen f. l.c. 135; Otophora pulchella (Ridley) Merr. l.c. (1921) 359,
Masamune l.c. 431; Otophora bijuga Radlk. in Engler l.c. (1932) 765, Masamune l.c. 429; Otophora glandulosa Radlk. in
Engler l.c. (1932) 763, Masamune l.c. 430; Otophora latifolia Ridl., Kew Bull. (1933) 190, nom. illeg., Radlkofer in Engler
l.c. (1934) 1494, Masamune l.c. 430.
Shrub or tree 1.510(15) m tall, 215 cm diameter. Twigs 2.520 mm diameter, smooth or
lenticellate, glabrous or sometimes variably fulvous hairy, glabrescent. Leaves without or
sometimes with a rather strongly reduced terminal leaflet, each with 18(14) pairs of
leaflets, mostly glabrous; petioles 0.532 cm long; rachis terete to laterally flattened, often
marginate to exceptionally narrowly winged in the upper part; pseudostipules very rarely
absent, mostly persistent, ovate, obovate, elliptic, or orbicular, sometimes very oblique, 0.2
10 x 0.110 cm, base cordate to obtuse, apex obtuse to rounded or exceptionally furcate,
reticulately or palmately, rarely pinnately veined. Leaflets opposite to alternate, thinchartaceous to coriaceous, often greyish above, brown beneath, with scattered sunken glands
above, sparsely to densely finely pitted beneath, or exceptionally with minute pitted warts on
both surfaces, glabrous or sometimes hairy on the midrib beneath; ovate- to obovate-oblong
to narrowly lanceolate, 940 x212 cm; base asymmetric or symmetric, subcordate to acute,
mostly attenuate, in asymmetric leaflets sometimes rounded at the lower half and cuneate at
the upper half, apex obtuse to acuminate, rarely acute, acumen short, broad and obtuse to
long, slender and acute, sometimes mucronate; midrib sharply raised to rounded beneath;
lateral veins 0.85 cm apart, straight to curved, none to all looping and joining; petiolules
absent to 3 cm long. Inflorescences terminal, axillary or borne on leafless twigs or stems,
solitary or sometimes clustered, unbranched or if branched either with a few to several
ascending long-branches appearing from near the base, or from all over and pyramidal, to 75
cm long, glabrous; cymes subsessile, rarely with a stalk up to 1 cm long, few- to severalflowered, in the upper part flowers often solitary, sometimes sticky (race glandulosa);
pedicels filiform, 0.31(1.5) cm long. Flowers scentless; sepals 45, glabrous or very
sparsely glandular-ciliolate, outer 2 sometimes smaller, elliptic, orbicular, or obovate, 24x
1.53 mm, dark red, rarely yellow to white, margins, especially of the inner ones, petaloid,
crenulate to fimbriate-ciliolate; petals 45, blade broadly ovate or elliptic to obovate, 1.53x
12 mm, dark red, rarely yellow to white, glabrous, claw short, ciliate or rarely hairy outside,
either with 2 auricles or with a small auricle and a reflexed scale, scales and auricles ciliate;
stamens 58, anthers 1.22.2 mm, yellow to white, glabrous or hairy; ovary 23(4)-locular,
glabrous, stigma subsessile, slightly lobed. Fruits ovoid, ellipsoid, subglobose, or
transversely ellipsoid, rarely distinctly lobed, 13 x0.62 x0.52 cm (fresh fruit to 4 cm
diameter), white when young, dark red to blackish when ripe; fruit wall thin, fleshy when
fresh; septum usually interrupted, mostly reduced to a rib all around; pedicels patent, slender,
to 1.5 cm long. Seeds mostly 2, subglobose to subellipsoid, flattened on the axial side, 823x
618 mm, 418 mm thick, hilum orbicular to lanceolate, to 6 x 34 mm.
318
Vernacular names. Sabahbelingasan (Kedayan), indulus (Dusun Sungei), kelinga (Dusun

Banggi), lulupitan (Murut), tamud tamud (Dusun West Coast).
Distribution. Lower Myanmar, Thailand, Indo-China, Sumatra, Peninsular Malaysia,
Borneo, Java, Philippines, Sulawesi, Lesser Sunda Is., and Maluku. Common in Sabah and
Ecology. In primary and secondary mixed dipterocarp forests, on ridges, along rivers, in
swamps, behind beaches, and in abandoned plantations; on various types of well-drained or
periodically inundated, nutrient-rich or nutrient-poor soil, including clay, sands, acid and
basic soil; from sea level to 1400 m. Flowering and fruiting mainly in December to May.
Fruits sweet, eaten and dispersed by birds, wild pigs and deer.
Uses. In Peninsular Malaysia, the roots are used in traditional medicine. The wood is hard,
durable, and heavy and has been used for house-building in Melaka. Sometimes cultivated for
its edible fruits (Burkill, EPMP (1935) 1614; Jansen et al. PROSEA 2 (1991) 343).
Taxonomy. In Borneo there are 3 main races, acuminata, fruticosa, and glandulosa,
the last one with 2 ecotypes. Race acuminata is characterized by its acute leaflets,
inflorescences borne on leafless twigs or stems, 5-merous flowers, and completely 3-locular
fruits; it is restricted to the Kapuas basin in Kalimantan. Race fruticosa has obtuse leaflets,
axillary or sometimes terminal inflorescences, 4-merous flowers, and incompletely 2-locular
fruits; it is apparently restricted to N Borneo. Race glandulosa is characterized by its
obtuse leaflets, axillary inflorescences (or borne on leafless twigs or stems), 5-merous
flowers, densely short-hairy anthers, and incompletely 2-locular fruits; its lowland ecotype is
rather wide-spread, but its mountain ecotype is restricted to Mt Kinabalu. The mountain
ecotype differs from the lowland ecotype in its more slender petiole and rachis, smaller
pseudostipules that are earlier caducous, narrower leaflets with a longer attenuate apex, and
the midrib which is more rounded beneath.
6. Lepisanthes kinabaluensis Leenh.
Fig. 14.
(of Mt. Kinabalu)
l.c. (1969) 73, l.c. (1994) 639; Whitmore, Tantra & Sutisna 312. Type: Chew & Corner RSNB 4998, Borneo, Sabah, Mt.
Kinabalu, Mesilau river (holotype K; isotype L).
Tree to 10 m tall, or shrub. Twigs 0.81 cm diameter, densely fulvous short-tomentose,

glabrescent. Leaves imparipinnate, each with 714 pairs of leaflets; axial parts hairy; petioles
513 cm long; pseudostipules orbicular to broadly ovate, 25 x2.54 cm, base deeply
cordate, apex rounded or obtuse to short-acuminate, pinnately to reticulately veined, distant
from the normal leaflets. Leaflets more-or-less opposite, parchment-like, mostly greyish
green above, greenish beneath, glabrous or the midrib fulvous short-tomentose above, with
scattered, glandular-pitted warts on both surfaces; lanceolate to oblong, parallel-sided, 12
20 x3.55 cm; base asymmetric to symmetric, cuneate or lower half rounded, apex attenuateacuminate, acumen long, pointed or obtuse; lateral veins 12 cm apart, straight to slightly
curved, more or less distinctly looping and joining near or at some distance from the margin.
Inflorescences terminal, pyramidal, c. 25 cm long, red, densely short-hairy to subglabrous;
branches patent; cymes short-stalked to sessile, several-flowered; pedicels to 5 mm long.
Flowers: sepals deep
319
Fig. 14. Lepisanthes kinabaluensis. A, fruiting leafy twig; B, fruit; C, female flower; D, ovary; E, petal from inside; F,
stamen; G, seed in longitudinal section. (AB from RSNB 4998, CG from RSNB
320
rose-red, sparsely ciliolate in the apical part, outer 2 ovate, 3.55 x2.53 mm, inner 3 elliptic
to rounded, 45 x2.55 mm, rather thin, the greater part petaloid, sometimes grading to
petals; petals 4, pink outside, darker towards the base, whitish inside, claw c. 0.5 mm,
ciliolate, blade widely rhomboid, c. 2.5 x3.5 mm, ciliolate at base, sparsely ciliolate at the
upper part, elsewhere glabrous or appressed hairy outside, scale bilobed or the lateral parts
reflexed, the central part erect and hood-shaped, c. 0.4 as long as the blade; disc glabrous or
short-hairy; stamens 710, anthers c. 2 mm long; ovary 3-locular, 3-angular, obovoid, 3lobed, glabrous, stigma sessile, dome-shaped. Fruits 3-lobed, often 1 or 2 lobes not
developed and style nearly basal, lobes spreading, ellipsoid, c. 3.2 x 2.2 cm, scurfy, glabrous,
brown, yellow, orange, or purplish red when ripe.
Distribution. Uncommon and endemic to Sabah (Mt. Kinabalu, Crocker Range).
Ecology. In montane forest, at 12002000 m. Flowering in March to April; fruiting in
January, March to April, June.
7. Lepisanthes multijuga (Hook. f.) Leenh.
Fig. 12I.
(Latin, multus = many, jugum = leaflet pair)
l.c. (1969) 73, l.c. (1994) 642; Whitmore, Tantra & Sutisna l.c. 312. Basionym: Nephelium multijuga Hook. f., Trans. Linn.
Soc. 23 (1860) 164. Type: Motley 328, Borneo, Labuan (K). Synonyms: Otophora multijuga (Hook. f.) Merr. l.c. (1921)
358, Radlkofer in Engler l.c. (1932) 774, Masamune l.c. 430; Otophora tricocca Radlk. in Merrill, PEB (1929) 174, nom.
nud., Masamune l.c. 431.
Tree or shrub to 12 m tall, 10 cm diameter. Twigs 1.52 cm diameter, glabrous. Leaves

imparipinnate, each with (15)3040 pairs of leaflets; axial parts fulvous tomentose,
glabrescent; petioles 36(8) cm long; pseudostipules 1 or 2 pairs, ovate to suborbicular,
2.57.5x2.55 cm, base cordate, apex acute, pinnately to reticulately veined, more or less
intergrading with the normal leaflets by intermediates. Leaflets opposite to alternate,
subsessile, chartaceous, blackish brown to greenish grey above, medium- to dark-brown
beneath; densely covered with glandular-pitted warts above, sparsely so beneath; linearlanceolate, to 18 x 3 cm; base rounded to subcordate, asymmetric, apex attenuate-acuminate,
obtuse to acute; midrib densely fulvous tomentose above, very sparsely hairy to glabrous
beneath, lateral veins 11.5 cm apart, slightly curved, obscurely looping and joining at a
distance from the margin. Inflorescences terminal, broadly thyrsoid, 2530 cm long,
subglabrous, sparsely branched; branches obliquely patent, to 45(60) cm long, narrowly
thyrsoid with scattered, short-stalked to sessile, condensed, few-flowered cymes; pedicels
0.54 mm long. Flowers red; sepals sparsely appressed short-hairy to glabrous outside,
sparsely glandular ciliolate, outer 2 elliptic to ovate, 2.53x1.82.5 mm, inner 3 orbicular,
with a broad petaloid margin, 34x2.5-4 mm; petals rather densely long-ciliate, sparesely
hairy at the base or glabrous outside, blade broadly ovate, slightly subtruncate at base, c. 2x
2.5 mm, claw c. 0.5 mm, scale slightly bilobed, about 0.3x as high as the blade, slightly
recurved, ciliate; disc glabrous; stamens (6)8, anthers 1.51.8 mm long, long-ciliate to
completely woolly; ovary 3-locular, deeply grooved, glabrous, style conical. Fruits deeply 3lobed (if only 1 lobe is developed, style located near the base above the scars of fallen sterile
lobes), lobes spreading, oblique-ellipsoid, c. 22x13 mm, scabrous, glabrous, chocolatecoloured or yellow or orange when fresh. Seeds hazelnut-shaped, c. 12 x 9 mm.
321
Distribution. Endemic to Sabah (also on Labuan Is.), Sarawak (one collection, from the
Kelabit Highlands), and Brunei.
Ecology. In primary and secondary forests, up to 1000 m. Flowering in July, September,
November; fruiting in January, September, November, December.
8. Lepisanthes ramiflora (Radlk.) Leenh.
(Latin, ramus = twig, flos = flower; flowering on leafless twigs)

l.c. (1969) 81, l.c. (1994) 647; Anderson l.c. 312. Basionym: Otophora ramiflora Radlk., Sapind. Holl.-Ind. (1879) 32, 85,
in Engler l.c. (1932) 758, Merrill l.c. (1921) 359, Masamune l.c. 431. Lectotype (Leenhouts, 1969): Beccari PB 364,
Borneo, Sarawak (FI).
Small tree or shrub to 22 m tall, 20 cm diameter. Twigs 57 mm diameter, densely pustular

lenticellate, sparsely hirsute, early glabrescent. Leaves pari- or rarely imparipinnate, each
with (3)46 pairs of leaflets, 4050 cm long; petiole and rachis sparsely hairy to
subglabrous, wings to 5 mm wide; petioles 618 cm long; pseudostipules ovate, 0.83.5 x0.5
2 cm, base cordate, apex acute to rounded, pinnately veined. Leaflets subopposite, sessile,
chartaceous, sometimes slightly bullate, grey or brown above, brown to greenish beneath;
oblong or sometimes slightly obovate, to 33 x10.5 cm; base asymmetric, narrowed, cordate,
apex attenuate-acuminate, acute; midrib rounded beneath, sparsely hirsute at least near the
base, lateral veins 12.5 cm apart, more or less curved, at least those in the upper half
distinctly looping and joining at some distance from the margin. Inflorescences several to
many fascicles borne on knobs of leafless twigs and stems, unbranched or hardly branched,
racemose, c. 1.5 cm long, few-flowered; pedicels patent, slender, 1.55 mm long. Flowers 4merous; sepals pale pink, outer 2 elliptic, c. 2.5 x2 mm, with petaloid margin, inner ones
broadly ovate, c. 3 x2 mm, for the greater part petaloid; petals white, sparsely long-ciliate
near the base, blade semi-transversely elliptic, c. 1 x1.8 mm, claw c. 0.4 mm long, scale
broad, rounded to bilobed, about 0.30.5x as high as the blade, slightly hairy; disc pale
yellow, glabrous; stamens 57, anthers c. 1 mm long, glabrous, cream-coloured; ovary
suborbicular, flattened, 2-locular, pale yellowish, style short, stigma dome-shaped, slightly
bilobed, white. Fruits transversely ellipsoid, slightly bilobed, c. 1 x1.5 cm, crimson to dark
purple, incompletely 2-locular. Seeds ellipsoid.
Distribution. Endemic to Sarawak (around Kuching).
Ecology. In primary and secondary forests on sandstone, from sea level to 100 m. Flowering
in January to February, July to September, November; fruiting in January, October,
November.
Uses. The fruits are edible.
9. Lepisanthes rubiginosa (Roxb.) Leenh.
Fig. 12F.
(Latin, rubiginosus = rust-coloured; the indumentum)
l.c. (1969) 82, l.c. (1994) 648 ; Yap l.c. 446; Whitmore, Tantra & Sutisna l.c. 313. Basionym: Sapindus
rubiginosa Roxb., Pl. Corom. 1 (1796) 44, t. 62. Type: Roxb. Pl. Corom. 1 (1796) t. 62. Synonyms:
Erioglossum rubiginosum (Roxb.) Blume l.c. (1847) 118, Merrill l.c. (1921) 358, Radlkofer in Engler l.c.
322
(1932) 693, Masamune l.c. 426, Backer & Bakhuizen f. l.c. 134, Corner, WSTM 3rd ed. 2 (1988) 679, pl. 209.
Shrub or small tree to 16 m tall, 28 cm diameter. Indumentum ferrugineous to fulvous,

sometimes silvery grey. Twigs c. 5(15) mm diameter, densely short-hairy when young.
Leaves each with (2)36(9) pairs of leaflets, often with a pseudoterminal leaflet, velvety
when young; petioles, 7.512(20) cm long, densely short-hairy, late glabrescent. Leaflets
stiff-chartaceous, greyish green to grey above, yellowish green to reddish brown beneath,
densely short-hairy on midrib and lateral veins on both sides, sparsely hairy on the blade
beneath (velvety to the touch), glabrescent; elliptic to lanceolate, (4.5)6.518(25)x (2-)
3.58.5(11) cm; base rounded to broadly cuneate, apex obtuse to acute or acuminate, often
mucronulate; midrib prominent above, rather strongly raised and rounded beneath, lateral
veins 812(16) on either side, nearly straight to slightly curved, bent at the margin, except
for few upper ones, not distinctly joining, hardly prominent above, slightly prominent
beneath, intercostal veins and veinlets inconspicuous above, densely reticulate-scalariform
and faintly raised beneath; petiolules to 5(10) mm long. Inflorescences 2535(50) cm
long, densely ferrugineous tomentose, branches often long, ascending, spicate; cymes nearly
sessile to short-stalked, glomerulous to distinctly branched, few- to several-flowered; bracts
and bracteoles small, subulate; pedicels 12(5) mm long. Flowers sweet-scented; sepals
orbicular-ovate, slightly concave, margin sometimes petaloid, ciliate, glabrous or with a few
hairs inside, outer two 1.22.2 x 1.22 mm, ciliate, inner three 1.82.8 x23 mm, obtuse;
petals 4 (or 5), blade 24 x1.52.2 mm, crenulate in the upper half, white to yellowish,
sparsely long-ciliate in the narrowed lower part, sparsely hairy at base outside, abaxial petals
shorter and narrower than the adaxial ones, and with a shorter scale, claw 0.51 mm long,
long-ciliate in the upper part, scale 1.53 mm long, quadrangular to more or less bilobed,
slightly narrowed from base to apex, bearded, the appendage deeply bilobed, the lobes
sometimes also bilobed, hairy; abaxial stamens longer than the adaxial ones, stamen filaments
flattened, white long-hairy, in male flowers (1.5)35 mm long, in female flowers c. 1.5 mm
long, anthers oblong-ovoid, c. 0.8 mm long, glabrous; ovary 3-lobed, 1.21.8 x22.2 mm,
densely appressed hairy, style cylindric, c. 2.2 mm long, bent near the obscurely 3-lobed
apex, sparsely appressed hairy in the lower two third; pistillodes in male flowers c. 0.8 mm
high, long-hairy. Fruits 13-lobed, lobes spreading, 813 x78 mm, faintly keeled, dark
purple to nearly black when ripe, subglabrous; endocarp thin but hard and tough, glabrous.
Seeds oblong-ellipsoid, 911 x 4 x 4 mm, hilum basal, small.
Vernacular names. Sabahborobogan (Dusun Bundu), damai (Bajau), lipupudsu (Dusun),
suang rason (Bajau Laut).
Distribution. Continental SE Asia from northern India to Indo-China and SE China, Malesia
and NW Australia. In Sabah, rather common; also in Brunei and Kalimantan.
Ecology. In young secondary forest, shrub growth, forest edges, road sides, riverbanks, and
inner mangroves; up to 1200 m; on well-drained or periodically inundated, nutrient-rich or
nutrient-deficient soils, including heavy clay, sand, and limestone. Flowering and fruiting
throughout the year. Flowers much visited by Xylocopa bees.
Uses. In India, where the trees seem to be bigger, the timber is said to be valuable, but in
Malesia it is used only for firewood and sometimes (in Java) for rice-pounders and tool-
323
handles. A decoction of the roots and leaves, sometimes also of fruits and seeds, is used
medicinally against fever. The young leaves are eaten as a vegetable, and the astringent but
sweet fruits are relished as a titbit, mainly by children (Burkill, EPMP (1935) 938; Heyne,
Nutt. Pl. Ned. Indi, ed. 2 (1927) 989; Ochse & Bakhuizen, Ind. Groenten (1931) 648, f.
396).
10. Lepisanthes senegalensis (Juss. ex Poir.) Leenh.
(of Senegal)
Fig. 12D
l.c. (1969) 85, l.c. (1994) 651; Anderson l.c. 312; Whitmore, Tantra & Sutisna l.c. 313. Basionym: Sapindus senegalensis
Juss. ex Poir., Encycl. 6 (1805) 666. Type: Adanson & Geoffroi f., s.n., Herb. Jussieu 11386, Senegal (P). Synonym:
Aphania dasypetala Radlk., Fedd. Repert. 18 (1922) 333, in Engler l.c. (1932) 714, Masamune l.c. 425.
Treelet, shrub, or tree to 24 m tall, 44 cm diameter. Young parts densely to sparsely

appressed fulvous to ferrugineous short-hairy. Twigs 26 mm diameter, smooth to
verruculose by many small, orbicular lenticels. Leaves sometimes with a pseudoterminal
leaflet, with 14(6) pairs of leaflets or sometimes simple; lower pair of leaflets sometimes
attached near or at the base, smaller and more caducous than the other ones; petioles absent to
14 cm long, often strongly swollen at base; rachis terete to keeled above. Leaflets chartaceous
to subcoriaceous, greyish green (rarely olive or brown) above, greyish or yellowish green to
light brown beneath, densely to sparsely set with minute pellucid dots; elliptic to lanceolate,
735 x215 cm (simple leaves to 60 x18 cm); base acute to obtuse, more or less attenuate (in
sessile simple leaves subcordate, symmetric or asymmetric), apex obtuse to acuminate,
mucronulate, acumen to 5 cm long, blunt to acute; midrib faint to slightly raised above,
prominent and rounded beneath, lateral veins 720 per side, 13(4 in the large simple
leaves) cm apart, straight to curved, looping and joining at some distance from the margin,
slightly raised on both surfaces, shorter intermediate veins strongly developed between two
lateral veins, intercostal veins rather densely reticulate, slightly raised on both surfaces,
especially beneath; petiolules swollen, 215 cm long. Inflorescences hairy to subglabrous, in
fascicles of a few to several, unbranched or sparsely slender-branched racemes to 40 cm long,
or unbranched or widely sparsely branched thyrses to 60 cm long with a strong peduncle and
rachis; cyme-like parts distant, short-stalked, 3-flowered; pedicels 14(8) mm long. Flowers
slightly fragrant, female ones apparently developing first; sepals 5, rarely 4, mostly dark red,
concave, outer 2 oblong-ovate to orbicular, apex rounded, 13 x0.81.2 mm, sometimes with
a narrow, petaloid, crenulate margin, sparsely ciliolate, inner ones 34 x3 mm, mostly with a
broad, petaloid, crenulate to fimbriate, ciliolate margin; petals 5, rarely 4, creamish or
greenish white, imbricate, short-clawed to sessile, blade elliptic to oblong, widest below to
about the middle, 2.54.5x12 mm, entire, rounded, membranous but often thickened
towards the centre and the base, ciliate at least near the base, other parts mostly glabrous,
sometimes subsericeous outside, scale minute to two-fifths the length of the blade, simple to
deeply bilobed, rarely divided into two minute auricles, sparsely ciliate to long-hairy
(exceptionally glabrous); disc entire (more or less interrupted if one of the petals is
suppressed), annular to saucer-shaped; stamens 57(9), filaments flattened or terete,
narrowed from base to apex, 13.5 mm long, white, woolly in the lower two third, rarely
subglabrous, anthers ellipsoid to ovoid, 11.8 mm long, light yellow when young, purplish
brown when old, connective mostly apiculate; ovary 2(3)-lobed, 1.53 mm across,
subsessile or with a stalk to 1.5 mm long, mostly with some stiff hairs at the style base, style
cylindric, 12 mm long, straight to slightly curved, stigma sometimes decurrent halfway
down the style, purple; pistillode often long-pilose in the apical
324
part. Fruits 2-lobed (often only 1 lobe developed), sessile or with a stalk to 2 mm long, lobes
short-ellipsoid to globose, 815 x515 mm, smooth, dark red to black when ripe; endocarp
eggshell-like, light brown. Seeds ovoid to globose, more-or-less attenuate towards the hilum,
78 x 67 mm; testa shiny brown to black, hilum circular.
Vernacular names. Sabahkelinga gaba (Dusun Banggi). Sarawakmelingkat (Iban).
Distribution. Tropical Africa, Madagascar, Sri Lanka and the SW Deccan Peninsula, NE
India, E Pakistan, Myanmar to Indo-China and Hainan, the Andamans and Nicobars, and
Malesia (all regions except the Lesser Sunda Islands). Common in Sabah, less so in Sarawak;
also in Brunei.
Ecology. In mixed dipterocarp forest and secondary forest, from sea level to 1400 m; on
well- drained as well as marshy, fertile to infertile, acid to basic soils, including clay, sand,
marl and coral limestone rocks. Flowering mainly in July to November; fruiting mainly in
January to April.
Uses. The wood is hard, heavy, and durable, and is used, in Peninsular Malaysia, for houseposts. The fruits are edible (Burkill, EPMP (1935) 190).
Notes. Two races (montana and dasypetala) occur in Borneo. The latter differs from the
former in its flowers with 4 petals that are rather densely sericeous outside, ovary with a
longer stalk, leaflets with an inconspicuous venation and lateral veins that are more distinctly
looping and joining at a slightly greater distance from the margin.
11. Lepisanthes tetraphylla (Vahl) Radlk.
(Greek, tetra = four, phullon = leaf; leaf of four leaflets)
Fig. 12J.
Sitzungsber. Math. Phys. Cl. Knigl. Bayer. Akad. Wiss. Mnch. 8 (1878) 276, in Engler l.c. (1932) 743, f. 15; Ridley FMP
5 (1925) 301; Leenhouts l.c. (1969) 39, 63, l.c. (1994) 630; Anderson l.c. 312; Yap l.c. 447; Whitmore, Tantra & Sutisna l.c.
313. Basionym: Sapindus tetraphylla Vahl, Symb. 3 (1794) 54. Type: Koenig, s.n., Herb. Vahl, India (C). Synonyms: L.
heterolepis Blume l.c. (1847) 153, Merrill l.c. (1921) 358, Radlkofer in Engler l.c. (1932) 734, Masamune l.c. 427; L.
petiolaris Radlk., Fedd. Repert. 18 (1922) 334, in Engler l.c. (1932) 733, Masamune l.c. 427.
Mostly shrub or treelet, sometimes tree to 23 m tall, 20 cm diameter. Twigs to 2.5 cm

diameter, glabrous or variably fulvous to ferrugineous hairy, mostly early glabrescent, mostly
conspicuously lenticellate. Leaves with 210 pairs of leaflets; axial parts glabrous or variably
hairy; petioles 550 cm long. Leaflets chartaceous to parchment-like, glabrous or variably
hairy; lanceolate to elliptic, ovate to obovate, 755 x210(20) cm; base slightly (or strongly)
asymmetric to symmetric, acute to rounded or subcordate, apex tapering to abruptly
acuminate, acumen short to long, obtuse to acute, lateral veins variable, at least the upper
ones (rarely all) looping and joining or exceptionally connected by an intramarginal vein,
slightly raised or sometimes sunken above, slightly raised to prominent beneath, intercostal
veins and veinlets finely reticulate; petiolules 225 mm long. Inflorescences 270 cm long,
mostly densely short-hairy; peduncles very short to long; main branches often racemose;
cymes patent, sessile or with a stalk to 0.5 cm long, to 7-flowered; pedicels to 6 mm long;
bracts ovate-lanceolate to subulate, to 5 mm long, rarely ovate and to 7 x 6 mm. Flowers
325
white, sometimes greenish white, cream, or pink, sweet-scented; outer two sepals ovate to
suborbicular, 1.27 x1.26 mm, densely (sometimes partly glandular) ciliolate, glabrous to
sparsely hairy inside, inner 3 sepals elliptic to orbicular, ovate to obovate, 2.26.2 x1.86
mm, margins mostly narrowly to broadly petaloid, glabrous outside, crenulate to fimbriatedenticulate, indumentum otherwise like that of outer sepals; petals 2.210 mm long, claw
0.52(4) mm long, sparsely hairy inside, blade elliptic to oblong to ovate, 1.24 mm wide,
more-or-less dentate in the upper part, mostly up to two-thirds sericeous outside, rarely
subglabrous, margin glabrous or sparsely to densely woolly ciliate below the insertion of
scale, rarely rest of the blade (partly glandular) ciliate outside, glabrous inside or the base
rarely sparsely hairy, scale mostly well-developed (rarely represented by a hairy rim only),
entire to deeply 24-lobed, glabrous, ciliate or sparsely to densely woolly inside, without or
more often with a crest varying in shape from a small wart to a deeply bilobed duplication of
the scale or 2 brushes; stamen filaments 1.56.5 mm long, sparsely to densely woolly in the
upper half, or rarely also in the lower half, anthers broadly ellipsoid to oblong, ovoid to
obovoid, 0.82.5 mm long, connective broad and obtuse or rarely narrow and pointed, woolly
to glabrous; ovary and lower part of style densely hairy to subglabrous. Fruits turning to
yellowish, grey, or greyish pink on ripening, flattened ellipsoid, shortly obovoid, or
subglobose, slightly lobed, the lobes rarely keeled, 1.55 cm diameter. Seeds brown; testa
papery, probably partly fleshy when fresh.
Vernacular names. Sabaharit-arit (Dusun), bansisi (Dusun Labuk). Sarawakjung
(Iban).
Distribution. SE Asia, from Sri Lanka and the Deccan Peninsula to Hainan, Sumatra,
Peninsular Malaysia, Java, Timor, Borneo, Philippines, N Sulawesi (one collection), and New
Guinea. Common in Sabah and Sarawak; also in Brunei and Kalimantan.
Ecology. In primary mixed dipterocarp and secondary forests, in forest edges, on riverbanks,
and in open country; on well-drained or swampy or periodically inundated localities, on all
kinds of soil; to 1200 m. Flowering throughout the year; fruiting in November to July.
Uses. The wood of some forms is heavy, hard, and close-grained, and good for furniture
(Burkill, EPMP (1935) 1332, under L. kunstleri).
12. LITCHI Sonn.
(after the Chinese plant-name li-tchi)
P.W. Leenhouts
Voy. Ind. Or. Chine 2 (1782) 230, t. 129; Radlkofer in Engler, Pflanzenr. 98 (1932) 914; Backer & Bakhuizen f., FJ 2 (1965)
137; Leenhouts, Blumea 24 (1978) 398, FM 1, 11 (1994) 653; Yap, TFM 4 (1989) 447.
Monoecious trees. Indumentum of solitary, simple or 2-branched hairs. Leaves spirally

arranged or sometimes partly, especially towards the inflorescence, nearly opposite,
paripinnate, each with 1-4(5) pairs of leaflets, without pseudostipules; petiole and rachis
not winged. Leaflets mostly subopposite, dull beneath, margin entire to undulate in the upper
part.
326
Inflorescences terminal and axillary, thyrsoid; bracts triangular. Flowers radially

symmetrical, unisexual; calyx 45-lobed, densely appressed short-hairy on both sides, tube
cupular, lobes to 0.30.5 the length of the tube, apert in bud, equal, not petaloid; petals
absent; disc annular, small, without appendages; stamens 611, far exserted in male flowers,
short in female and in seemingly bisexual but functionally male flowers, filaments variably
hairy, anthers glabrous; ovary short-stalked, 2(3?)-locular, style terminal, shorter than the
ovary, stigma consisting of 2(3?) long, spreading to recoiled lobes; ovules 1 per locule,
basal. Fruits nearly always with only 1 locule developed, hardly stalked, ellipsoid, ovoid, or
globose, not winged, indehiscent, glabrous, spiny to scaly; pericarp thin, coriaceous to rather
hard; endocarp glabrous. Seeds glabrous, partly or completely enveloped by an entire,
translucent, juicy arillode; embryo straight, cotyledons thick, superposed, radicle basal.
Distribution. Monotypic. SE China, Peninsular Malaysia, Borneo, Philippines, SE New
Guinea; cultivated elsewhere.
Litchi chinensis Sonn.
Fig. 15.
(of China)
l.c. 230, t. 129; Radlkofer in Engler l.c. (1932) 917, f. 21; Backer & Bakhuizen f. l.c.137; Poilane, J. Agr. Trop. Bot. Appl.
12 (1967) 541; Leenhouts l.c. (1978) 398, l.c. (1994) 654; Yap l.c. 447. Type: Sonnerat 1062, China (G, P).
subsp. chinensis
Tree to c. 35 m high, c. 1 m diameter. Twigs 2.53.5 mm diameter, appressed brown-hairy

when young, early glabrescent. Leaves with 24(5) pairs of leaflets; petioles swollen and
slightly hollowed at base, 1.27 cm long, early glabrescent to glabrous, often pustular
lenticellate; rachis flat or grooved with a fine central rib (lower part) or keeled (upper part)
above, rounded beneath. Leaflets chartaceous to coriaceous, mostly rather stiff, smooth,
shiny above, glabrous to fairly densely appressed minute-hairy beneath; elliptic or obovate,
(3)811(16) x1.84 cm; base acute, more or less attenuate, symmetric to sometimes
asymmetric, margin slightly recurved, apex gradually or abruptly tapering acuminate, acumen
obtuse to acute or sometimes rounded to slightly emarginate; midrib narrowly grooved above,
prominent and rounded, lateral veins 0.51 cm apart, inconspicuous on both sides, intercostal
veins coarsely reticulate, veinlets minutely tessellate, distinct; domatia absent; petiolules 38
mm long, narrowly and deeply grooved above, swollen toward the base. Inflorescences
ferrugineous-strigose, 1530 cm long, few-branched; cymules lax, 512-flowered, stalks to 5
mm long; pedicels slender, 23(4) mm long; bracts 0.52 mm long. Flowers greenish white
or yellowish, fragrant; calyx 4(5)-lobed, c. 1.5 x2 mm; disc variably hairy to glabrous;
stamens 6(10), filaments filiform, to 2.5 mm long, rather densely hairy all over, anthers
ellipsoid, apiculate to emarginate at apex, c. 1 mm long; ovary c. 2 x5 mm, with spreading
lobes, densely warty, stalk 11.5 mm long, style terete, c. 1 mm long. Fruits c. 3.5 x3 cm,
bright red to purplish when ripe, nearly smooth, scaly, or densely set with flat, pyramidal,
acute warts of up to 1 mm high. Seeds ellipsoid, c. 2 x1.5 cm; testa shiny blackish brown;
hilum basal, circular, 67 mm diameter; arillode bluish white or light yellow to pinkish, to 5
mm thick when fresh.
Vernacular names. Lychee (popular local name). Sarawakpareh (Iban).
327
Fig. 15. Litchi chinensis. A, fruiting leafy twig; B, female flower; C, immature male flower; D, mature male flower. (A from
FRI 36409, BD from FRI 27888.)
328
Distribution. Probably originating from the northern parts of Indo-China or from SE China;
widely cultivated mainly in subtropical regions. In Malesia, possibly indigenous in Peninsular
Malaysia, Borneo and the Philippines, and rarely cultivated or naturalized and apparently
hardly ever fruiting. Rarely cultivated in Sabah, Sarawak and Brunei; apparently not
indigenous.
Ecology. Prefers a hot and wet climate with at least a short cool dry season, but no frost. It
seems to favour compact, moist, and fertile soils, and is mostly grown at low altitudes on
riverbanks, along dykes, between ditches or ponds, etc. The mountain lychee, which has
smaller and more prickly fruits and may represent the original form, however, is grown on
well-drained land in the hills. The fruits are dispersed by bats and parrots.
Uses. Cultivated for its fruits mainly in SE China, Florida, Hawaii, Madagascar and South
Africa, from where they are exported fresh, dried, or canned. The wood is also highly prized,
reported to be hard, durable, and taking a fine polish (Groff, Ding & Groff, Lingn. Agr. Rev.2
(1924) 34; Burkill, EPMP (1935) 1546; Menzel l.c. 191).
Notes. Although probably one of the best-known genera in the family, to the taxonomist
Litchi remains full of mystery regarding its history, phytogeography, and taxonomy (for more
detailed discussion on its origin and morphological variation, see Leenhouts l.c. 1994).
13. MISCHOCARPUS Blume, nom. cons.

(Greek, mischos = stalk, karpos = fruit; the stalked fruits)
R.W. J. M. van der Ham

Bijdr. Fl. Ned. Ind. (1825) 238, Rumphia 3 (1849) 166; King, J. As. Soc. Beng. 65, 3 (1896) 447; Merrill, EB (1921) 362;
Ridley, FMP 1 (1922) 507; Radlkofer in Engler, Pflanzenr. 98 (1933) 1288; Masamune, EPB (1942) 427; Backer &
Bakhuizen f., FJ 2 (1965) 140; van der Ham, Blumea 23 (1977) 251, FM 1, 11 (1994) 658; Corner, WSTM 3rd ed. 2 (1988)
681; Yap, TFM 4 (1989) 449; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 313.
Shrubs or large trees, monoecious or dioecious. Indumentum of mostly appressed, single

hairs only. Leaves paripinnate, each with 16 pairs of leaflets; upper leaflets larger;
pseudostipules and wings absent; petioles subterete to dorsiventrally flattened. Leaflets
alternate to subopposite; base symmetric to slightly asymmetric, margin entire, flat to
revolute, apex emarginate, rounded to acuminate, often mucronulate; lower surface not
papillose, red glands absent, domatia often present in the axils of lateral veins; lateral veins
looping and joining in the upper part to indistinctly so in the lower half to three-quarters,
intermediate lateral veins present, intercostal veins forming a very regular, dense reticulate
pattern, more or less prominent on both surfaces; petiolules present. Inflorescences thyrses,
pseudoterminal, axillary, or borne on leafless twigs, nearly always branching; cymules 17(
10)-flowered; pedicels 13(5) mm long; bracts triangular to lanceolate. Flowers unisexual;
calyx spreading or cup-shaped, 5(6)-lobed, lobes connate for up to two-thirds, persistent in
fruit, lobes subequal, slightly imbricate at base; petals absent or up to 5, minute to slightly
longer than the calyx, apert, clawed or not, blade basally usually with 2 auricles or scales,
without crest; disc
329
uninterrupted or interrupted, annular or cup-shaped; stamens (5)8(9), exserted, filaments

filiform, anthers basifixed, latrorse or latero-introrse, connective with a lighter coloured wart
at the top; ovary (2)3(4)-locular, stalked or subsessile, style apical, the upper part split into
3 more or less recurved stigmatic lobes, ovules one per locule. Fruit a loculicidal capsule,
nearly always stalked, not lobed, not winged, red, smooth; the locules about equally
developed but the seeds abortive in 1 or 2 locules; stalk hollow; valves thin to almost woody,
usually shrivelled after dehiscence; pericarp slightly fleshy; endocarp glabrous or variably
hairy, sclerenchymatic, complete and lining the valves and distal parts of septa, or incomplete
and present along the sutures only. Seeds hanging by the pseudofunicle; testa shiny, chestnutbrown, nearly completely covered by a thin-fleshy, translucent arillode, with an abaxial
pseudofunicle descending into the stalk; cotyledons superposed.
Distribution. About 15 species; from SE Asia, throughout Malesia to Australia; two species
Ecology. In primary and sometimes secondary forests, from sea level (rocky sea coasts; salt
water creeks) to 3000 m. The fruits may be attractive to animals because of the showy,
slightly fleshy arillode.
Key to Mischocarpus species
Leaflets (1)26 per side; intercostal veins usually coarsely reticulate. Calyx lobes hardly
connate. Petals 5 or reduced in number. Filaments usually puberulous; anthers more or less
hairy..............................................................................................................1. M. pentapetalus
Leaflets 13(4) per side; intercostal veins densely and finely reticulate. Calyx connate for
one-fourth to one-third its length. Petals absent. Filaments and anthers glabrous.......................
...........................................................................................................................2. M. sundaicus
1. Mischocarpus pentapetalus (Roxb.) Radlk.
(Greek, penta = five, petalon = petals; the flower)
Fig. 16F.
Sapind. Holl.-Ind. (1879) 43, in Engler l.c. (1933) 1293; van der Ham l.c. (1977) 271, l.c. (1994) 662; Whitmore, Tantra l.c.
215; Yap l.c. 449; Whitmore, Tantra & Sutisna l.c. 313. Basionym: Schleichera pentapetala Roxb., Fl. Ind. ed. 2 (1832)
275. Type: M.R. Smith (3041?), 1811, India ( holotype BM). Synonyms: M. sumatranus Blume l.c. (1847) 168, King l.c.
448, Merrill l.c. (1921) 362, Ridley l.c. (1922) 508, Masamune l.c. 427, Anderson l.c. 312, Corner l.c. 681; Mischocarpus
fuscescens Blume l.c. (1847) 169, King l.c. 447, Merrill l.c. (1921) 362, Masamune l.c. 427, Backer & Bakhuizen f. l.c. 141.
Shrub or tree to 15(25) m tall; all young parts puberulous, partly glabrescent. Leaves each
with (1)25(6) pairs of leaflets; petioles 325 cm long. Leaflets parchment-like to
coriaceous; ovate to elliptic, (5)720(40) x(1)26(15) cm; base rounded or angular;
domatia usually present in the axils of lateral veins below; midrib prominent above, angular
to rounded in the basal part, or in larger leaflets completely so or rather flat and broad, lateral
veins (6)920(25) per side, at least prominent on the lower surface, hardly distinct from
intercostal veins, intercostal veins usually coarsely reticulate, hardly or not prominent above,
prominent beneath; petiolules 312 mm long. Inflorescences axillary and pseudoterminal;
330
Fig. 16. Mischocarpus sundaicus (AE); M. pentapetalus (F). A, fruiting leafy twig; B, fruit; C, male flower; D, male flower
with two sepals removed; E, bisexual flower; F, fruit. (AB from SAN 50950, CD from SAN 116099, E from SAN 121647,
F from PNH 34128.)
331
main axis to 40 cm long, secondary axes 120 cm long; cymules to 11 mm long; bracts
triangular, 68 x1.54 mm, those of the secondary axes caducous. Flowers: calyx lobes
hardly connate to one-fourth its length, triangular to ovate, 0.752 x12 mm, subcoriaceous
to slightly fleshy, 1(3)-veined, upper part puberulous to pubescent outside, or rarely very
sparsely so in the upper part, puberulous inside; petals 5 or less, usually unequal, ovate to
elliptic, to 2 x1 mm, glabrous or pubescent at the base, not clawed, usually auriculate; disc
annular, (1)1.52(3) mm diameter, sometimes irregularly lobed, rarely with lobes
protruding between the stamens, puberulous or glabrous; stamens 7 or 8, 34 mm long,
filaments usually puberulous, rarely glabrous, anthers papillose, sparsely pubescent or rarely
glabrous; ovary puberulous. Fruits (0.8)12(3) cm long, glabrous; stalk 211 x12 mm;
seed-bearing locules globose to ellipsoid to obovoid, 3-angular or triquetrous in cross section,
0.41.4 cm diameter, glabrous or completely rather densely hairy inside; endocarp
incomplete (2 mm thick at either side of the suture) to complete, usually less so in the fertile
than in the sterile locules. Seeds to 8(12) mm long, ellipsoid to globose; cotyledons
subcollateral, both or only the lower one folded, lower one usually smaller.
Distribution. From India, Myanmar, SW China, throughout SE Asia, to W Malesia.
Relatively frequent in Sabah and Sarawak; also in Kalimantan and possibly in Brunei.
Ecology. In primary and secondary forests, at (300)8002000 m. In Sabah and Sarawak,
mainly found on fertile clay loam soil. Flowering and fruiting throughout the year, but the
main periods differ from region to region.
2. Mischocarpus sundaicus Blume
Fig. 16AE.
(of the Sunda islands)
l.c. (1825) 238; King l.c. 447; Merrill., En. Philip. 2 (1923) 513; Radlkofer in Engler l.c. (1933) 1299; Masamune l.c. 428;
Backer & Bakhuizen f. l.c. 141; van der Ham l.c. (1977) 281, l.c. (1994) 667; Anderson l.c. 312; Whitmore & Tantra l.c.
215; Corner l.c. 681; Yap l.c. 449; Whitmore, Tantra & Sutisna l.c. 312. Lectotype (van der Ham 1977): Anonymous, s.n. (=
L. sheet no. 908.269749), Java, Nusa Kambangan (L).
Shrub or tree to 10(30) m tall; young parts and inflorescences puberulous. Leaves each with
13(4) pairs of leaflets or simple; petioles 111 cm long. Leaflets parchment-like to
subcoriaceous; ovate to elliptic, 417(26) x1.57(10) cm; base rounded or angular, margin
slightly revolute, domatia usually present in the axils of lateral veins below; midrib prominent
above, broad to rather narrow, rounded or often locally to completely angular beneath, lateral
veins 815 per side, moderately curved, prominent beneath; intercostal veins densely and
finely reticulate, slightly raised to flat above, more prominent toward the margin beneath;
petiolules 38 mm long. Inflorescences axillary and pseudoterminal, 1.525 cm long,
branched; cymules short-stalked; pedicels 13 mm long; bracts triangular, to 6 x mm,
usually caducous. Flowers: calyx connate for one-fourth to one-third its length, triangular,
0.652 x0.51 mm, subcoriaceous, puberulous to pubescent in the lower half or rarely in the
upper half, glabrous to sparsely hairy inside; petals absent or to 3 (in Peninsular Malaysia);
disc annular, 12 mm diameter, glabrous or sparsely puberulous; stamens (6)8(9), to 4 mm
long, filaments glabrous, anthers not papillose, glabrous; ovary puberulous. Fruits 717(
20) mm long, glabrous or very sparsely puberulous; stalk 214 x11.5 mm; seed-bearing
lobes globose to ellipsoid, round to faintly triangular in cross section, glabrous to hairy along
332
the sutures inside; endocarp completely sclerenchymatic. Seeds globose to ellipsoid, to 7 mm

long; cotyledons superposed, about equal.
Distribution. India, S China, throughout SE Asia and Malesia. Relatively common in Sabah
and Sarawak; also in Kalimantan and possibly in Brunei.
Ecology. In primary and secondary lowland, hill, and lower montane forests, from sea level
to 1600 m. Flowering mostly in JanuaryAugust; fruiting mostly in MayDecember.
14. NEPHELIUM L.
(Greek, nephelion = a little cloud)

rambutan (Malay)
P.W. Leenhouts
Mant. Pl. (1767) 18; King, J. As. Soc. Beng. 65, 3 (1896) 432; Merrill, EB (1921) 359; Ridley, FMP 1 (1922) 498; Radlkofer
in Engler, Pflanzenr. 98 (1932/33) 950; Masamune, EPB (1942) 428; Brown, FTSB (1955) 317; Backer & Bakhuizen f., FJ 2
(1965) 138; Anderson, CLTS (1980) 312; Leenhouts, Blumea 31 (1986) 373, FM 1, 11 (1994) 669; Whitmore & Tantra,
CLS (1986) 215; Corner, WSTM 3rd ed. 2 (1988) 682; Yap, TFM 4 (1989) 450; Whitmore, Tantra & Sutisna, CLK 2, 1
(1990) 313.
Medium-sized to tall trees or rarely shrubs, dioecious or sometimes monoecious. Indumentum

of solitary simple hairs. Leaves spirally arranged, paripinnate, 1-foliolate or each with 15(
18) pairs of leaflets, without pseudostipules; neither petiole nor rachis winged. Leaflets
alternate to more rarely opposite, mostly distinctly glaucous and finely papillate beneath,
domatia often present; margin entire or rarely repand; venation open (except in N.
subfalcatum). Inflorescences axillary, pseudoterminal, terminal, or borne on leafless twigs
and stems (N. cuspidatum). Flowers unisexual, radially symmetrical; sepals (4)5(6), free
to connate for more than halfway, valvate or sometimes slightly imbricate, all equal, not
petaloid, entire, hairy inside, sparsely short-hairy outside, not ciliate, without glands; petals
5(6) or 14 reduced, or often absent, if present shorter than the calyx, clawed, welldeveloped, with a bilobed scale without appendages, hairy on both sides, entire; disc entire,
often slightly lobed, without appendage; stamens 410, exserted in male flowers, filaments
rather densely long-hairy at least in the basal part, anthers nearly always with at least a few
hairs; ovary short-stalked, (1)2(4)-locular, mostly warty, densely hairy, style usually welldeveloped, stigmas fairly long, spreading to finally recurved; ovules 1 per locule, half
enveloped by an outgrowth of the placenta. Fruits 1(2)-lobed; stalk short to inconspicuous;
lobes ellipsoid to subglobose, warty to spiny, or exceptionally nearly smooth; fruit wall thinto thick-coriaceous or rarely corky to nearly woody, glabrous inside, indehiscent or
apparently often dehiscing irregularly at the upper part or splitting into two or more or less
equal valves. Seeds: hilum nearly basal; micropylar wart apical or mostly subapical;
sarcotesta covering the whole seed, except for the micropylar region or at least perforated in
front of the micropyle, sometimes with a collar-like outgrowth around the hilum; endotesta
tough or exceptionally rather hard; embryo erect with the plumule at the micropylar end,
equal cotyledons, straight suture and fully included plumule and radicle or transverse with
lateral plumule, often very unequal cotyledons (the hilar one larger), curved to sigmoid
suture, and exposed radicle.
333
Distribution. Twenty two species; SE Asia from Yunnan and Assam to Hainan, Peninsular
Malaysia, Sumatra, Jawa, Borneo, Philippines, Sulawesi, and Maluku; 15 species in Sabah
and Sarawak.
Ecology. Midstorey trees of rain forest at low to medium altitudes; outside Borneo sometimes
also in deciduous or savannah forest. The fruits are mainly dispersed by monkeys and fruitbats, though in some species dispersal by water has been reported.
Uses. The sarcotesta of a few species is edible, but only N. lappaceum and N. ramboutan-ake
are commonly cultivated for their commercially important fruits. The timber is hardly of any
commercial importance.
Taxonomy. Identification is hardly possible without more or less mature fruits. Therefore,
under many species notes have been given regarding differences with other comparable
species from the same area.
Key to Nephelium species
1. Ovary 1-locular.............................................................................................8. N. maingayi
Ovary 2(4)-locular, very rarely 1-locular...........................................................................2
2. Fruit wall hard, corky, 36 mm thick......................................................2. N. compressum
Fruit wall coriaceous or eggshell-like, rarely more than 1(7) mm thick............................3
3. Fruits warty, knobbly or spiny.............................................................................................4
Fruits with filiform, strap-shaped, tongue-shaped or spine-like appendages.......................6
4. Fruits glabrous. Leaflets ovate to elliptic....................................................1. N. aculeatum
At least the upper half of the fruit-appendages densely puberulous. Leaflets elliptic to
obovate................................................................................................................................5
5 Leaf axes densely tomentulose, sometimes glabrescent. Leaflets hairy above, except
(sometimes only) for the midrib, early glabrescent; midrib and lateral veins densely
tomentose beneath. Calyx to 3 mm high....................................................4. N. daedaleum
Leaf axes sparsely puberulous to glabrous. Leaflets glabrous above, sparsely sericeous
beneath, sometimes glabrescent. Calyx to 1.75 mm high..................10. N. melanomiscum
6. Fruit appendages distinctly puberulous all over..................................................................7
Fruits subglabrous or exceptionally with a hair-tuft on top of the appendages...................9
7. Petiolules not grooved above. Leaflets slightly falcate........................7. N. macrophyllum
Petiolules grooved above. Leaflets symmetrical.................................................................8
8. Intercostal veins and veinlets densely reticulate, faintly conspicuous above, inconspicuous
beneath. Leaflets drying brown..................................................................9. N. meduseum
Intercostal veins and veinlets more or less laxly reticulate and conspicuous on both sides.
Leaflets drying green.................................................................13. N. reticulatum (in part)
334
9. Domatia present.................................................................................................................10
Domatia absent..................................................................................................................14
10. Reticulation dense on the upper side of leaflets.................................................................11
Reticulation lax on the upper side of leaflets.....................................................................12
11. Fruits to 2.5x1.5 cm; fruit wall to 1 mm thick; appendages to 10 mm long. Intercostal
veins reticulate.............................................................................................5. N. havilandii
Fruits to 6x3.5 cm; fruit wall to 2.5 mm thick, appendages usually more than 10 mm
long. Intercostal veins often tending to be scalariform.................6. N. lappaceum (in part)
12. Inflorescences axillary or pseudoterminal, ending in a vegetative bud; infructescences
thus lateral. Fruits with bulbous-based spines, or sometimes warty. Leaflets when dried
often curled or rolled-up.....................................................12. N. ramboutan-ake (in part)
Inflorescences usually truly terminal, not ending in a vegetative bud; infructescences thus
also terminal. Fruit appendages subfiliform or sometimes more or less tongue-shaped.
Leaflets when dried mostly flat.........................................................................................13
13. Leaflets parchment-like, apex not mucronate; midrib raised above; intermediate lateral
veins well-developed; intercostal veins reticulate. Fruits to 3 x2.25 cm; appendages sparse
and short, to 7.5 mm long..........................................................................15. N. uncinatum
Leaflets coriaceous, apex usually mucronate; midrib usually sunken above, intermediate
lateral veins mostly inconspicuous; intercostal veins tending to be scalariform. Fruits to
6 x3.5 cm; appendages dense, to 2 cm long....................................9. N. lappaceum (in
part)
14. Fruit appendages of long spines. Leaflets when dried often curled or rolled-up...................
............................................................................................12. N. ramboutan-ake (in part)
Fruit appendages mostly filiform. Leaflets mostly flat when dried...................................15
15. Leaflets, at least on the midrib, nearly always hairy below...............................................16
Leaflets glabrous below or at most with a few minute hairs along the midrib..................17
16. Twigs and leaves usually densely hairy, rarely glabrous. Inflorescences axillary and/or
pseudoterminal.........................................................................................3. N. cuspidatum
Twigs and leaves sparsely hairy to glabrous. Inflorescences mostly truly terminal..............
......................................................................................................6. N. lappaceum (in part)
17. Apical part of fruit appendages nipple-like, 12 mm long. Leaflets hardly short- and
broad-acuminate.......................................................................................11. N. papillatum
Apical part of fruit appendages thread- or hair-like, to 15 mm long. Leaflets rather
abruptly caudate-acuminate...............................................................................................18
18. Reticulation of intercostal veins and veinlets dense and not elongatedly parallel to the
lateral veins. Apical part of fruit appendages to 10 mm long, slightly hairy.........................
...................................................................................................13. N. reticulatum (in part)
Reticulation of intercostal veins and veinlets lax and elongatedly parallel to the lateral
veins. Apical part of fruit appendages to 15 mm long, glabrous............14. N. subfalcatum
335
1. Nephelium aculeatum Leenh.
(Latin, aculeatus = with spine-like appendages; the fruit)

l.c. (1986) 382, l.c. (1994) 672. Type: Meijer SAN 48559, Borneo, Sabah (holotype L; isotypes K, SAN).
Medium-sized tree. Leaves each with 4 pairs of leaflets; petioles 810 cm long, 2 mm thick,
terete; rachis tomentulose to subglabrous. Leaflets stiff parchment-like, glabrous above,
puberulous on midrib and lateral veins beneath, in between veins fairly densely sericeous;
domatia absent; ovate to elliptic, 919 x4.55.5 cm; base rounded to obtuse, slightly
attenuate, usually parallel-sided in the lower two third, apex gradually acuminate, acumen
rather long and broad; midrib grooved above, lateral veins 0.751 cm apart, hardly raised
above, intermediate lateral veins absent, intercostal veins fairly densely scalariform, slightly
raised on both sides, veinlets finely reticulate, slightly raised above, only partly visible
beneath; petiolules 58 mm long, narrowly deeply grooved, without a rib above.
Inflorescences probably axillary. Flowers unknown. Fruits 12-lobed, oblique-ellipsoid, c.
3 x 2 x 1.75 cm, glabrous; appendages fairly dense, spine-like, c. 2 mm long.
Distribution. Endemic to Borneo; In Sabah, uncommon.
Ecology. Secondary forest. Fruiting in August.
Uses. Fruits edible.
2. Nephelium compressum Radlk.
Fig. 17D.
(Latin, compressus = laterally flattened; the fruit)
Sapind. Holl.-Ind. (1879) 9, 28; Merrill l.c. (1921) 360; Radlkofer in Engler l.c. (1933) 980; Masamune l.c. 428; Anderson
l.c. 317; Leenhouts l.c. (1986) 382, l.c. (1994) 672; Whitmore, Tantra & Sutisna l.c. 313. Type: Beccari PB 1268, Borneo,
Sarawak (holotype M; isotypes BO, FI, K, NY).
Tree. Leaves each with 35-pairs of leaflets; petioles 4.512 cm long, 24 mm thick,
grooved or subterete towards the apex above; rachis tomentose, glabrescent. Leaflets stiffcoriaceous, tomentose on the midrib above, soon glabrescent, densely tomentose on the
midrib, lateral veins and intermediate lateral veins beneath, thinly in between; domatia
absent; elliptic to obovate, 4.518 x2.757.5 cm; base rounded (upper leaflets), obtuse or
acute, sides curved, margin repand, apex apiculate or with a short, broad, acute to rounded
acumen; midrib slightly raised above, lateral veins 0.51.25 cm apart, sunken above,
intermediate lateral veins well-developed; intercostal veins densely scalariform, slightly
sunken or flat above, visible beneath, veinlets minutely reticulate, visible above, obscure
beneath; petiolules 25 mm long, broadly and shallowly grooved with a strong median rib
above. Inflorescence a widely branched terminal or pseudoterminal thyrses. Flower: sepals
free or nearly so, 22.2 mm long; petals 5, 1.251.8 x1.11.4 mm, white, blade woolly
outside, base sometimes woolly inside, claw 0.30.4 mm long, woolly outside, scale 0.20.25
mm high, consisting of 2 connate, reflexed, rounded lobes; disc glabrous; stamens 7 or 8;
ovary 2-locular. Fruits flattened ellipsoid, 44.5 x2.5 x1.251.75 cm, tomentulose, partly
glabrescent, rugose-warty; fruit wall hard, corky, 36 mm thick.
Distribution. Endemic to Sarawak, near Kuching, uncommon.
336
Fig. 17. Nephelium maingayi (AC); N. compressum (D); N. cuspidatum (E); N. daedaleum (F). A, flowering leafy twig; B,
female flower; CF, fruits. (AB from SAN 65365, C from SAN 38799, D from Beccari PB 1268, E from S. 27687, F from
SAN 50490.)
337
Ecology. Flowering in March; fruiting in December. This species is characterized by its

peculiar fruits which appear to be dispersed by water rather than by animals at least as judged
from the thick corky fruit wall and the thin sarcotesta.
Taxonomy. Vegetatively, the species is closely related to N. cuspidatum var. eriopetalum.
The best characters to distinguish these two taxa are: the slightly bullate leaflets with sunken
lateral veins and usually slightly sunken intercostal veins on the upper surface in N.
compressum, against the hardly bullate leaflets with narrowly grooved lateral veins and
usually slightly raised intercostal veins in N. cuspidatum var. eriopetalum. In flowering
specimens, N. compressum differs from N. cuspidatum var. eriopetalum by its rather large
and widely branched, thyrsoid, terminal or pseudoterminal inflorescences, the nearly free and
slightly imbricate sepals, and the well-developed corolla. In N. cuspidatum var. eriopetalum,
the inflorescences are mainly axillary or sometimes even born on leafless twigs and stems,
and composed of a few spike-like branches; the sepals are more connate and valvate; and the
petals are few and reduced in size or absent.
3. Nephelium cuspidatum Blume
(Latin, cuspidatus = tapering gradually; the leaflet apex)
Fig. 17E.
Rumphia 3 (1847) 110; Merrill l.c. (1921) 360; Radlkofer in Engler l.c. (1933) 977; Masamune l.c. 428; Anderson l.c. 312;
Leenhouts l.c. (1986) 385, l.c. (1994) 674; Yap l.c. 451; Whitmore, Tantra & Sutisna l.c. 215. Type: Korthals, s.n. (= L.
sheet no. 908. 269-1369, 1389, 1470 and 1490), Borneo (L). Synonyms: N. dasyneurum Radlk., Rec. Bot. Surv. India 3
(1907) 354, in Engler, Planzenr. 98 (1933) 978; N. obliquinerve Radlk. l.c. (1907) 354, in Engler l.c. (1933) 978.
Tree to 40 m tall, 80 cm diameter, sometimes with small buttresses, rarely a shrub. Leaves
each with (1)29(13) pairs of leaflets; petioles 2.521 cm long, 16 mm thick, terete or
subterete with a longitudinal groove above; rachis mostly persistently hairy. Leaflets
coriaceous or chartaceous, glabrous to variably hairy along the midrib and on the lower
lateral veins or rarely hairy all over above, variably hairy all over beneath, between lateral
veins often minutely sericeous, domatia absent; elliptic to linear, 635 x 1.7512.5 cm; base
acute to broadly rounded, exceptionally subcordate, mostly variably attenuate, sides curved to
straight and parallel, apex rounded to acute, mostly acuminate, acumen to 2.5 cm long,
slender to sometimes broad, acute to sometimes obtuse; midrib sunken to slightly raised
above, lateral veins 0.52 cm apart, slightly raised to slightly grooved above, intermediate
lateral veins variably developed, intercostal veins and veinlets either together closely or
sometimes laxly reticulate, or the former more or less clearly scalariform, at least the veinlets
often hardly visible beneath; petiolules 27.5(15) mm long, variably grooved or sometimes
flat above, with or without a median rib. Inflorescences mostly pseudoterminal in the axils of
upper leaves, sometimes terminal, or borne on leafless twigs or stems, often long pendulous
racemes or spikes. Flower: sepals hardly to halfway connate, 1.12.5 mm long; petals mostly
absent, if present often reduced in number; disc hairy or glabrous; stamens (4)78(9);
ovary 2-locular. Fruits ellipsoid to sometimes globose, 24 x 23 cm, glabrous or sometimes
slightly hairy at the tip of the appendages; appendages dense, filiform to narrowly strapshaped, up to 2 cm long, or sometimes ligulate and 56 mm long, straight or often curved or
curled, globose at base, pyramidal, or triangular and in the latter case often connate; fruit
wall thin-coriaceous.
338
Vernacular names. Sabahkalambuko (Murut), lemag (Dusun Kinabatangan), manggalum

(Kadazan), mengalim (Dusun Kinabatangan), rambutan (Malay). Sarawakbuah kebuau
(Iban), buah mersilat (Punan), buah sibau (Iban), kapusong (Kayan, Land Dayak).
Distribution. Myanmar, Thailand, Cambodia, Vietnam, Peninsular Malaysia, Sumatra, W
Java, Borneo, and Palawan. Common in Sabah and Sarawak.
Ecology. Usually an understorey tree of primary or sometimes secondary mixed dipterocarp
forest on well-drained land, mainly on ridges and slopes, rarely on plains or river banks;
preferably on fertile clay loam derived from igneous rocks, rarely on sandstone or limestone;
from sea level to 800 m. Flowering mainly in November to March; fruiting mainly in May to
September.
Uses. Locally cultivated as a fruit tree. The wood is sometimes used for construction but is
not of a very good quality (Burkill, EPMP (1935) 1544, 1548; Heyne, Nutt. Pl. Ned.-Indi,
ed. 2 (1927) 997).
In Sabah and Sarawak, five varieties with two subvarieties are recognized.
Key to varieties and subvarieties
1. Intercostal veins laxly reticulate, prominent on both sides of leaflets. Inflorescences
terminal.................................................................................................................................
var. robustum (Radlk.) Leenh.
l.c. (1986) 391, l.c. (1994) 677. Basionym: N. robustum Radlk., Leafl. Philip. Bot.
5(1913) 1607, in Engler l.c. (1933) 966. Type: Elmer 12934, Philippines, Palawan,
Puerto Princesa, Mt. Pulgar (holotype M; isotypes BO, FI, L, NY, U).
Leaflets c. 35 x510 cm, margins straight and parallel to slightly curved; midrib flat to
sunken above, lateral veins slightly raised to sunken above, intermediate lateral veins
only slightly developed; petiolules 715 mm long. Flowers: sepals 2550% connate,
1.31.8 mm long; petals absent. Fruit appendages dense, narrowly strap-shaped to
filiform, gradually thickened and broadened toward the base, curled, c. 1.5 cm long.
Palawan (one collection) and Borneo. Common in Sabah; also in Brunei and
Kalimantan.
Intercostal veins finely reticulate, often inconspicuous to hardly visible on the upper side
of leaflets. Inflorescences pseudoterminal or axillary..........................................................2
2. Leaflet lower surface in between lateral veins glabrous or sparsely appressed-hairy (the
venation visible). Sepals 2040% connate..........................................................................3
Leaflet lower surface in between lateral veins minutely sericeous or more-or-less hairy
(the venation obscure). Sepals hardly to slightly connate....................................................4
3. Leaf rachis hairy. Largest leaflets 2035 cm long, lateral veins not conspicuously dense,
those in the central part of leaflets to 2 mm apart. Fruit appendages 1020 mm
long...........
var. eriopetalum (Miq.) Leenh.
l.c. (1986) 389, l.c. (1994) 676; Yap l.c. 451. Basionym: N. eriopetalum Miq., Fl.
Ned. Ind.,
339
Suppl. (1861) 508, Ridley l.c. (1922) 502, Radlkofer in Engler l.c. (1933) 979,
Anderson l.c. 312, Corner l.c. 684. Lectotype (Leenhouts, 1986): Junghuhn, s.n. (= L.
sheet no. 908. 2691390), Sumatra (L).
Leaflets 312.5 cm broad, margin nearly straight and parallel to mostly slightly
curved; midrib slightly raised to slightly sunken above, lateral veins finely grooved or
flat above, intermediate lateral veins often many and well-developed; petiolules 210
mm long. Flowers: sepals 1.82.5 mm long; petals absent to 5 and reduced in size.
Fruit appendages dense, strap-shaped to sometimes filiform, straight to sometimes
curled, base swollen, pyramidal or broadened.
Peninsular Malaysia, Sumatra, Borneo, W Java. Common in Sabah, uncommon in
Sarawak; also in Brunei.
Leaf rachis early glabrescent. Largest leaflets to 17 cm long; lateral veins dense, those in
the central parts of leaflets to 0.75 mm apart. Fruit appendages to 8 mm
long......................
var. multinerve (Radlk.) Leenh.
l.c. (1986) 389, l.c. (1994) 676. Basionym: N. multinerve Radlk., Sapind. Holl.-Ind.
(1879) 9, 27, in Engler, l.c. (1933) 979, Merrill l.c. (1921) 360, Masamune l.c. 429.
Type: Beccari PB 2820, Borneo, Sarawak (FI).
Leaflets 68.5 broad, margins curved; midrib almost glabrous, narrowly grooved
above, lateral veins almost glabrous, slightly raised above, intermediate lateral veins
hardly developed; petiolules 512.5 mm long. Flowers: sepals c. 1.75 mm long; petals
not seen (present according to Radlkofer l.c., 1933). Fruit appendages very dense,
arranged in longitudinal rows, narrowly triangular to strap-shaped.
4. Leaflets linear, margins parallel, glabrous or early glabrescent beneath; petiolules
narrowly shallowly grooved above, sometimes with a faint median rib. Twigs early
glabrescent. Inflorescences pseudoterminal. Fruit appendages ligulate-triangular...............
var. cuspidatum subvar. cuspidatum
Leaflets 1.755 cm broad, apex acuminate, acumen long and slender; midrib sunken
above, lateral veins flat to slightly grooved above, intermediate lateral veins often
well-developed, petiolules 47.5 mm long. Flowers: sepals 1.11.2 mm long; petals
absent, or one and reduced in size. Fruit appendages dense, 56 mm long, slightly
curved.
Endemic to Borneo; uncommon in Sabah.
Leaflets elliptic, margins gradually curved, more-or-less sericeous beneath; petiolules
broadly grooved with a strong median rib above. Twigs densely hairy. Inflorescences
axillary. Fruit appendages narrowly strap-shaped with bulbous base....................................
var. ophioides (Radlk.) Leenh. subvar. beccarianum (Radlk.) Leenh.
l.c. (1986) 390, l.c. (1994) 677. Basionyms: N. ophelioides Radlk. l.c. (1879) 77, 78,
Ridley l.c. (1922) 502, Corner l.c. 686. Type: Maingay KD 543, p.p., Malacca
(holotype B, destroyed; isotype K); N. beccarianum Radlk. l.c. (1879) 9, 27, in Engler
l.c. (1933) 978, Merrill l.c. (1921) 359, Masamune l.c. 428, Anderson l.c. 312.
Lectotype (Leenhouts, 1986): Beccari PB 2279, Borneo, Sarawak, Mt. Matang (FI).
Leaflets 25 cm broad, midrib slightly raised to slightly sunken above, lateral veins
flat to slightly grooved above, intermediate lateral veins often rather strongly
developed; petiolules 27 mm long. Flowers: sepals c. 1.8 mm long; petals absent
(present according to Radlkofer l.c., 1933). Fruit appendages to 1 cm long.
Endemic to Borneo; common in Sabah and Sarawak.
340
4. Nephelium daedaleum Radlk.
(Latin, daedaleus = with sinuous, irregularly plaited or jagged lines; the fruit wall)
Fig. 17F.
Sapind. Holl.-Ind. (1879) 9, 27, in Engler l.c. (1933) 980; Merrill l.c. (1921) 360; Masamune l.c. 429; Leenhouts l.c. (1986)
391, l.c. (1994) 677. Type: Beccari PB 2818, Borneo, Sarawak (holotype FI; isotype K).
Tree to 33 m tall, 70 cm diameter, sometimes with buttresses. Leaves each with 13(5)
pairs of leaflets; petioles 29.5 cm long, 12 mm thick, terete to subterete; rachis
tomentulose, sometimes glabrescent. Leaflets parchment-like, hairy all over above, early
glabrescent except the midrib, tomentose on midrib and lateral veins beneath, densely
sericeous in between lateral veins, domatia absent; elliptic, 6.518 x2.57 cm; base acute to
rounded, margins curved, apex not to abruptly acuminate, acumen short and broad to long
and slender, obtuse; midrib slightly raised above, lateral veins 0.51.25 cm apart, slightly
grooved above, intermediate lateral veins often well-developed, intercostal veins finely
reticulate, inconspicuous above, invisible or sometimes slightly raised and somewhat
scalariform beneath; petiolules 59 mm long, variably grooved, and mostly with a median rib
above. Inflorescences terminal. Flowers: sepals almost free, c. 3 mm long; petals 4 or 5,
1.752 x1.52 mm, blade auricled to funnel-shaped, densely woolly outside, claw to 1 mm
long, densely woolly outside; disc glabrous; stamens 8 or fewer; ovary 2-locular. Fruits
flattened-ellipsoid, base bulging at the abaxial side, c. 4 x2.25 cm, c. 1.5 cm thick, densely
short-hairy, fairly densely set with pyramidal warts of c. 1.5 mm high, deeply irregularly
fissured; fruit wall coriaceous, 22.5 mm thick.
Vernacular names. Sabahrambutan (Malay). Sarawakkalas (Iban).
Distribution. Endemic to Borneo; common in Sabah, uncommon in Sarawak.
Ecology. Primary lowland kerangas or mixed dipterocarp forest on slopes or ridges; on
sandstone, sandy clay, or loam soils; up to 570 m. Flowering in May, September; fruiting in
June, August, September, November.
Uses. Sometimes cultivated for its edible fruit.
5. Nephelium havilandii Leenh.
(G. D. Haviland, 18571901, first Sarawak Medical Officer)

l.c. (1986) 394, l.c. (1994) 678. Type: Haviland 2241, Borneo, Sarawak (holotype K; isotypes L, SING).
Tree c. 8 m tall. Leaves each with 12 pairs of leaflets; petioles 2.255 cm long, 11.5 mm
thick, slightly hollowed above; rachis subglabrous. Leaflets coriaceous, glabrous above,
thinly minutely sericeous beneath, domatia absent or present as exposed glands or covered
by a tuft of hairs; elliptic, 69 x34.25 cm; base slightly asymmetric, acute, attenuate,
margins curved, apex narrowly rounded, short-acuminate or not; midrib slender and slightly
raised above, lateral veins 0.51.5 cm apart, slightly raised above, intermediate lateral veins
well-developed or sometimes absent, intercostal veins reticulate, moderately coarse and
slightly raised above, inconspicuous beneath; petiolules 35 mm long, broadly rather deeply
grooved and often with a faint median rib above. Inflorescences axillary. Flowers: sepals
free, 1.52.5 mm long; petals 5, elliptic, 0.81.4 mm long, with a short claw, without a scale,
either glabrous outside
341
and woolly in the lower half inside, or woolly mainly in the lower half on both sides; disc
woolly; stamens c. 7; ovary 2-locular. Fruits ellipsoid, c. 22.5 x1.25 cm, glabrous,
appendages dense, 810 mm long, base bulbous, apical part triangular in cross section,
curved; fruit wall coriaceous, c. 0.5 mm thick.
Distribution. Endemic to Borneo; Sarawak, near Kuching, uncommon; also very uncommon
in W Kalimantan.
Ecology. Flowering in October; fruiting in February, November.
6. Nephelium lappaceum L.
(Latin, lappaceus = bur-like; the fruit appendages)

Mant. Pl. 1 (1767) 125; Ridley l.c. (1922) 499; Radlkofer in Engler l.c. (1932) 957, fig. 24; Backer & Bakhuizen. f. l.c.
(1965) 138; Anderson l.c. 312; Leenhouts l.c. (1986) 398, l.c. (1994) 680; Whitmore & Tantra l.c. 215; Corner l.c. 684; Yap
l.c. 453; Whitmore, Tantra & Sutisna l.c. 314. Neotype (Leenhouts, 1986): Bogor Botanic Garden III.H.10 (= Carocci-Buzi
190, Nedi 12, Sutrisno 71) (L; isoneotypes BO, M, NY, U).
Tree or sometimes shrub. Leaves 1-foliolate or each with up to 8 pairs of leaflets; petioles
1.516 cm long, 13.5 mm thick, terete, subterete, or sometimes grooved above; rachis
variably hairy, early to late glabrescent. Leaflets coriaceous, glabrous or sometimes slightly
hairy on the midrib above, variably hairy beneath, domatia common to absent; ovate to
obovate, 528 x210.5 cm; base acute to rounded, attenuate or not, margins strongly curved
to nearly straight and parallel, apex acute to truncate; midrib slightly raised to slightly sunken
above, lateral veins 0.52 cm apart, slightly raised to slightly sunken above, intermediate
lateral veins mostly inconspicuous, intercostal veins finely or coarsely reticulate, the veinlets
often tending to be scalariform, often raised above; petiolules 512 mm long, broadly and
shallowly grooved with or without median or lateral rib or narrowly grooved or flat without
rib above. Inflorescences axillary, pseudoterminal, or sometimes terminal. Flowers: sepals
almost free to more than halfway connate, 0.72.1 mm long; petals mostly absent, sometimes
up to 4, reduced in size, claw c. 1.1 mm long, blade c. 0.5 x0.5 mm, margin infolded and
connate towards the base, glabrous or with a few long hairs outside, margin long-ciliate,
woolly inside; disc hairy or glabrous; stamens (4)58(9); ovary 2(3)-locular. Fruits
hardly stalked, ellipsoid to subglobose, to 6 x3.5 cm, glabrous; fruit wall coriaceous, to 2.5
mm thick; appendages sparse or dense, base bulbous or broad, tapering towards the strapshaped or filiform, more or less curved, 0.52-cm-long upper parts.
Vernacular names. Preferred namerambutan (Malay). Sabahrangalau (Dusun Ranau).
Sarawakbuah abong (Kenyah), buah maha (Punan Tutoh), lung bipuyoh (Berawan),
sangan (Bidayuh Sadong), situgog (Bidayuh).
Distribution. Yunnan, Hainan, Indo-China, Peninsular Malaysia, Sumatra, Java, Borneo,
Philippines, and Sulawesi. Common in Sabah and Sarawak; also in Brunei and Kalimantan.
Taxonomy. The only constant morphological difference between N. lappaceum and N.
ramboutan-ake is in the fruit appendages. In N. lappaceum, the basal part of fruit appendage
is small and inconspicuous and the upper part is thread-like and long; in N. rambutan-ake, on
the other hand, the basal part of fruit appendage is relatively large, triangular to ovoid, and
342
tapering towards the short and broad, often thick and stiff apical part. In Sabah and Sarawak,
3 varieties are recognised.
Key to varieties
1. Leaflets hardly glaucous beneath, mostly widest above the middle, margins strongly
curved, apex rarely acuminate; lateral veins widely spreading and strongly curved.............
var. lappaceum
Yap l.c. 453; Lenhouts l.c. (1994) 682. Synonyms: N. glabrum Reinw. ex Blume, Cat.
(1923) 111, nom. nud., Masamune l.c. 428; N. subferrugineum Radlk. l.c. (1879) 9,
77, in Engler l.c. (1933) 956, Masamune l.c. 428, Anderson l.c. 313; N. obovatum
Ridl., Kew Bull. (1933) 191, Radlkofer in Engler l.c. (1934) 1500, Masamune l.c.
429, Anderson l.c. 313.
Tree or shrub to 27 m tall, 70 cm diameter; buttresses to 1.5 m high. Leaves 1foliolate or each with up to 5 pairs of leaflets; petioles 1.512 cm long; rachis variably
hairy, glabrescent. Leaflets 522 x2.510.5 cm, glabrous above, sparsely hairy on the
midrib and lateral veins beneath, in between lateral veins mostly glabrous, sometimes
sparsely or densely appressed-hairy, domatia present; base obtuse or rounded, apex
obtuse or rounded, sometimes truncate to slightly emarginate, apiculate or slightly
acuminate, rarely tapering into a cuneate and acute acumen; lateral veins 0.752 cm
apart, intercostal veins and veinlets coarsely to finely reticulate; petiolules 210 mm
long. Flowers: sepals 45(7), mostly slightly connate, 1.12.1 mm long, mostly
thinly to densely appressed short-hairy outside, mostly appressed long-hairy to woolly
inside; stamens 58. Fruits large; fruit wall thin.
Thailand, Peninsular Malaysia, Sumatra, Borneo, W and C Java, Philippines, and
possibly Ceram. Commonly cultivated for its fruits, also in other parts of the tropics,
and doubtless not rarely escaped and become naturalized. In Sabah and Sarawak
common; also in Brunei and Kalimantan.
Leaflets usually distinctly glaucous beneath, mostly widest about or below the middle,
margins often nearly straight and parallel, apex usually acuminate; lateral veins fairly
steep and only slightly curved.............................................................................................2
2. Leaflets mostly to only 4 cm wide; midrib often glabrous beneath, intercostal veins and
veinlets reticulate, with the veins often tending to be scalariform.........................................
var. pallens (Hiern) Leenh.
l.c. (1986) 402, l.c. (1994) 683. Basionym: N. mutabile Blume var. pallens Hiern in
Hooker f., Fl. Br. Ind. 1 (1875) 687. Lectotype (Leenhouts, 1986): Maingay 1527 =
KD 454, p.p., Malacca (K). Synonym: N. chryseum Blume l.c. (1847) 105, Merrill l.c.
(1921) 360, Radlkofer in Engler l.c. (1933) 962, Masamune l.c. 428, Anderson l.c.
312.
Tree or shrub to 44 m tall, 125 cm diameter; buttresses to 4 m tall. Leaves each with
(1)35(8) pairs of leaflets; petioles 2.512 cm long; rachis densely short-hairy to
glabrous. Leaflets 5.512(20) x(2)34(7) cm, sometimes slightly sickle-shaped,
upper surface glabrous or sometimes slightly puberulous on the midrib, lower surface
mostly minutely sericeous, domatia common to absent; base acute to sometimes
obtuse or rounded, apex narrowly rounded to acute or tapering into rounded or acute
acumen; lateral veins 0.51(2) cm apart; petiolules (1.5)38 mm long. Flowers:
sepals 45, rarely 6, up to halfway connate, 12 mm long, sparsely to densely
appressed, short-hairy outside, densely velutinous inside; stamens (4)58(9). Fruits
343
to 5 x 3.5 cm; stamens (4)58(9). Fruits to 5 x 3.5 cm; fruit wall 1.52.5 mm thick.
China, Thailand, Indo-China, Peninsular Malaysia, Sumatra, Borneo, Philippines and
Sulawesi. Common in Sabah, uncommon in Sarawak; also in Brunei and Kalimantan.
Leaflets to 10 cm wide; midrib densely tomentulose, intercostal veins mostly distinctly
scalariform, sometimes tending to be coarsely reticulate, veinlets reticulate........................
var. xanthioides (Radlk.) Leenh.
l.c. (1986) 403, l.c. (1994) 683. Basionym: N. xanthioides Radlk. l.c. (1879) 9, 77, in
Engler l.c. (1933) 966, Beccari, For. Born. (1902) 601, Merrill l.c. (1921) 360,
Masamune l.c. 429. Type: Beccari PB 2849, Borneo, Sarawak (FI).
Tree to 30 m tall, 60 cm diameter; buttresses to 2 m high. Leaves each with 25 pairs
of leaflets; petioles 1016 cm long; rachis tomentulose. Leaflets 928 x310 cm,
upper surface glabrous or sometimes sparsely puberulous on the basal half of the
midrib, lower surface densely tomentulose on the midrib or sparsely to densely
appressed short-hairy on the midrib and lateral veins or sparsely minute-hairy,
domatia rare; base obtuse to rounded, apex tapering into a slender, rounded or acute
acumen; lateral veins 0.61.75 cm apart; petiolules 512 mm long. Flowers: sepals 4
5, to halfway connate, 0.71.6 mm long, densely hairy on both sides; stamens 56.
Fruits c. 3 x 2 cm; fruit wall c. 1 mm thick.
Endemic to Borneo. Uncommon in Sabah and Sarawak; also in Kalimantan.
7. Nephelium macrophyllum Radlk.
(Greek, makros = large; phullon = leaf)
Sapind. Holl.-Ind. (1879) 9, 27, in Engler l.c. (1933) 973; Merrill l.c. (1921) 360; Masamune l.c. 428;
Anderson l.c. 313; Leenhouts l.c. (1986) 406, l.c. (1994) 684. Type: Beccari PB 2500, Borneo, Sarawak (holotype FI;
isotypes M, NY).
Tree to 24 m tall, 40 cm diameter. Leaves each with 12 pairs of leaflets; petioles 47 cm

long, 12.5 mm thick, subterete to obtuse-angular above; rachis glabrous or sometimes very
sparsely minutely puberulous. Leaflets thick-coriaceous, glabrous above, sparsely puberulous
on midrib and lateral veins beneath, in between lateral veins densely minute-sericeous,
domatia present; elliptic, 1022.5 x510 cm; base acute to rounded, slightly attenuate,
margin curved, apex tapering into short acute acumen; midrib slightly raised above, lateral
veins 13 cm apart, slightly sunken above, few intermediate lateral veins well-developed,
intercostal veins tending to be scalariform, slightly raised or hardly visible above, slightly
raised beneath, veinlets finely reticulate, slightly raised to hardly visible above, hardly visible
beneath; petiolules 1.57 mm long, slightly bulging above. Inflorescences pseudoterminal.
Flowers known only from remains under the fruit; sepals probably only slightly connate, c. 2
mm long; petals present (?); disc glabrous; ovary 2- rarely 3-locular. Fruits ellipsoid, c.
3.5 x2 cm, appendages dense, c. 7.5 mm long, base bulbous, upper part tongue-shaped,
slightly curved, densely ferrugineous-puberulous; fruit wall coriaceous, c. 2 mm thick.
Distribution. Endemic to Borneo; Sarawak, from the Baram valley to Kuching, uncommon;
also in Brunei.
Ecology. In primary mixed dipterocarp forest, on sandy clay and clay loam soils, to 1000 m.
Fruiting in September.
344
Taxonomy. The gross morphology of fruits of the present species may superficially resemble
that of N. ramboutan-ake. However, in the latter species the fruits are glabrous and the
leaflets much thinner. N. macrophyllum sometimes shows some vegetative characters similar
to those of N. melanomiscum. However, the former is characterized by asymmetric, slightly
sickle-shaped leaflets, whereas the latter has symmetric leaflets. The fruits of both species are
distinctly different.
8. Nephelium maingayi Hiern
(A.C. Maingay, 18361869, botanist of the East India Company)
Fig. 17AC.
In Hooker f., Fl. Br. Ind. 1 (1875) 688; Radlkofer in Engler l.c. (1933) 964; Anderson l.c. 313; Leenhouts l.c. (1986) 407, l.c.
(1994) 685; Yap l.c. 454. Syntypes: Maingay 1120, 1120A, 1524A, Malaya (K).
Tree to 40 m tall, 90 cm diameter; sometimes buttresses to 1.4 m high. Leaves 1-foliolate or

each with up to 3(5) pairs of leaflets; petioles 110 cm long, 13 mm thick, terete to
subterete; rachis early glabrescent or sometimes glabrous. Leaflets parchment-like, glabrous
or sometimes very sparsely hairy on midrib and lateral veins beneath, domatia absent; elliptic
to obovate, 5.7522 x2.759 cm; base rounded to acute, mostly attenuate, margin curved,
apex without or with an obtuse to acute acumen; midrib sunken to slightly raised above,
lateral veins 0.52.5 cm apart, slightly grooved to slightly raised above, intermediate lateral
veins absent, intercostal veins and veinlets coarsely reticulate, slightly raised on both sides;
petiolules 417.5 mm long, grooved, and without rib above. Inflorescences axillary to
terminal. Flowers: sepals from less than halfway to nearly completely connate, 11.3 mm
long; petals absent; disc hairy or glabrous; stamens 46; ovary 1-locular, style lateral, with 1
stigma. Fruits more-or-less flattened-ellipsoid, 22.75 x1.251.75 cm, x11.25 cm thick;
stalk 23 mm long; remnant of style present in the form of a small point or hook just above
the fruit stalk; surface variably warty, slightly puberulous around and especially beneath the
style remnant, otherwise glabrous; fruit wall coriaceous, 11.5 mm thick.
Vernacular names. Sabahbuah sungkit (Kedayan), kolomundui (Kadazan). Sarawak
mujau (Iban), serait (Malay).
Distribution. Peninsular Malaysia, Sumatra, Borneo. Common in Sabah and Sarawak; also
Ecology. In primary mixed dipterocarp and secondary forest on flat land (often in peat
swamps and periodically flooded riverbanks), slopes, and ridges; on peat, sandy or lownutrient clay soils, mostly at low to medium altitudes, exceptionally up to 10001600 m.
Flowering mainly in JanuaryApril, JulyOctober; fruiting in JanuaryApril, August
November.
Uses. The timber is of a good quality and is used for many purposes; the sarcotesta is edible
but of no economic importance (Heyne, Nutt. Pl. Ned.-Indi, ed. 2 (1927) 998; Burkill,
EPMP (1935) 1544, cited as N. glabrum).
9. Nephelium meduseum Leenh.
Fig. 18B.
(Latin, medusa = one of the three Gorgons, whose head bears serpents; the curled fruit appendages)
345
Fig. 18. Nephelium papillatum (A); N. meduseum (B); N. ramboutan-ake (C); N. uncinatum (D). A, fruiting leafy twig; BD,
fruits. (A from SAN 41845, B from S. 32399, C from FRI 10648, D from Hotta 12984.)
346
l.c. (1986) 409, l.c. (1994) 686. Type: Yii S. 41142, Borneo, Sarawak, 1st Div., G. Gaharu, Sabal FR (holotype K; isotypes
KEP, L, MO, SAN, SAR).
Tree to 27 m tall, 50 cm diameter; buttresses to 1.2 m high. Leaves each with 25 pairs of
leaflets; petioles 27 cm long, 1.52 mm thick, subterete; rachis puberulous, glabrescent.
Leaflets thin-coriaceous to stiff parchment-like, upper surface puberulous along the midrib,
mostly early glabrescent, lower surface glabrescent or glabrous except for the long-hairy
midrib and lateral veins, domatia absent; elliptic, 512.5 x2.55 cm; base acute to obtuse,
slightly attenuate, margin curved, apex attenuate-acuminate with a fairly long, broad and
obtuse acumen, or acutely apiculate, or not acuminate; midrib slender and slightly raised
above, lateral veins 0.751 cm apart, slightly sunken above, intermediate lateral veins mostly
well-developed though usually only a few per leaflet, intercostal veins and veinlets finely
reticulate and slightly raised above, veins coarsely reticulate to scalariform beneath, veinlets
reticulate and slightly raised to inconspicuous beneath; petiolules 312 mm long, broadly
shallowly grooved, mostly with a broad but not very strong median rib. Inflorescences
terminal. Flowers (only known from old ones and the remains under the fruit): sepals hardly
connate; petals at least 3, c. 1.6 mm long, claw slender, c. 1 mm long, blade ovate, c. 0.7 mm
wide, margin incurved at the basal half, sparsely woolly (except the apex) outside, glabrous
inside; disc glabrous; ovary 2-locular. Fruits ellipsoid to subglobose, 3.254 x2.53 cm,
densely fulvous puberulous; appendages dense, filiform, curled, c. 15 mm long, swollen at
base; fruit wall coriaceous, c. 1 mm thick.
Vernacular name. Sarawakrambutan hutan (Malay).
Distribution. Endemic to Borneo; in Sarawak uncommon; also in Brunei and Kalimantan.
Ecology. In primary mixed dipterocarp forest on hills, ridges, and slopes, on yellow sandy
clay, to 450 m. Fruiting in January and October.
Uses. The sarcotesta is edible.
10. Nephelium melanomiscum Radlk.
(Greek, melanos = black, miskos = stalk; the leaf rachis)

l.c. (1879) 74, in Engler l.c. (1933) 972; Merrill l.c. (1921) 360; Masamune l.c. 429; Anderson l.c. 313; Leenhouts l.c. (1986)
410, l.c. (1994) 687. Type: Beccari PB 3918, Borneo, Sarawak, Rejang river (FI). Synonym: Nephelium xerospermoides
Radlk., Leafl. Philipp. Bot. 5 (1913) 1608, in Engler l.c. (1933) 976, Anderson l.c. 313.
Tree to 25 m tall, 45 cm diameter; buttresses to 1.2 m high. Leaves 1-foliolate or each with
up to 5 pairs of leaflets; petioles 16 cm long, 12 mm thick, terete to subterete, sometimes
grooved above; rachis sparsely puberulous to glabrous. Leaflets parchment-like, glabrous or
puberulous on the base of the midrib above, sparsely minute-sericeous beneath, sometimes
glabrescent, domatia absent or present; elliptic to obovate, 3.514 x1.56 cm; base acute to
rounded, slightly attenuate, margin curved, apex rounded, emarginate, or tapering into a short
to long, narrow to broad, acute to rounded acumen; midrib slightly raised to sunken above,
lateral veins 0.751.5 cm apart, slightly raised to grooved above, intermediate lateral veins
mostly well-developed, intercostal veins and veinlets mostly well-differentiated, especially
347
beneath, more or less finely reticulate, often more prominent above than beneath; petiolules
38 mm long, broadly to narrowly deeply grooved above, with or without a median rib.
Inflorescences terminal, or sometimes pseudoterminal. Flowers: sepals less than 3065%
connate, 1.31.75 mm long; petals (according to Radlkofer) 2 or 3, reduced in size; disc
glabrous; stamens 6; ovary 23-locular. Fruits ellipsoid, c. 3.75 x2.5 cm, densely fulvous
puberulous at least in the upper half of the appendages; appendages dense, 1.52 mm high,
not differentiated into a basal and an apical part, those at the base of the fruit appear as
longitudinal ribs, those in the central part as triangles, and those towards the upper part as
pyramidal warts; fruit wall fairly hard, c. 0.75 mm thick.
Vernacular name. Sabahharut (Kedayan).
Distribution. Borneo, Philippines (Mindanao); uncommon in Sabah and Sarawak; also in
Brunei.
Ecology. Primary and old secondary forests on slopes, riverbanks, and hill tops on clay-rich
soil, and on fertile alluvial soil in mixed dipterocarp forest; to 1350 m. Flowering in October;
fruiting in January.
Uses. The sarcotesta is edible.
11. Nephelium papillatum Leenh.
Fig. 18A.
(Latin, papillatus = having papillae; the fruit wall)
l.c. (1986) 413, l.c. (1994) 688. Type: Mikil SAN 41845, Borneo, Sabah, Tambunan Dist., Mt. Trus Madi (holotype L;
isotype SAN).
Tree to 36 m tall, 85 cm diameter; buttresses to 2 m high. Leaves each with 13 pairs of

leaflets; petioles 25 cm long, 11.5 mm thick, subterete; rachis glabrous. Leaflets stiff
parchment-like, glabrous, domatia absent; elliptic, 4.510.5 x2.54.5 cm; base acute to
obtuse, attenuate, margin curved, apex hardly obtuse-acuminate; midrib slightly sunken to
flat above, lateral veins 0.51.25 cm apart, slightly raised above, intermediate lateral veins
well-developed, making the venation more or less irregular, intercostal veins and veinlets
rather coarsely reticulate, slightly more prominent above than beneath; petiolules 58 mm
long, narrowly deeply grooved without a median rib above. Inflorescences axillary, or
somewhat pseudoterminal. Flowers (known only from remains under the fruit): sepals 5, at
least sparsely puberulous outside; disc glabrous; stamens 7; ovary 2-locular. Fruits ellipsoid,
c. 2.25 x1.75 cm, glabrous; appendages fairly dense, c. 3 mm long, pyramidal, apical part
nipple-like, to 2 mm long; fruit wall rather woody, c. 1 mm thick.
Distribution. Endemic to Borneo; uncommon in Sabah.
Ecology. Primary lower montane to montane forest, at 13501950 m. Fruiting in November.
12. Nephelium ramboutan-ake (Labill.) Leenh.
(after a local name, rambutan ake)
Fig. 18C.
l.c. (1986) 415, l.c. (1994) 689; Yap l.c. 454. Basionym: Litchi ramboutan-ake Labill. in DC., Bull. Soc. Phil.
Math. Paris 2 (1801) 161 (Litsea). Type: Herb. Jussieu 11382 (P). Synonym: Nephelium mutabile Blume l.c.
(1847) 104, Merrill l.c. (1921) 360, Ridley l.c. (1922) 501 (excl. var. pallens), Radlkofer in
348
Engler l.c. (1933) 967, Masamune l.c. 429, Backer & Bakhuizen f. l.c. 138, Anderson l.c. 313, Corner l.c. 686.
Tree mostly less than 10 m, sometimes to 40 m tall, 80 cm diameter; buttresses to 2.4 m high.
Leaves 1-foliolate or each with 17 pairs of leaflets; petioles 0.7511.5 cm long, 0.752.5
mm thick, terete to subterete; rachis densely hairy to glabrous. Leaflets thin parchment-like to
thin-coriaceous, upper surface puberulous on the midrib to glabrous, lower surface sparsely
puberulous on the base of the midrib, densely minute-sericeous all over, to sometimes
glabrous, domatia mostly common, sometimes scarce or absent; narrowly elliptic, 4
20 x1.7511 cm; base acute or in lower leaflets obtuse to rounded, attenuate, margin mostly
curved, sometimes nearly straight, apex mostly acuminate, the acumen usually short, broad,
obtuse; midrib slightly raised or rarely slightly sunken above, usually appearing as a slender
rib, lateral veins 0.52 cm apart, slightly sunken to sometimes slightly raised above,
intermediate lateral veins variable, intercostal veins mostly reticulate, tending to be somewhat
scalariform or sometimes conspicuously so, slightly raised above, hardly visible beneath;
petiolules 38 mm long, mostly narrowly and deeply grooved without or with only a faint
median rib, or sometimes broadly and shallowly grooved with a stronger rib. Inflorescences
axillary or pseudoterminal. Flowers: sepals slightly or up to halfway connate, 12.75 mm
long; petals absent; disc glabrous; stamens 58; ovary 2(3)-locular. Fruits ellipsoid to
subglobose, 46.5 x2.55 cm, glabrous; pericarp coriaceous, to 7 mm thick, coarsely spiny
with the spines up to 1.5 cm long, bulbous and often confluent at base, or covered with short
tongue-shaped knobs.
Vernacular names. Sabahlongitam (Dusun), meritam (Kedayan, Malay), rambutan hutan
(Malay). Sarawakbuah mua, ma, melenjau, mujau, pudun (Iban).
Distribution. Assam, Myanmar, Peninsular Malaysia, Sumatra, Java (doubtful), Borneo,
Philippines, Maluku. Common in Sabah and Sarawak; also in Brunei and Kalimantan.
Ecology. Primary mixed dipterocarp or sometimes secondary forest on flat lands, slopes,
riverbanks, rarely in swamps; usually on clay, loam, more rarely on rocky soil, particularly
shale, basalt, or limestone; from sea level to 1950 m. Flowering mainly in February to April
and July to September; fruiting in May to July and October to December.
Uses. Cultivated as a fruit tree; the timber is also used for construction (Seibert, PROSEA 2
(1991) 233).
Taxonomy. This species is a rather variable one. The typical form found in Peninsular
Malaysia, Sumatra, Java, and Borneo, is characterized by its thin parchment-like leaflets
which, unlike those of most other species of Nephelium, are commonly rolled up in the
herbarium specimens, drying reddish brown above and glaucous beneath. The midrib is
slightly raised and slender above; the lateral veins are distinctly curved and rather steeply
ascending; domatia are common; and the reticulations are fairly lax and usually visible above.
The most distinct form restricted to Borneo is characterized by its leaves, each with 13
pairs of leaflets; the leaflets are relatively large, thin-coriaceous, glabrous above and often so
beneath, and lacking in domatia. The midrib is hardly raised to slightly sunken and more
rounded above; the lateral veins are dense, steeply ascending and nearly straight; the
349
intercostal veins and veinlets are clearly different from one another, with the former being
mostly rather densely scalariform and raised on both sides, and the latter being more faintly
and laxly reticulate. This form resembles N. lappaceum var. xanthioides but the latter
differs in its leaflets having midrib which is densely hairy beneath and at the base slightly
sunken above; and by its terminal inflorescences and infructescences.
N. ramboutan-ake may easily be confused with Dimocarpus longan var. echinatus because
these two taxa possess a strong resemblance in leaf and fruit characters. The main differences
are that in the former, the indumentum consists of solitary hairs and the twigs are brownish,
whereas in the latter, the indumentum is of stellate hair-tufts, and the twigs are white. The
fruits of N. rambutan-ake may also resemble those of N. macrophyllum; for differences
between the two species see there.
N. rambutan-ake is also closely allied to N. lappaceum var. pallens. However, the latter can
be distinguished by its midrib which is flat to sunken above, instead of mostly raised;
intercostal veins and veinlets which are slightly raised above, instead of inconspicuous; and
the rather dense instead of lax reticulations.
13. Nephelium reticulatum Radlk.
(Latin, reticulatus = net-like; the leaflet intercostal veins and veinlets)

l.c. (1879) 9, 27, in Engler l.c. (1932) 955; Beccari, For. Born. (1902) 600, 601; Merrill l.c. (1921) 360; Masamune l.c. 429;
Leenhouts l.c. (1986) 419, l.c (1994) 691. Type: Beccari PB 2819, Borneo, Sarawak (FI).
Tree to 25 m tall, 70 cm diameter, sometimes with small buttresses. Leaves each with 4(7)
pairs of leaflets; petioles 618.5 cm long, 1.53 mm thick, terete to slightly hollowed above;
rachis thinly puberulous, glabrescent. Leaflets parchment-like, glabrous or sometimes
sparsely puberulous on the midrib and sparsely appressed minute-hairy all over beneath,
domatia absent; narrowly elliptic to ovate, 6.520 x 2.55.5 cm; base rounded to acute,
attenuate, margin slightly curved, apex abruptly to gradually acuminate, acumen short to
fairly long, cuneate, acute; midrib appearing as a slightly sunken fine rib above, lateral veins
0.751.25(1.5) cm apart, slightly raised above, intermediate lateral veins variably
developed, intercostal veins and veinlets coarsely to minutely reticulate, slightly raised on
both sides; petiolules 210 mm long, narrowly and deeply to broadly and shallowly grooved,
without rib or with 3 ribs. Inflorescences terminal and axillary. Flowers often male and
female in the same inflorescence; sepals nearly free or up to c. 40% connate, 11.2 mm long;
petal absent or 5 to 1.8 mm long, claw c.1 mm, blade c. 1.5 mm wide, sparsely hairy outside,
densely hairy on the lower half inside; disc glabrous; stamens 58; ovary 2-locular. Fruits
ellipsoid, c. 4 x 2.5 cm; appendages dense, narrowly strap-shaped, bulbous to triangular at
base, curved, to c. 1 cm long, thinly puberulous, glabrescent; fruit wall coriaceous, c. 1 mm
thick.
Distribution. Endemic to Borneo; uncommon in Sabah, more common in Sarawak; also in
Kalimantan.
Ecology. Primary forest on flat or hilly country; to 400 m. Flowering in April, May, July,
August, October; fruiting in January.
Uses. Occasionally locally cultivated for its fruits.
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14. Nephelium subfalcatum Radlk.
(Latin, sub = somewhat, falcatus = sickle-shaped; the leaflets)

Rec. Bot. Surv. India 3 (1907) 353, in Engler l.c. (1933) 973; Leenhouts l.c. (1986) 420, l.c. (1994) 692; Yap l.c. 455. Type:
Forbes 3092, Sumatra, Palembang, Rawas river (holotype M; isotypes K, L, SING).
Tree to 35 m tall, 60 cm diameter; buttresses to 2 m high. Leaves 1-foliolate or each with up

to 5 pairs of leaflets; petioles 1.57 cm long, 0.751.5 mm thick, terete to subterete; rachis
sparsely puberulous, mostly glabrescent, or glabrous from the beginning. Leaflets thinly
parchment-like to coriaceous, glabrous or sometimes sparsely appressed minute-hairy along
the midrib beneath, domatia absent; narrowly ovate to elliptic, 415 x1.55 cm; base acute to
rounded, attenuate, margins slightly to sometimes strongly curved, sometimes nearly parallel,
apex obtuse or rounded or attenuate-acuminate, acumen short, broad and rounded to long,
slender and acute; midrib sunken to sometimes slightly raised above, lateral veins 0.51.5 cm
apart, slightly raised above, more or less distinctly looping and joining near the margin,
venation rather irregular because of the often great number of variably developed
intermediate lateral veins, intercostal veins clearly differentiated from veinlets, coarsely
reticulate, slightly raised on both sides; petiolules 310 mm long, narrowly and deeply or
sometimes slightly grooved above, without a median rib, mostly with swollen lateral ribs.
Inflorescences axillary, pseudoterminal, or terminal. Flowers: sepals variably connate, 12
mm long; petals absent; disc glabrous; stamens (6)8; ovary 2-locular. Fruits ellipsoid,
3.253.75 x2.5 cm, glabrous; appendages more-or-less dense, strap-shaped to filiform,
curved, to c. 15 mm long, base bulbous to triangular, confluent or not; fruit wall hard,
coriaceous, to 3 mm thick.
Distribution. Sumatra, Peninsular Malaysia, Borneo. Uncommon in Sabah, more common in
Sarawak; also in Brunei.
Ecology. Primary mixed dipterocarp forest on slopes and ridges, on sandy and loamy soils
mainly below 500 m, exceptionally to 975 m. Flowering in August; fruiting in December to
February.
Taxonomy. Compared to most specimens from Peninsular Malaysia and Sumatra, the
Bornean material is less hairy, the margin of the petiolules is not swollen, the apex of the
leaflets is long, slender acute-acuminate, and the fruit appendages are somewhat longer. But
these differences are not sharp, they do not hold for all specimens, and so a subdivision into
infra-specific taxa seems unwarranted.
15. Nephelium uncinatum Radlk. ex Leenh.
Fig. 18D.
(Latin, uncinatus = hook-like; the fruit appendages)
l.c. (1986) 421, l.c. (1994) 692; Yap l.c. 455. Type: Elmer 21708, Borneo, Sabah, Tawau (holotype M; isotypes BO, L, NY,
SING, U).
Tree to 25(40?) m tall, 45 cm in diameter; buttresses to 1.5 m high. Leaves 1-foliolate or

each with up to 37(18) pairs of leaflets; petioles 39 cm long, 11.5 mm thick, terete;
rachis densely minute-hairy, rarely glabrescent. Leaflets parchment-like, puberulous in the
basal part of the midrib to subglabrous above, sparsely hairy on the midrib and lateral veins
beneath, in between lateral veins minutely sericeous, domatia present; narrowly elliptic to
351
obovate, sometimes slightly falcate, 4.7511 x1.53.5 cm; base acute, decurrent, margin
slightly curved, apex tapering to fairly abruptly acuminate, acumen short to long, broad,
obtuse to acute; midrib raised above; lateral veins 38 mm apart, nearly patent, slightly raised
above, intermediate lateral veins well-developed, intercostal veins and veinlets clearly
different, coarsely reticulate, slightly raised but rather inconspicuous on both sides; petiolules
(1)24 mm long, broadly shallowly grooved with a strong median rib above. Inflorescences
terminal, pseudoterminal, or axillary in the upper leaves. Flowers: sepals c. 2550% connate,
in male flowers 11.1 mm, in female ones 1.41.5 mm long; petals absent; disc in male
flowers fairly strongly developed, with the lobes protruding between the stamens, in female
flowers less conspicuous, glabrous or sparsely hairy; stamens 5 or 6; ovary 2-locular,
sometimes in the same specimen also 1-locular. Fruits ellipsoid to subglobose, 2.753 x2
2.25 cm, glabrous, sparsely warty; warts thick, tapering into or more abruptly terminated by
filiform, curved appendage of up to 7.5 mm long; fruit wall coriaceous, 12 mm thick.
Distribution. Peninsular Malaysia, Sumatra, Borneo. Common in Sabah, uncommon in
Ecology. Primary and sometimes old secondary forests, mainly on hill slopes and ridges on
well-drained soils, rarely in swamp; mostly on sandy soils, to 330 m. Flowering mainly in
April to June, sometimes August to October and December; fruiting in December, March.
Uses. In Kalimantan, sometimes cultivated for its fruits.
15. PARANEPHELIUM Miq.
(Greek, para = resembling; near to the genus Nephelium)
M. Davids
Fl. Ned. Ind. Suppl. (1861) 509; Merrill, PEB (1929) 176; Radlkofer in Engler, Pflanzenr. 98 (1933) 1321; Masamune, EPB
(1942) 431; Anderson, CLST (1980) 313; Davids, Blumea 29 (1984) 425, FM 1, 11 (1994) 693; Whitmore & Tantra, CLS
(1986) 216; Corner, WSTM 3rd ed. 2 (1988) 686; Yap, TFM 4 (1989) 456; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990)
314.
Trees or sometimes shrubs, monoecious or very rarely dioecious. Indumentum of solitary

simple hairs only. Leaves imparipinnate or paripinnate, each with 16 pairs of leaflets;
pseudostipules absent; petiole and rachis terete, not winged. Leaflets smooth beneath; margin
entire to dentate; venation open, if margin dentate then each second lateral vein ending in a
tooth. Inflorescences borne on leafless twigs, axillary or terminal. Flowers unisexual or very
rarely bisexual, radially symmetrical; sepals (4)5(7), slightly connate to free, all equal to
very unequal, not petaloid, white to dark yellow hairy outside, white sericeous inside, margin
entire, ciliate, glands absent; petals (4)5(7), longer than the sepals, distinctly clawed to
broadly truncate at base, usually glabrous to laxly woolly outside, glabrous inside, blade
variably developed to nearly completely reduced, scale usually larger than the blade,
emarginate to divided into two lobes, orange woolly especially at the upper margin on both
sides, crest absent; disc composed of a flat ring adnate to the torus except for the margin,
thinly fleshy, glabrous, margin with an erect rim to tubular collar up to 1 mm high; stamens
352
58(9), distinctly exserted in male flowers, glabrous, latrorse; ovary sessile, densely
tuberculate, each tubercle bearing a stiff erect hair, elsewhere glabrous to minute-hairy, (1)
3(4)-locular, ovules 1 per locule, style apical, longer than the ovary, laxly to densely shortstrigose, stigma flat or with recurved lobes to 1.5 mm long. Fruits loculicidal capsules,
subglobose, sessile, not winged, smooth, rough, ribbed, or warty to densely spiny, dehiscing
into 34 mostly unequal valves or tearing apart at random; fruit wall thick, fibrous-woody,
hairy inside. Seeds usually 1 per fruit, subglobose to slightly 23-lobed; arillode absent;
white hilar spot covering up to 75% of the seed; membranous remains of the septa and the
undeveloped ovules tightly pressed against the seed.
Distribution. Four species; in SE Asia from Yunnan and Myanmar to Hainan, Vietnam, and
Thailand, Sumatra, Peninsular Malaysia, Borneo, and the Philippines. In Sabah and Sarawak,
2 species.
Ecology. Mainly medium-sized trees in the lower storeys of various kinds of lowland forest.
The seeds are said to be dispersed by monkeys.
Uses. Of slight importance, only the seeds are edible; oil pressed from the seeds was formerly
used for lamp-oil.
Key to Paranephelium species
Leaflet lateral veins straight, abruptly curving near the margin; intercostal veins densely
scalariform (lax if leaflets large and petiolules at least 4 mm diameter)...................1. P. joannis
Leaflet lateral veins curving gradually; intercostal veins laxly reticulate (sometimes laxly
scalariform if leaflets large, but petiolules at most 4 mm diameter)...............2. P. xestophyllum
1. Paranephelium joannis Davids
Fig. 19E.
(Joannes A.G. Davids, father of M. Davids)
l.c. (1984) 434, fig. 2, 3a, l.c. (1994) 695, fig. 64a; Whitmore, Tantra & Sutisna l.c. 314. Type: Endert 3460, Central E
Borneo, W Kutei, near Petah (holotype L).
Shrub or tree to 24 m tall, 1040(60) cm diameter; buttresses to 1.5 m high or with stiltroots. Leaves each with 23 pairs of leaflets, sparsely tomentose, glabrescent; petioles 825
cm long, 37 mm thick. Leaflets coriaceous, tomentose on midrib above and on midrib,
lateral veins, intercostal veins and veinlets beneath; elliptic to obovate, 1050 x422 cm; base
symmetric or asymmetric, acute, margin entire, apex emarginate, acute or cuspidate; midrib
more or less raised in a groove above; lateral veins straight, abruptly curving near margin,
sunken to slightly raised, intercostal veins densely to laxly scalariform; petiolules 722 mm
long, 18 mm thick. Inflorescences axillary or terminal, stout and often clustered, 2038 cm
long, densely yellow-tomentose or strigose. Flowers sweetly fragrant, white; sepals 5(6),
connate at base, deltoid to ovate, 0.22.3 x0.61.8 mm, acute to mucronate; petals 5, blade
small to absent (but mind the scale!), narrowly spathulate, 01.7 x01 mm, glabrous to pilose
outside, scale lobed, 1.52.1 x 0.92 mm; disc to 1 mm high, 2.53.5 mm diameter; stamens
353
Fig. 19. Paranephelium xestophyllum (AD); P. joannis (E). A, flowering leafy twig; BC, fruits; D, seed; E, fruit. (A from
Whitmore & Kade 3311, B from SAN 79936, C from Kostermans 13238, D from Boerlage, s.n., E from Kostermans 12670.)
354
78, filaments 24.5 mm long; ovary 3-locular. Fruits 2.54 x3.54.5 cm, yellowish or
brownish or red to black, laxly to densely spiny, glabrous to densely short-strigose.
Vernacular name. Sarawakkiah (Iban).
Distribution. Endemic to Borneo; uncommon in Sabah and Sarawak; also in Brunei and
Kalimantan.
Ecology. Primary forest, often on riverbanks, sometimes on slopes; on clay or sandy soil,
sandstone or limestone; from sea level to 450 m. Flowering in MarchApril and August
Sepember, November; fruiting in JulyNovember and JanuaryFebruary.
Uses. The seeds are edible when cooked.
2. Paranephelium xestophyllum Miq.
Fig. 19AD.
(Greek, xestos = smooth, phullon = leaves)
l.c. (1861) 198, 509; Merrill, Enum. Philipp. Fl. Pl. 2 (1923) 514; Radlkofer in Engler l.c. (1933) 1324; Davids l.c. (1984)
437, fig. 1, 3bi, l.c. (1994) 696, fig. 646d, 65; Yap l.c. 457; Whitmore & Tantra l.c. 216; Whitmore, Tantra & Sutisna l.c.
314. Lectotype (Davids, 1984): Teijsmann HB 4218, Sumatra, Lampong Province, G. Batin (L; isolectotype K). Synonym:
Paranephelium nitidum King, J. As. Soc. Beng. 65, 3 (1896) 450, Merrill l.c. (1929) 176, Masamune l.c. 431, Anderson l.c.
313.
Shrub or tree to 20(40) m tall, 1045(75) cm diameter, often with stilt-roots to 60 cm, or
with buttresses to 70 cm high. Leaves each with 16 pairs of leaflets; rachis more or less
densely minute-hairy; petioles 0.817 cm long, 14 mm thick. Leaflets thick-chartaceous;
elliptic (or in the lowermost pair ovate), 342 x1.114 cm; base symmetric (or strongly
asymmetric in the lowermost pair), narrowly cuneate to obtuse, attenuate or not, margin
entire, more or less undulate, apex acute to rounded, acuminate or not; midrib more or less
raised in a furrow to slightly prominent above, minute-hairy on both sides (especially at
base), lateral veins gradually curving, sunken or flat to slightly raised in a groove and hairy
beneath, intercostal veins more or less laxly reticulate or scalariform; petiolules 120 mm
long, 0.84 mm thick. Inflorescences usually borne on leafless twigs, occasionally axillary or
terminal, delicate and clustered, 230 cm long, laxly to densely velutinous and hispid.
Flowers fragrant, white to greenish white to yellowish; sepals (4)5(6), free or connate at
base, equal to very unequal, narrowly triangular to broadly ovate or linear, acute to
acuminate, 2-tipped, 0.82 x0.51.8 mm; petals (4)5(7), 12.5 x0.72.7 mm, glabrous to
woolly outside, scale emarginate to divided into 2 lobes, 1.12.2 x1.23 mm; disc 0.71.5
mm high, 12.8 mm diameter, at base with a few hair-tufts alternating with petals; stamens
58 (or 9), filaments to 4 mm long; ovary (2)3(4)-locular. Fruits to 7 cm diameter, more or
less 23-lobed or globose, brown, yellow or grey, smooth, laxly to densely more-or-less
irregularly gibbose, warty or spiny, glabrous.
Vernacular names.
Kinabatangan).
Sabahambuakat,
ampungot,
bambakat,
tombuakat
(Dusun
Distribution. China (Yunnan), Myanmar, Thailand, Indo-China, Peninsular Malaysia,

Sumatra, Borneo, and the Philippines. Relatively common in Sabah and Sarawak; also in
355
Ecology. Understorey trees of various kinds of forest, often along rivers, on hill-slopes, rarely
in seasonal swamps and on well-drained ridges or summits, on clay, podsol, sand over
igneous rock, sandstone or limestone; most typically in mixed dipterocarp forest on leached
clay-rich soils; from sea level to 1100 m. Flowering mainly in FebruaryJuly, fruiting in
MaySeptember.
Uses. Good firewood but rarely used for timber. Seeds are edible when baked or boiled;
lamp-oil can be obtained from the pressed seeds.
16. POMETIA J.R. Forst. & G. Forst.

(P. Pomet, 16581699, a French writer)
kasai (Malay)
Based on the work by the late
M. Jacobs
Char. Gen. Pl. (1775) 55, t. 55; King, J. As. Soc. Beng. 65, 3 (1896) 440; Merrill, EB (1921) 360; Ridley, FMP 1 (1922) 503;
Radlkofer in Engler, Pflanzenr. 98 (1932) 924; Masamune, EPB (1942) 431; Jacobs, Reinwardtia 6 (1962) 109, FM 1, 11
(1994) 698; Backer & Bakhuizen f., FJ 2 (1965) 138; Anderson, CLTS (1980) 313; Whitmore & Tantra, CLS (1986) 216;
Corner, WSTM 3rd ed. 2 (1988) 687; Yap, TFM 4 (1989) 457; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 314.
Synonyms: Irina Blume, Bijdr. Fl. Ned. Ind. (1825) 229.
Medium-sized to large monoecious trees; often buttressed, with a red exudate when cut.
Indumentum mainly of solitary hairs, sometimes mixed with some small tufts of hairs. Leaves
spirally arranged, paripinnate; basal leaflets stipule-like, often strongly reduced and
caducous; petioles pulvinate; young leaves crimson, very conspicuous; leaves at the base of
inflorescences often reduced to pseudostipules. Leaflets opposite to alternate, lower ones
always smaller; lower surface often with large orbicular glands at least on both sides near
the base, or in the axils of some lateral veins, or sometimes scattered all over the surface, or
exceptionally on the marginal incisions; petiolules broadly attached, nearly always with 2
lateral grooves, or with a broad flat groove, or narrowly winged above. Inflorescences
terminal and sometimes in the axils of upper leaves, thyrsoid; branches mostly long and
racemose; cymules short, of about the same length, patent or nearly so, once-dichasial,
branches bostrycoid (ringlet-like), condensed, axes usually partly connate, distal cymules
reduced to 2 seemingly collateral, bracteate flowers; uppermost flowers solitary; bracts
narrowly triangular to filiform; bracteoles absent; pedicels terete, slender, articulate,
lengthened and swollen in fruit. Flowers unisexual, radially symmetrical; sepals 5, slightly to
more than halfway connate, valvate in bud, 2 outer ones usually slightly smaller, not
petaloid, entire, persistent in fruit; petals 5 (rarely absent), much shorter to distinctly longer
than the calyx, not or hardly clawed, nearly entire, without appendages; disc annular,
pulvinate, not lobed, more or less wavy; stamens 5 (6), in male flowers long-exserted,
filaments filifrom, hairy mainly in the lower half or glabrous, anthers densely minutely
papillose; ovary sessile, cordate, 2(3)-locular, style about as long as or longer than the
ovary, ovules 1 per locule. Fruits sessile, mostly only 1 locule developed, indehiscent,
smooth, glabrous, red to black
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when ripe; exocarp hard, rather thin; mesocarp rather thick and very juicy, white,
semitransparent, tasting sweet, in the dried fruit irregularly split into two fibrous or corky
layers, one inside the pericarp, the other covering the seed. Seeds oblique ovoid, red-brown,
completely enveloped by a thin fleshy arillode, hilum orbicular, c. 5 mm diameter.
Distribution. Two species; Sri Lanka, Andaman and Nicobar Is., Indo-China, Taiwan,
throughout Malesia and the Pacific to Fiji, Samoa, and Tonga.
Ecology. Lower storey and canopy trees of tropical rain forest, primary as well as secondary,
at low to medium altitudes. Dispersal probably mainly by fruit-bats and by birds (Lane-Poole,
For. Res., 1925, 109; Sody, Indon. J. Nat. Sc. 111 (1955) 195), possibly also to some extent
by water as at least some forms of P. pinnata are common along river banks and as the fruits
are buoyant for a few days (Guppy, Observ. Natur. Pacific 2 (1906) 532). In both species,
witches broom is a common and conspicuous pathological feature; they represent mostly
repeatedly dissected leaves, but sometimes also (parts of) inflorescences; the origin is
unknown.
Taxonomy. Pometia seems to be closely allied to Dimocarpus with which it shares
characters such as the orbicular glands on the lower side of the leaflets and, when the leaflets
are incised, the typical venation. They also have similarities in floral structure, the free
arillode, and the tendency to develop pseudostipules.
Key to Pometia species
Leaflets acicular to coarsely dentate; lateral veins sunken above, venation hardly or not
prominent................................................................................................................1. P. pinnata
Leaflets entire; lateral veins slightly raised above, venation prominent on both sides................
..................................................................................................................................2. P. ridleyi
1. Pometia pinnata J.R. Forst. & G. Forst.
Fig. 20.
(Latin, pinnatus = with 2 rows of leaflets; the leaf)
l.c. 55, t. 55; King l.c. 441; Merrill l.c. (1921) 375; Ridley l.c. (1922) 504; Radlkofer in Engler l.c. (1932) 929; Masamune
l.c. 431; Desch, Mal. For. Rec. 15 (1954) 534, t. 107, f. 2; Brown, FTSB (1955) 318; Jacobs l.c. (1962) 120, l.c. (1994) 701;
Backer & Bakhuizen f. l.c. 139; Burgess, TBS (1966) 443, f. 52; Anderson l.c. 313; Whitmore & Tantra l.c. 216; Corner l.c.
688; Yap l.c. 457; Whitmore, Tantra & Sutisna l.c. 314. Type: Forster, s.n., New Hebrides, Namoka (holotype BM; isotype
W). Synonyms: Irina tomentosa Blume l.c. (1825) 230; Pometia tomentosa (Blume) Teijsm. & Binnen., Cat. Hort. Bogor.
(1866) 214, Radlkofer in Engler l.c. (1932) 934, Masamune l.c. 413; Irina alnifolia Blume, Rumphia 3 (1847) 117; Pometia
alnifolia (Blume) King l.c. 442, Ridley l.c. (1922) 504, Radlkofer in Engler l.c. (1932) 928, Masamune l.c. 431, Corner l.c.
688; Nephelium acuminatum Hook.f., Trans. Linn. Soc. 23 (1860) 164; Pometia acuminata (Hook. f.) Radlk., Sapind. Holl.Ind (1879) 9, Merrill l.c. (1921) 175, Radlkofer in Engler l.c. (1932) 933, Masamune l.c. 431, Anderson l.c. 313.
Tree to 50 m tall, 1.4 m diameter; buttresses to 5 m high, spreading to 3 m wide, to 15 cm

thick. Young parts very early to late glabrescent. Leaves to more than 1 m long, each with 4
13 pairs of leaflets; petiole and rachis glabrous to densely hairy; pseudostipules elliptic to
357
Fig. 20. Pometia pinnata. A, flowering leafy twig; B, detail of gland on lower surface of leaflet; C, fruit. (A from Hoogland
5048, B from Kings coll., s.n., C from Clemens 5329.)
358
ovate, 0.43x0.255 cm, straight to strongly falcate. Leaflets parchment-like to coriaceous,

glabrous or variably hairy, variably glandular; ovate to obovate, slightly to distinctly falcate,
632 x213 cm; margin acicular or coarsely dentate, apex acute to abruptly or gradually
acuminate, acumen to 2 cm long, broad to slender, mostly mucronate, lateral veins sunken
above, 1.52.5 cm apart, every 2nd vein ending in or protruding from a marginal tooth, those
in between gradually curving towards the margin and becoming feebler, intercostal veins
hardly or not prominent; petiolules 1.54 mm long, Inflorescences erect to drooping, 1570
cm long, mostly hairy; pedicels 14 mm long, slender to filiform, articulate in the lower one
fifth; bracts usually absent apart from those of the primary flowers of the cymules. Flowers:
calyx 12.5 mm diameter, sepals variably connate, lobes 0.51.5 x0.31.2 mm, variably hairy
outside, hairy or glabrous inside; petals shorter or longer than calyx, 0.41.3 x0.31.6 mm,
hairy or sometimes glabrous outside; stamen filaments 26 mm long, glabrous to completely
hairy, anthers 0.51 mm long; ovary 0.81.2 x2.5 mm, style 25.5 mm long. Fruits 1.5
5 x13 cm; fruit wall in the lower part c. 1 mm, in the upper part to at least 7 mm thick.
Seeds to 2.5 x1.5 cm.
Vernacular names. Sabahdatanut (Dusun), dipangah (Dusun Banggi). Sarawak enselan
(Iban Selayar), kasai, kasai daun besar (Malay), merabong (Malay, Pusa), rapanah, selan
(Melanau), silak (Iban).
Distribution. As for the genus.
Ecology. Primary and secondary forests; well-drained land, swamps, or temporarily
inundated habitats; slopes, ridges and plains, in some regions especially common on
riverbanks; on various soils; from sea level to 1700 m. Flowering and fruiting throughout the
year, mainly seasonally.
Uses. Throughout its area, the wood is used for several purposes, the fruits are eaten, and
according to a few reports a decoction of the bark can be used medicinally. In Sarawak, the
Selayar Iban use this tree for curing chickenpox: the patient is bathed in water in which small
pieces of or powder from the bark is boiled (Heyne, Nutt. Pl. Ned.-Indi ed. 2 (1927) 999;
Burkill, EPMP (1935) 1797).
2. Pometia ridleyi King ex Radlk.
(H.N. Ridley, 18551956, botanist and former Director of Singapore Botanic Gardens)
In Engler l.c. (1932) 927; Jacobs, l.c. (1962) 119, l.c. (1994) 703; Whitmore & Tantra l.c. 216; Corner l.c. 688; Yap l.c. 458;
Whitmore , Tantra & Sutisna l.c. 314. Type: Goodenough 1099, Malaya (holotype CAL; isotypes K, M, SING).
Tree to 40 m tall, 1.5 m diameter; buttresses steep, to 3 m high. Young parts very early
glabrescent. Leaves each with 68 pairs of leaflets; petiole and rachis subglabrous;
pseudostipules more or less elliptic, 0.61 x0.20.5 cm, strongly falcate. Leaflets chartaceous,
glabrous except for a few scattered hairs on the midrib beneath, without gland or
exceptionally with a gland in the axils of 1 or 2 lateral veins; more or less elliptic, 7.5
15 x2.85.5 cm, middle ones hardly to distinctly falcate, margin entire, apex gradually to
rather abruptly acuminate, acumen acute, to 1 cm long; lateral veins 0.51 cm apart, all alike,
gradually bending towards the margin and more or less distinctly looped and joined, slightly
359
raised above, intercostal veins prominent on both surfaces; petiolules 1.52 mm long.
Inflorescences erect, 1725 cm long, subglabrous; pedicels c. 2 mm long, rather slender,
articulate up to one fourth to one third above the base; bracts present under all flowers.
Flowers: calyx c. 2.5 mm diameter, sepals connate for 1520%, lobes c. 1.5 x0.75 mm,
glabrous on both sides; petals shorter than the calyx, 0.65 x0.81 mm, glabrous outside;
stamen filaments c. 2.2 mm long, very sparsely hairy at base, anthers c. 0.75 mm long; style
c. 3.5 mm long. Fruits c. 2 x 2 cm.
Distribution. Sumatra, Peninsular Malaysia, Borneo. In Sabah, uncommon.
Ecology. Primary lowland mixed dipterocarp; outside Borneo also in bamboo forests; to 200
m. Flowering in July, September.
17. RHYSOTOECHIA Radlk.
(Greek, rhusos = wrinkled, toichos = wall; the inner side of the fruit wall)
B. Etman
Sapind. Holl.-Ind. (1879) 61, 62, in Engler, Pflanzenr. 98 (1933) 1209; Merrill, EB (1921) 361; Masamune, EPB (1942) 432;
Etman, Blumea 39 (1994) 41, FM 1,11 (1994) 704.
Small to medium-sized trees or shrubs. Indumentum, if present, mostly of simple, appressed

hairs. Leaves imparipinnate; petiole and rachis terete, slightly winged or not; petioles
pulvinate. Leaflets opposite to alternate; base symmetric to slightly asymmetric, margin
entire; lower surface with domed cells, domatia absent; petiolules pulvinate, sometimes
absent. Inflorescences axillary, subterminal or borne on leafless twigs, paniculate or
thyrsoid; bracts and bracteoles usually not persistent in fruit. Flowers radially symmetrical,
seemingly bisexual; sepals 5, subpersistent in fruit, outer 2 slightly to distinctly smaller than
inner 3, inner ones with a petaloid margin; petals 5, distinctly clawed, scales absent to welldeveloped, crest absent; disc entire, slightly lobed, glabrous; stamens (7)8, filaments
especially towards the base pilose or velutinous, anthers often with a few hairs; ovary 2(3)
locular, sericeous, ovule one per locule, style usually glabrous, elongating in fruit. Fruits
subcordate to reniform, with one to all lobes developing, loculicidal; fruit wall rugose to
ribbed and often laxly hairy outside, densely papillose inside; stalk absent to very distinct.
Seeds obovoid to globose; arillode (or sarcotesta) cup-shaped, covering the seed except at
the apex; hilum round; pseudohilum round to reniform; cotyledons secondarily collateral,
about equal in size.
Distribution. Fourteen species; Borneo, the Philippines, Sulawesi, Maluku, New Guinea, and
Australia. In Sabah, 1 species, not yet reported from Sarawak.
Ecology. In lowland rain forest (often coastal) to montane forest.
Rhysotoechia koordersii Radlk.
(S.H. Koorders, 18631919, Forest Officer of the Dutch East Indies at Bogor)
Fig. 21.
In Engler & Prantl, Nat. Pfl. Fam. 3 (1907) 206, in Engler l.c. (1933) 1213; Etman, Blumea 39 (1994) 63, FM 1, 11 (1994)
708. Type: Koorders 18844, Celebes, Menado (holotype M; isotypes BO, K, L).
360
Fig. 21. Rhysotoechia koordersii. A, fruiting leafy twig; B, fruit. (From SAN 92985.)
361
Tree to 20 m tall, 25 cm diameter. Bark smooth, dark green to blackish brown, inner bark
yellowish to ochre, thin. Leaves each with 14 pairs of leaflets; petioles 211 cm long,
slightly angular to terete, sometimes slightly winged below the lowermost pair of leaflets,
slightly ribbed, glabrous; rachis 2.520 cm long, angular, slightly winged below pairs of
leaflets, ribbed, glabrous. Leaflets usually subopposite, sometimes opposite, coriaceous;
glabrous on both surfaces; obovate to elliptic, 623 x111 cm; base sometimes very slightly
asymmetric but never distinctly so, acute to attenuate, margin not to slightly recurved, apex
obtuse to abruptly narrowing and distinctly acuminate; midrib prominent, smooth to angular,
lateral veins 0.53 cm apart, upper ones distinctly looped, intermediate lateral veins curved
towards the base, intercostal veins laxly reticulate, slightly raised above; petiolules usually of
pulvinus only, to 10 mm long, glabrous. Inflorescences axillary, thyrsoid, branching at the
base, to 19 cm long, sparsely puberulous; cymules 1-flowered; bracts and bracteoles 0.51
mm long; pedicels 49 mm long. Flowers c. 7 mm diameter; sepals glabrous on both sides,
ciliate, outer ones broadly ovate, 2.53x2.5 mm, inner ones orbicular, 33.5 x3 mm; petals
broadly obovate, c. 2.5 x2 mm, pilose outside, sparsely so inside, margins coarsely lobed,
slightly thickened towards the base, densely pilose, apex rounded, claw c. 0.8 mm high,
pilose on both sides, scales absent; disc glabrous; stamens 8, filaments c. 3 mm long,
velutinous towards the base, anthers c. 0.5 mm long, puberulous; ovary c. 1 mm high, densely
sericeous , style c. 2 mm high, sparsely puberulous. Fruits obcordate, attenuate at base, 1.5
1.7 x1.31.5 cm, rugose and very sparsely puberulous outside, smooth inside; stalk slender,
57 mm long; lobes 13, well-developed, 0.81x 0.60.7 cm; style c. 0.5 mm long.
Distribution. Borneo (Sabah: Kota Belud and Beluran districts), Sulawesi. Also known from
Kalimantan.
Ecology. In lowland and hill forests, from sea level to 700 m. Flowering in April; fruiting in
May.
18. TRIGONACHRAS Radlk.
(Greek, trigonos = triangular, achras = pear-shaped; the fruit)
P.W. Leenhouts
Sapind. Holl.-Ind. (1879) 46, in Engler, Pflanzenr. 98 (1933) 1243; King, J. As. Soc. Beng. 65, 3 (1896) 445; Ridley, FMP 1
(1922) 507; Corner, WSTM 3rd ed. 2 (1988) 689; Leenhouts, Blumea 33 (1988) 204, FM 1, 11 (1994) 734; Yap, TFM 4
(1989) 459.
Trees, monoecious. Indumentum of solitary simple hairs only. Leaves paripinnate, each with
19 pairs of leaflets; pseudostipules absent; neither petiole nor rachis winged. Leaflets
opposite to alternate, parchment-like; margin entire; upper surface without wax, below
sometimes with brownish glands in the axils of lateral veins. Inflorescences axillary or
pseudoterminal, thyrsoid, hairy, more or less glabrescent; cymes lax, 1few-flowered.
Flowers radially symmetrical, unisexual; sepals 5, free, slightly imbricate, all about equal or
sometimes the outer two slightly smaller, sometimes thinner towards the entire margin,
without glands; petals 5, longer than the calyx, distinctly clawed, claw densely ciliate, blade
entire, scales 1(2), erect, nearly as long as the blade, densely woolly, not crested; disc
362
uninterrupted, annular, rather thick, glabrous; stamens (7)8(9), in male flowers long
exserted, filaments filiform, broadened toward the base, anthers obovoid, latero-introrse to
latrorse; ovary sessile, densely hairy, 3-locular, ovule 1 per locule, septa complete, style
apical, about as long as the ovary, stigma grooved or slightly lobed. Fruits distinctly stalked,
loculicidal capsules, dehiscing into 3 equal valves, not winged, smooth, minute-hairy or
glabrous outside, usually densely woolly inside; fruit wall hard-fleshy. Seeds ellipsoid; testa
smooth, shiny, black to brown; hilum nearly basal, elliptic, fairly big, covering less than one
third of the seed; arillode absent.
Distribution. Eight species; Peninsular Malaysia, Sumatra, Borneo, the Philippines,
Sulawesi, and New Guinea. In Sabah and Sarawak, three species, including two undescribed
ones.
Ecology. Lowland primary as well as secondary forests, often in marshy habitats.
Key to Trigonachras species
1. Mature fruits hairy...............................................................................................1. T. acuta
Mature fruits glabrous.........................................................................................................2
2. Leaves each with 35 pairs of leaflets; leaflets elliptic, apex acute.......................................
...........................................................................................................2. Trigonachras sp. A
Leaves each with 7 pairs of leaflets; leaflets ovate, apex tapering into an acute
acumen.......
...........................................................................................................3. Trigonachras sp. B
1. Trigonachras acuta (Hiern) Radlk.
Fig. 22.
(Latin, acutus = pointed; the leaflet apex)
l.c. (1879) 46, in Engler l.c. (1933) 1245; King l.c. 445; Ridley l.c. (1922) 507; Corner, l.c. 689; Leenhouts l.c. (1988) 206,
l.c. (1994) 735, fig. 79; Yap l.c. 459. Basionym: Cupania acuta Hiern in Hooker, f., Fl. Br. Ind. 1 (1875) 677. Types:
Maingay KD 445, Malay Peninsula (holotype K; isotypes CGE, L).
Tree to 25 m tall, 50 cm diameter, with buttresses. Leaves each with 38 pairs of leaflets;
petioles 1.56 cm long, 12 mm thick. Leaflets opposite to alternate, slightly falcate, thin to
rather stiff parchment-like; glabrous or with some appressed hairs on the lower side of the
midrib, with small glands in the axils of lateral veins beneath; ovate to elliptic, 4.512 x1.54
cm; base symmetric to asymmetric, obtuse, acute or rounded, slightly attenuate, apex usually
tapering into an acute acumen, or acute to obtuse; lateral veins 0.51.5 cm apart, spreading,
strongly curved, ending free, intercostal veins fairly coarsely reticulate, equally raised on
both sides; petiolules subterete to grooved above, 36 mm long. Inflorescences
pseudoterminal and terminal, 1020 cm long, without or with a few long branches, usually
few-flowered, fulvous-puberulous. Flowers: sepals broadly ovate, 1.31.75 x1.251.75 mm,
fairly densely short-sericeous on both sides to rather sparsely so inside, margin ciliate; petals
23 x1.62.5 mm, claw with a few hairs outside, orbicular, glabrous, scales 2, slightly more
than half as long as the petal; stamens 7 or 8, filaments to 5.5 mm long, fairly densely
patently long-hairy except for the upper part, anthers c. 1 mm long, glabrous; ovary 3(4)locular, shortly brown-hairy,
363
Fig. 22. Trigonachras acuta. A, flowering leafy twig; B, fruit. (From SAN 40845.)
364
slightly 3-angular, tapering into the at least equally long style, stigma grooved. Fruits moreor-less triangular-clavate, apiculate, 4.55 x2.253 cm, brown-tomentulose, red when fresh;
fruit wall c. 1 mm thick, hard, thick and fleshy when fresh; endocarp rugose, sparsely to
densely woolly. Seeds c. 1.5 x1 cm, black and shiny; hilum basal, transversely elliptic, c. 7x5
mm.
Distribution. Sumatra, Peninsular Malaysia, and Borneo (Sabah, uncommon).
Ecology. In primary and secondary forests, often in swamps, along riverbanks, along road
sides, on slopes and ridges, often on sandy soils; from sea level to 225 m. Flowering in April
June; fruiting in February, JulySeptember, November.
2. Trigonachras sp. A
Leenhouts l.c. (1988) 207, l.c. (1994) 739.
Tree to 30 m tall, 35 cm diameter. Leaves each with 35 pairs of leaflets; petioles 3.54 cm
long, 1.52 mm thick. Leaflets opposite, stiff parchment-like, glabrous, without glands;
elliptic, 510 x1.754 cm, slightly falcate; base hardly to distinctly asymmetric, acute,
strongly attenuate, apex acute; lateral veins 11.5 cm apart, spreading, straight, ending free,
intercostal veins coarsely reticulate, slightly raised on both sides. Inflorescences
pseudoterminal, 1215 cm long, in fruit sparsely branched, fairly densely puberulous.
Flowers (only known from remains under the fruit): sepals sparsely hairy outside, densely
hairy inside. Fruits (probably young) triangular-obovoid, c. 2.75 x1.5 cm, narrowed toward
the base, apiculate, glabrous; fruit wall somewhat fleshy, at least 1 mm thick; endocarp
densely woolly.
Distribution. Borneo: Sabah, 1 collection, SAN 75360, Sandakan Distr. (L, SAN).
Ecology. Low hill at c. 180 m. Young fruits collected in July.
3. Trigonachras sp. B
Leenhouts l.c. (1988) 209, l.c. (1994) 739.
Tree c. 25 m tall, c. 65 cm diameter; with buttresses. Leaves each with 7 pairs of leaflets;
petioles c. 5 cm long, 2 mm thick. Leaflets subopposite, membranous (young?), glabrous
above, tomentulose on the midrib beneath, small glands present in the axils of lower lateral
veins; ovate, 7.511.5 x34 cm, slightly falcate; base asymmetric, apex tapering into an
acute acumen; lateral veins c. 1 cm apart, straight to slightly curved, ending free, intercostal
veins densely reticulate above, coarsely reticulate beneath, slightly raised on both sides;
petiolules slightly grooved above, 35 mm long. Fruits triangular ellipsoid, c. 3 x2 cm,
stalked, apiculate, rugose, glabrous.
Distribution. Borneo: Sarawak, one collection, S. 28037 (L, SAR ).
Ecology. On limestone slope, grey sandy soil; at c. 250 m. Fruiting noted in September.
Taxonomy. Possibly related to another unnamed species from Mindanao, the Philippines.
365
19. TRISTIROPSIS Radlk.
(Greek, opsis = resembling; closely allied to Tristira)
P.W. Leenhouts
In Durand, Ind. Gen. Phan. (1888) 76; in Engler, Pflanzenr. 98 (1932) 861; Anderson, CLST (1980) 314; Leenhouts, FM 1,
11 (1994) 742; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 315.
Trees, monoecious. Indumentum of solitary, simple hairs only. Leaves bipinnate or the upper
ones at least forked, with several leaflets per branch; pseudostipules absent; petiole and
rachis not winged; petioles pulvinate, more or less flattened above. Leaflets alternate to
subopposite, chartaceous, base oblique, margin entire. Inflorescences thyrses, axillary
towards the end of the branches; peduncles pulvinate, branches few, spreading, bearing few
to many, short- to long-stalked, few- to many-flowered (in the upper parts reduced to a
solitary flower) cymes; pedicels articulate at base. Flowers: sepals 5, free, imbricate,
concave, the two outer ones slightly or hardly smaller, fleshy, margin petaloid or not, ciliate,
densely appressed short-tomentose outside; petals absent or 5, slightly shorter than the
sepals, clawed or not, sericeous towards the base outside, margin ciliate, scale solitary,
somewhat bifid, hairy, not crested; disc uninterrupted; stamens 8(13), hardly or not
exserted, filaments filiform, anthers latrorse, ovate-oblong, basifixed, cleft at base for a third
at most, apex of connective usually rounded-apiculate by a gland which is dark when dry;
ovary densely appressed fulvous-hairy, conical-ovoid, faintly 3(5)-angular, 3(5)-locular,
tapering into a short style with a 3-grooved stigma; ovules 1 per locule. Fruits
subdrupaceous, (1)3(5)-locular, shortly attenuate to short-stalked at base, shortly apiculate
at apex, densely minutely tomentose, subglabrescent, variably hairy inside; fruit wall thin;
epicarp fibrous; endocarp woody. Seeds brown; hilum basal, orbicular; arillode absent.
Distribution. Three species; from the Philippines and Borneo eastwards to the Marianas,
Christmas Is., Queensland, and the Solomon Is. (Leenhouts & van Balgooy, Blumea Suppl. 5
(1966) 198, map 109). One species in Sabah and Sarawak.
Ecology. Canopy trees of primary and secondary forests at low to medium altitudes.
Tristiropsis ferruginea Leenh.
Fig. 23.
(Latin, ferrugineus = rusty brown; the indumentum)

Blumea 13 (1966) 395, l.c. (1994) 746, fig. 82ac. Type: Carr SF 27276, Borneo (holotype SING).
Tree to 30 m tall, 40 cm diameter. Leaves: petioles 610 cm long; ultimate parts of rachis
flattened to keeled above. Leaflets alternate, bearded in the axils of lateral veins, glabrous or
thinly tomentose on the midrib and on the lateral veins beneath; ovate, 510 1.83 cm; base
acute to obtuse, decurrent, apex attenuate-acuminate, acumen acute; midrib slender and
slightly raised above, lateral veins 0.51 cm apart, curved to straight, only the upper ones
looped and joined near the margin, slightly raised on both sides, but more strongly so
beneath, intermediate lateral veins more or less well-developed, parallel to the lateral veins;
petiolules to 2 mm long, keeled above, pulvinate. Inflorescence a thyrse, to 17 cm long,
densely tomentulose; peduncles 25 cm long; cymes with 12-mm-long stalks; pedicels 12
mm long.
366
Fig. 23. Tristiropsis ferruginea. A, fruiting leafy twig; B, petal from inside; C, ovary. (A from Kostermans 21330, BC from
Anderson 4626.)
367
Flowers: sepals dark purple, hardly or not persistent in fruit, outer 2 broadly ovate, inner
ones broadly obovate, c. 2.5 x22.2 mm, rather densely appressed hairy inside, margin entire,
not petaloid; petal blade transversely half-elliptic, c. 1 x1.5 mm, margin at below the insertion
of the scale densely woolly, upper part crenulate and densely ciliate, woolly inside, claw c.
1.25 mm long, densely woolly, scale about half the length of blade, slightly bilobed,
completely woolly; disc 5-lobed, the lobes in the centre deeply hollowed, hence each lobe
more or less annular, the parts towards the centre of the flower densely hairy, elsewhere
glabrous; stamens 8, filaments 12 mm long, rather densely woolly in the lower part, anthers
c. 0.8 mm long, glabrous. Fruits subglobose, c. 22 x17 mm, at base contracted into a c. 1mm-long stalk, densely ferrugineous tomentulose, (1)2(3)-locular, densely ferrugineous
velvety inside.
Distribution. Endemic to Borneo. In Sabah and Sarawak uncommon; also in Kalimantan.
Ecology. Primary forest on limestone; up to 500 m. Flowering in May, July; fruiting in
August, October.
20. XEROSPERMUM Blume
(Greek, xeros = dry, spermum = seed; the seed without arillode)
P.W. Leenhouts
Rumphia 3 (1847) 99; King, J. As. Soc. Beng. 65, 3 (1896) 431; Merrill, EB (1921) 359; Ridley, FMP 1 (1922) 496;
Radlkofer in Engler, Pflanzenr. 98 (1932) 936; Masamune, EPB (1942) 432; Backer & Bakhuizen f., FJ 2 (1965) 137;
Anderson, CLTS (1980) 314; Leenhouts, Blumea 28 (1983) 389, FM 1, 11 (1994) 746; Whitmore & Tantra, CLS (1986)
216; Corner, WSTM 3rd ed. 2 (1988) 690; Yap, TFM 4 (1989) 460; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 315.
Medium-sized trees or shrubs, androdioecious. Indumentum of solitary simple hairs only.

Wood reddish brown. Leaves paripinnate, unifoliolate or each with 3 pairs of leaflets;
pseudostipules absent; petioles slightly swollen at base, hollowed above; petiole and rachis
not winged. Leaflets opposite, neither papillose, nor glaucous beneath, usually with a few to
several orbicular flat glands in the axils of lateral veins or glands scattered all over the
surface mainly in the basal half, domatia absent; base somewhat symmetric, margin entire.
Inflorescences usually solitary in the lower leaf axils and tufted in the upper leaf axils; bracts
usually caducous. Flowers radially symmetrical, 4- or 5-merous, effectively unisexual; sepals
free or slightly connate, either all about equal or the outer two slightly smaller, the outer ones
usually with a narrow, the inner ones with a broad membranous margin to nearly completely
membranous; petals about equal to slightly shorter than the sepals, sessile to variably clawed,
without a scale, woolly ciliate; disc uninterrupted or interrupted, not lobed, in female flowers
inconspicuous; stamens (7)8(9), in male flowers hardly to distinctly exserted, anthers
dorsifixed, latrorse, in female flowers with indehiscent anthers bearing functional pollen;
ovary 2(3)-locular, deeply lobed, warty, style apical, columnar, broadened towards the apex,
stigma arched, elliptic with a longitudinal groove, deeply cleft in fruit, ovules 1 per locule,
nearly basally attached. Fruits with 1(2) lobes developed, in the latter case lobes widely
spreading, not winged; lobes ellipsoid to subglobose, indehiscent; fruit wall early glabrescent,
spiny, warty, colliculate or granular outside, smooth or slightly colliculate and glabrous inside;
368
spines broader than high. Seeds ellipsoid to subglobose, completely covered by a thin sarcotesta except for the
basal hilum; inside of testa with a pocket in which the radicle fits; embryo transverse, plumule lateral with a
relatively big radicle pointing downwards.
Distribution. Two species; Bangladesh, Assam, Myanmar, Indo-China, Peninsular Malaysia, Sumatra, Java, and
Borneo.
Ecology. Often common in the middle and lower storeys of lowland and lower montane rain forests. The fruits are
said to be eaten and dispersed by birds, monkeys, and fruit-bats.
Uses. The thin yellow to orange sarcotesta is edible, but is not of economic importance. The opinions on the timber
quality are rather divergent: fire wood, inferior timber, or a good, tough, and durable timber (Heyne, Nutt. PI. ed. 3
(1950) 997; Burkill, EPMP 2 (1966) 2313).
Key to Xerospermum species
Flowers 5-merous. Pedicels in fruit not much swollen, 1.52.5(3.5) mm thick.........................

....................................................................................................................... 1. X. laevigatum
Flowers 4-merous. Pedicels in fruit distinctly swollen, (3.5-)5-7 mm thick.................................
................................................................................................................... 2. X. noronhianum
1. Xerospermum laevigatum Radlk.
Fig. 24A, B.
(Latin, laevigatus= smooth and polished; the leaves)
Sapind Holl.-Ind. (1879) 23, 25; in Engler I.e. (1932) 949; King I.e. 431; Ridley I.e. 497; Burkill, EPMP 2 1935) 2272;
Anderson I.e. 314; Leenhouts I.e. (1983) 391, I.e. (1994) 748; Whitmore & Tantra I.e. 216; Yap I.e. 461. Type: Griffith KD 1006/1,
Burma (holotype K).
Shrub or tree to 36 m tall, 1 m diameter. Leaves: petioles 0.4-7 cm long; petiole and rachis glabrous. Leaflets
coriaceous, glabrous, without or with a few to several glands; elliptic, 4.5-18 x 1.75-10 cm; base acute to
rounded, decurrent, apex rounded to acute or variably acuminate; upper lateral veins more or less distinctly
joined, intermediate lateral veins variable, intercostal veinlets rather finely reticulate, slightly raised on both sides,
sometimes more so beneath than above; petiolules 0.3-1.5 cm long. Inflorescences to 20 cm long if solitary, to 5
cm long if tufted; well-developed axes with scattered, patent and short or erecto-patent and long side-branches;
branches and upper part of main axis bearing several lax and often several-flowered cymes; lower cymes longstalked, consisting of a central flower and two sometimes long and many-flowered monochasial branches; upper
cymes sessile and monochasial, often (if the axis is short) forming a dense cluster of branches and flowers; bracts
deltoid to lanceolate, to 1.5 mm long, sparsely hairy; pedicels 1.5-5 mm long. Flowers 5-merous; sepals free,
about equal and ovate to obovate, or the outer two distinctly smaller and concave, 1.6-2.8 x 1.4-2.5 mm, ciliate to
woolly ciliate, glabrous or nearly so; petals obovate to spathulate, 1-3 x 0.5-1.2 mm, larger in male than in female
flowers, whitish, variably woolly; disc uninterrupted or interrupted, yellow; stamens (7-)8, filaments 2-5 mm
long, woolly in the lower half to all over except for the apex, anthers 0.6-0.9 mm long, glabrous: Fruit lobes
globose to oblong-ellipsoid, 2.5-3.75 x 1.5-2.5 cm, densely spiny, orange to pinkish; fruit wall coriaceous, c. 0.5
mm thick; pedicels only slightly swollen, 1.5-2.5(-3.5) mm thick.
369
Distribution. Myanmar (Mergui Arch.), Peninsular Malaysia, Sumatra, and Borneo. Two
subspecies are recognised.
Key to subspecies
Disc in female flowers nearly always interrupted, in male flowers sometimes interrupted.
Leaflets broad-elliptic, distinctly acuminate........................................................
subsp. acuminatum (Radlk.) Leenh.
I.e. (1983) 393, I.e. (1994) 749. Basionym: Xerospermum acuminatum Radlk. I.e. (1879) 25,
Merrill I.e. 359, Masamune I.e. 432, Anderson I.e. 314 (as X. acuminatissimum). Type: Beccari
PB 3468, W Borneo (holotype FI; isotypes K, P).
Shrub or tree to 20 m tall, 24 cm diameter, with slight buttresses or with stilt-roots.
Leaves: petioles to 7 cm long; rachis to 7 cm long. Leaflets fairly strongly
dorsiventrally curved, acumen usually cuneate and acute, to 3 cm long.
Inflorescences if axillary 2 cm long or longer, glabrous or rarely sparsely hairy.
Flowers: outer sepals distinctly smaller than inner ones, glabrous except the margin,
fleshy or the inner ones partly petaloid; petals to 3 x 1.2 mm.
Endemic to Borneo. In Sabah and Sarawak, relatively common; also in Brunei and
Kalimantan. In peat swamp, kerangas and mixed dipterocarp forests, on sandy humic
yellow soils; to 30 m.
Disc interrupted. Leaflets usually oblong to elliptic, not or slightly acuminate ...
subsp. laevigatum
Leenhouts I.e. (1983) 392, I.e. (1994) 749; Yap I.e. 461. Synonym: Xerospermum unijugum
Radlk., Rec. Bot. Surv. Ind. 3 (1907) 351, in Engler I.e. (1932) 948, Ridley I.e. (1922) 497.
Shrub or tree to 36 m tall, 1 m diameter; buttresses to 2.5 m high, 2.3 m wide, thick,
branching. Leaves: petioles to 4.5 cm long; rachin 2-3 cm long. Leaflets flat to
slightly dorsiventrally curved. Inflorescences to 12 cm long, slightly thin-puberulous,
glabrescent. Flowers: sepals nearly equal, usually thin and petaloid; petals to 2 x 0.8
mm.
Distribution as for the species; common in Sabah and Sarawak; also in Kalimantan.
In primary and sometimes secondary mixed dipterocarp forest, on well-drained land;
to 700 m.
2. Xerospermum noronhianum (Blume) Blume

(Fernando Noronha, Spanish biologist in the 18th century)
Fig. 24C-F.
l.c. (1847) 100; Radlkofer in Engler I.e. (1932) 946; Backer & Bakhuizen/ I.e. 137; Leenhouts, I.e. (1983) 394, fig. 2a-c, e, l.c.
(1994) 750, fig. 84b-e; Whitmore & Tantra l.c. Ill; Yap l.c. 461. Basionym: Euphoria noronhiana Blume, Bijdr. Fl. Ned. Ind.
(1825) 234. Lectotype (Leenhouts, 1983): Blume, s.n. (=L sheet no. 908.272-748), Java (L). Synonyms: Xerospermum
lanceolatum Radlk. l.c. (1879) 7, 25, Merrill l.c. 359, Masamune l.c. 432; X. muricatum Radlk. in Engler & Prantl l.c. (1895)
331, fig. 168, Anderson l.c. 314; X. echinulatum Radlk. l.c. (1907) 350, Anderson l.c. 314; X. intermedium Radlk. l.c. (1907)
348, Anderson l.c. 314.
Tree to 25(-30) m tall, 30(-75) cm diameter, with buttresses. Leaves: petioles 1-7 cm long;
petiole and rachis glabrous or thinly to densely brownish or fulvous-puberulous, glabrescent.
Leaflets parchment-like to coriaceous; glands few to many; more-or-less elliptic, to 50 x30 cm;
base obtuse to acute, apex rounded to variably acuminate, lateral veins ending free except
371
for a few uppermost ones, intermediate lateral veins not conspicuous, intercostal veins and
veinlets finely to laxly reticulate, about equally raised on both sides to smooth or slightly
sunken on the upper side; petiolules 1-12 mm long. Inflorescences male or female; male
inflorescences to 25 cm long if solitary, or much shorter if tufted; female inflorescences tufted;
axes simple or with some short patent branches in the lower part; cymes few to many, sessile or
subsessile, very condensed, few- to several-flowered; distal flowers solitary; bracts deltoid to
lanceolate, to 1.5 mm long, sparsely hairy; pedicels c. 2 mm long. Flowers 4-merous; sepals free
or slightly connate, outer two usually slightly smaller than inner ones, ovate to obovate, l-2(-3) x
1-2.4 mm, glabrous on both sides or at base hairy inside, ciliate; petals obovate to broadly
spathulate, 1-2.8 x 0.5-1.7 mm, short- to long-clawed, blade ovate to transversely elliptic,
variably woolly except the base outside, sparsely hairy to glabrous inside; disc uninterrupted;
stamens 8(-9), filaments 1.5-2.5 mm long, woolly except at base and apex, to woolly or
glabrous in the upper half, anthers 0.3-0.8 mm long, glabrous or with a few hairs (ciliate).
Fruit lobes ellipsoid to subglobose or obovoid, 1.75-5 x 1.25-5 cm, aculeate, tuberculate,
colliculate to granulate, red or dark brown; fruit wall coriaceous, corky or woody, 0.65-2.5 mm
thick; pedicels strongly swollen, (3.5-)5-7 mm thick.
Vernacular names. Sabahresat (Dusun Kinabatantan). SarawakHat, kalas (Iban), tundun
biawak (Malay).
Distribution. As for the genus. Common in Sabah and Sarawak; also in Brunei and
Kalimantan.
Ecology. Primary and secondary rain forests, sometimes in peat swamp and kerangas forests, on
plains, slopes and crests; in well-drained places but also on riverbanks, marshy habitats, and
periodically flooded localities; on different kinds of soil including sandy, sandy-clay, fertile
volcanic loam, peat, subsoil granite, sandstone, and moist limestone; from sea level to 1500 m.
Flowering throughout the year but mainly in January-April and August-December; fruiting
mainly in January-August. The fruits are eaten and dispersed by birds and monkeys.
21. ZOLLINGERIA Kurz, nom. cons.
(H. Zollinger, 1818-1859, Swiss school teacher and plant collector)
F. Adema
J. As. Soc. Beng. 41, 2 (1872) 303; Radlkofer in Engler, Pflanzenr. 98 (1933) 724; Adema, Blumea 37 (1992) 73, FM 1, 11
(1994) 752.
Trees, monoecious. Indumentum of solitary simple hairs; glands absent. Leaves spirally
arranged, paripinnate, each with 18 pairs of leaflets; pseudostipules absent; petiole and rachis
winged. Leaflets opposite to alternate, margin entire. Inflorescences axillary, thyrsoid,
branched. Flowers unisexual, radially or rarely bilaterally symmetrical; sepals 5, free, slightly to
distinctly unequal, imbricate or valvate, not petaloid, hairy or glabrous outside; petals 5 or
occasionally 4, shorter or longer than the sepals, with or without scales or auricles; disc entire or
interrupted, glabrous; stamens 8, slightly exserted in male flowers, filaments hairy, anthers
glabrous; ovary 3-locular at base, higher up 1-locular, ovule 1 per locule, style 3-lined or 3lobed. Fruits 1-locular, 3-winged, indehiscent. Seeds / per fruit, without arillode or
sarcotesta.
372
Distribution. Three or four species in Myanmar, Laos, Thailand and Borneo; one species in
Sabah.
Zollingeria borneensis Adema
Fig. 25.
(of Borneo)
Blumea 37 (1992) 75, I.e. (1994) 754. Type: Elmer 20888, Borneo, Sabah, Tawau (holotype L; isotypes
A, F, U).
Tree c. 60 m tall, clear bole 30 m high, to 70 cm in diameter. Bark whitish, inner bark pale
yellow, c. 0.7 cm thick. Sapwood white. Indumentum ferrugineous tomentulose. Twigs silverygrey. Leaves each with (l-)2-3 pairs of leaflets; petioles 1-6.5 cm long, pulvinate; rachis 1-10.5
cm long; petiole and rachis subterete, finely grooved, patently short-hairy. Leaflets
chartaceous, glabrous; more or less elliptic, 9.5-20 x 2-6 cm; base slightly asymmetric,
broadly cuneate, apex acuminate, acumen rounded; midrib and lateral veins sparsely to densely
short-hairy, midrib prominent above, lateral veins 9-12 per side, 8-28 mm apart; petiolules
reduced to pulvinus, 1-4 mm long, subterete, short-hairy. Inflorescences pseudoterminal, 6-14
cm long; cymes several-flowered; bracts and bracteoles acicular, 0.6-12 x 0.1-0.4 mm, hairy on
both sides; pedicels 1.5-2.5 mm long, articulate near the base, angular, hairy. Flowers yellowish
to green; male ones somewhat smaller than the female ones; sepals triangular to spathulate,
valvate, thinned towards the margin especially at the apex, slightly unequal, 1.7-2.6 x 1.2-1.6
mm, apex rounded to retuse, appressed hairy on both sides; petals cuneate to spathulate, distinctly
clawed, 1.4 - 2.4 x 1.2-1.6 mm, ciliate, claw 0.4-0.7 mm long, hairy on both surfaces, auricles
ciliate; disc entire, annular or saucer-shaped; stamen filaments c. 1.5 mm long, anthers c. 0.4 mm
long, in female flowers respectively 1.2 - 1.6 mm long, 0.5-0.6 mm long; ovary hairy outside,
glabrous inside, style c. 1 mm long, stigma more or less capitate, 3-lobed; pistillode c. 3 x 1.9
mm. Fruits prismatoid, sharply deltoid in cross section, 6-7 x 2.5-5 cm; fruit wall thincoriaceous, striate and densely velutinous outside, glabrous inside. Seeds prismatoid, c. 33 x 14
mm.
Distribution. Endemic to Sabah; uncommon.
Ecology. Primary forest, at c.15 m. Flowering noted in February.
374
SCYPHOSTEGIACEAE
Taman Fajar, Lahad Datu,
Sabah, Malaysia
Hutchinson, Fam. Fl. Pl. 2nd ed. (1964) 326, f. 187a & b; Metcalfe, Reinwardtia 4 (1956) 99; van Steenis,
FM 1, 5 (1957) 297; van Heel, Blumea 15 (1967) 107; Anderson, CLTS (1980) 323.
A monogeneric family with one species endemic to Borneo.

Taxonomy. The systematic position of the family has been controversial. Stapf (FMK (1894)
217) assigned the only genus, Scyphostegia, to the Monimiaceae, but Hutchinson (l.c.) placed
it in a separate family, the Scyphostegiaceae, closely related to Moraceae. van Steenis (l.c.),
Metcalfe (l.c.), and van Heel (l.c.), on the other hand, though in agreement with Hutchinson,
considered the family as closely allied to the Flacourtiaceae. The interpretation of the
inflorescence structure used here differs from that of van Steenis l.c. and was kindly provided
by Dr. K. M. Wong.
SCYPHOSTEGIA Stapf
(Greek, scyphos = cup, stege = shelter; the cup-like floral bracts)

FMK (1894) 217, t. 17 (under Monimiaceae); Hutchinson l.c. (1964) 326, f. 187a & b; Baehni, Soc.
Phys. Hist. Nat. Geneva 54 (1937) 91, Ber. Schweiz. Bot. Ges. 48 (1938) 22, f. 13; Swamy, Proc. Nat.
Inst. Sc. India 19 (1953) 127, f. 138; Metcalfe l.c. 99; van Steenis l.c. 297; van Heel l.c. 107; Anderson
l.c. 323.
Dioecious shrubs or trees, without exudate. Twigs often strongly 34-angular, slender, more
or less zig-zag, brown, nodes swollen. Stipules free, triangular, caducous. Leaves simple,
alternate, pinnately veined. Inflorescence basic units of short-peduncled racemes, bearing
112 (in male) or 12 (in female) serially imbricate, tubular, obtuse bracts, with each bract
subtending a single flower; the tubular-bracteate raceme units solitary in the distal leaf axils
or arranged on a slender flowering main stalk terminal to a leafy branch. Flowers unisexual,
radially symmetrical, developing one at time from the base upwards on each raceme-like
unit; pedicel flattened, persistent; perianth 6-lobed (3 outer and 3 inner tepals), imbricate in
bud, orbicular, petaloid, connate, fleshy; male flowers: pedicels thin, dry and membranous,
lengthwise 2-veined; perianth lobes shorter than the tube, inner ones shorter than and alternate
with the outer ones, distinctly thickened at apex; stamens 3, connate into in a column with
common apical connective produced above the anthers, anthers 4-locular, extrorse, dehiscing
lengthwise; disc-glands 3, oblong, fleshy, opposite the stamens; female flowers: pedicels
shorter than in males; perianth lobes longer than the tube, equal, more or less persistent,
375
recurved in fruit; disc present, disc-glands absent; ovary superior, 1-locular, globose, style
discoid with a narrow central aperture, stigmatic lobes 912, fleshy, largely fused to form
a thick plate-like stigma with recurved margin; ovules numerous, crowded on the slightly
convex base of the ovary, surrounded at base by a fleshy, collar-like aril, placentation parietal.
Fruits fleshy, dehiscent capsules, crowned by persistent styles and stigmas, 912-ribbed, at
maturity breaking up at the mouth into 912 segments. Seeds numerous, dry, densely packed,
funiculate, shortly erect-patent hairy over whole surface, their needle-like bases surrounded
and partly joined with spongy tissue from the base.
Scyphostegia borneensis Stapf
Fig. 1.
(of Borneo)
l.c. 218, t. 17; Merrill, EB (1921) 272; Baehni l.c. (1937) 91; Masamune, EPB (1942) 305; Metcalfe l.c.
99; van Steenis l.c. 297; van Heel l.c. 107; Anderson, l.c. 323. Type: Haviland 1377, Borneo, Sabah, Mt.
Kinabalu, near Koung (holotype K).
Shrub or tree to 20 m tall, 40 cm diameter, low branching; crown spreading. Bark dark green
to dark brown or blackish, smooth, thin, sometimes lenticellate; inner bark yellow to brown or
grey. Wood white or slightly yellowish. Twigs sometimes lenticellate, with knob-like stipule
scars; stipules 12 1 mm. Leaves drying chartaccous; narrow-oblong to obovate-oblong, 7
18(24) 2.56.8 (8.5) cm; base rounded, often decurrent down the petiole, margin serrulate,
the teeth with thickened tips, apex acute to acuminate, acumen to c. 13 mm long; lateral veins
611 pairs, spreading at a narrow angle from the midrib, arching in the lower half and joining
in faint loops 12 mm from the margin, flat or sunken above; intercostal veins scalariform,
more or less parallel to the midrib; petioles 25(6) mm long, 1(3) mm thick, furrowed
above. Inflorescences: males 6.530 cm long, raceme units 12 cm long, peduncles 2.520
mm long, bracts 510 mm long, 23 mm wide, pedicels 46 mm long, c. 1 mm thick; females
58.5 cm long, raceme units 58 mm long, peduncles 1235 mm long, bracts 14 23 mm,
pedicels shorter than in the males. Flower: males: perianth tube 45 mm long, the outer lobes
orbicular-oblong or spathulate, c. 3 2 mm, the inner ones c. 2 1.5 mm, disc-glands c. 0.75
0.4 mm, staminal column 23 mm long, anthers c. 2 mm long; females: perianth tube c. 2
mm long, lobes c. 6 4 mm, ovary c. 4 mm. Fruits c. 18 15 mm. Seeds 56 0.81 mm,
funicle c. 3 mm long.
Vernacular names. Sabahkata manok (Dusun), sangkurat (Kedayan), tarukakang (Dusun),
tulau (Idahan). Sarawakdelako (Kelabit), rukam hantu (Iban).
Distribution. In Sabah, known from the Crocker Range through the interior to the east coast
and southwards from Sapulut to Tawau; also in Kudat at the northern most tip of Sabah. In
Sarawak, on the Rejang river from Kapit to Belaga and headwaters of the rivers to the north;
also on the Balleh river south of Kapit and on the Baram river near Lobang Garam. Also
known in Kalimantan.
Ecology. In Sabah, found in primary rain forest from flat and sloping ridge areas to steep-sided
riverbanks, streams and gorges, and also in logged and disturbed forests. In Sarawak, often
growing along banks of swift-flowing rivers, at 201700 m. Flowering and fruiting all the year
around with local variation.
376
Uses. In Sarawak, the leaves are pounded together with tuba roots and used as fish-poison.
Notes. This species can be mistaken for Flacourtia rukam (Flacourtiaceae) from the structure
of the tree and the similarity of the leaves. However, the 4-angular, zig-zag twigs, the serrulate
leaf margin, and the fruit being a dehiscent fleshy capsule rather than an indehiscent drupe
clearly distinguish S. borneensis from F. rukam.
378
TETRAMERISTACEAE
John B. Sugau
Sandakan, Malaysia
Hutchinson, Fam. Fl. Pl. (1959) 277; Merrill, EB (1921) 392 (as part of Theaceae); Ridley, FMP 1 (1922)
367 (as part of Ochnaceae); Masamune, EPB (1942) 474 (as part of Camelliaceae); Browne, FTSB
(1955) 259 (under Marcgraviaceae); Smythies, CST (1965) 134; Burgess, TBS (1966) 468; Cockburn,
TS 2 (1980) 105 (as part of Theaceae); Anderson, CLTS (1980) 329; Keng, TFM 4 (1989) 470.
Trees. Leaves spirally arranged, simple, entire, coriaceous, punctate with black glands
below; stipules none. Inflorescence axillary, a panicle with umbel-like clusters of flowers,
pedunculate, with conspicuous bracts. Flowers small, bisexual, 2-bracteolate; sepals 4 or 5,
free, imbricate, persistent; petals 4 or 5, free, imbricate; stamens 4 or 5, free, alternate with
petals, filaments flattened and slightly fused at the base, anthers basifixed, oblong-sagittate,
locules 2, separate, glandular at the base, opening longitudinally; ovary superior, locules 4 or
5, style one, stigma apically lobed or toothed; ovules one in each locule, basal, anatropous.
Fruit a globose to ellipsoid berry; exocarp leathery, mesocarp fleshy. Seeds 4 or 5, oblong;
endosperm copious; embryo straight.
Distribution. Two genera, one (Tetramerista) in W Malesia, the other (Pentamerista) in the
Guayana Highlands of S America.
Taxonomy. The Tetrameristaceae were formerly placed in various classifications under
either the Ochnaceae, Marcgraviaceae or Theaceae. They were treated as a separate family
by Hutchinson l.c. and other more recent systems of classification. The Marcgraviaceae
are distinct by having many ovules in its basically 1-locular ovary, and seeds with little or
no endosperm. The Ochnaceae and Theaceae are still regarded by many authors as being
heterogeneous and typically differ from the Tetrameristaceae by their numerous stamens
and axile placentation. In addition, Ochnaceae anthers dehisce mostly by terminal pores and
Theaceae seeds have little or no endosperm.
TETRAMERISTA Miq.
(Greek, tetra = four, meris = part; the 4-partite flower)

amat (Brunei Malay, Sabah), entuyut (Iban, Sarawak), tuyut (Dusun, Sabah)
Fl. Ind. Bat. Suppl. 534 (1861); Merrill l.c. 392; Ridley l.c. 367; Masamune l.c. 474; Smythies l.c. 134;
Burgess l.c. 468; Cockburn l.c. 105; Anderson l.c. 329; Keng l.c. 470.
379
Trees. Leaves large, spirally arranged, coriaceous. Inflorescence an axillary panicle with
umbel-like clusters of flowers, peduncle long. Flower 4-merous, small, radially symmetrical;
sepals 4, imbricate, persistent, the 2 outer ones larger; petals 4, persistent; ovary 4-angled, 4locular, septa thin; style one, stigma with 4 triangular lobes. Fruit at the base with persistent
petals, sepals and bracteoles. Seeds 4, oblong.
Distribution. Three species, confined to W Malesia. T. glabra is the only species found in
Sabah and Sarawak.
Taxonomy. The 4-merous flowers of Tetramerista distinguish it from Pentamerista, which is
a more recently discovered genus (in 1972) with 5-merous flowers.
Tetramerista glabra Miq.
Fig. 1.
(Latin, glaber = without hairs; referring to most parts of the plant)

l.c. 534; Ridley l.c. 367; Masamune l.c. 474 (as Tetramista glabra); Smythies l.c. 134; Anderson
l.c. 329; Keng l.c. 470; PROSEA 5 (1) (1994) 453. Type: Teysmann, s.n., W Sumatra, Sibolga district
(BO). Synonyms: Tetramerista crassifolia Hallier f. in Beih. Bot. Centralbl. 34, 2 (1916) 38, Merrill
l.c. 392 (as Tetramista crassifolia), Masamune l.c. 474 (as Tetramista crassifolia); Tetramerista
montana Hallier f. l.c. 39, Merrill l.c. 392 (as Tetramista montana), Masamune l.c. 474 (as Tetramista
montana).
Tree to 35 m tall, 110 cm diameter, fluted near the base; buttresses small and steep,
pneumatophores and stilt roots sometimes present. Bark fissured to flaky, rough, corky,
dark brown; inner bark thick, soft, fibrous, red to pinkish. Wood pale yellow, darkening on
exposure. Leaves mostly clustered at the distal portions of twigs, coriaceous, glabrous and
sometimes shiny on the upper surface, punctate with black glands below; oblanceolate, 822
48 cm; base attenuate, running down to the leaf-stalk base and forming narrow wings,
margin with small dark pit-glands, slightly recurved when dry, apex acute or rounded to
notched; midrib prominent and drying purple-brown on the lower side, flattened and drying
medium brown on the upper side, lateral veins 1115 pairs, flattened on upper side, slightly
prominent on the lower side, reticulations faint or invisible; petioles 23.5 cm long, winged.
Inflorescence a little- to much-branched panicle, peduncle 3.510.5 cm long, 34 mm thick,
rather stiffly erect, glabrous. Flowers 22.5 cm across, with 2 persistent bracteoles at the base
of the pedicel; pedicels 11.8 cm long; sepals oblong, 610 24 mm, glabrous; petals oblong
to lanceolate, 56 22.5 mm, glabrous; stamens 56.5 mm long, filaments 45 mm long;
anthers 11.5 mm long, glandular at the base; ovary broad-ellipsoid to cylindric, 34 1.52
mm, style 22.5 mm long. Fruits 1.53 cm across; fruit wall 23 mm thick. Seeds long-ovoid,
1015 58 mm, brown; germination epigeal.
Vernacular names. Sabahamat (Brunei Malay), punah (Malay), samondu (Brunei Malay),
suyut (Dusun), tuyot, tuyut (Brunei Malay, Dusun, Kedayan). Sarawakantuyut, entuyut
(Iban), kayu chelega (Punan Tutoh), kayu hujan (Malay), kayu tangiran (Berawan), punah
(Malay). Bruneiamat (Brunei Malay), terepit (Brunei Malay).
Distribution. Peninsular Malaysia, Sumatra and Borneo (Sabah, Sarawak, Brunei and
Kalimantan). In Sabah, found in the Sipitang, Weston, Kuala Penyu, Membakut, Beaufort,
Papar, Keningau and Tawau districts; in Sarawak, recorded for the Kuching, Sibu, Belaga,
Bintulu, Binatang, Miri and Baram districts.
380
Ecology. Lowland forests, usually in peat swamp forests, but occasionally also in kerangas
forest on white sands; lowlands up to 200 m. Occurrence of big trees is usually scattered but in
some peat swamp sites it may be more common. Young trees coppice freely (Browne l.c.).
Uses. Medium to heavy hardwood, suitable for indoor construction. The fresh logs sink in
water. Wong (DMT) gives a review of the timber characteristics.
382
ULMACEAE
E. Soepadmo & Z. Edi Hamli
Kepong, Malaysia
Merrill, EB (1921) 216; Masamune, EPB (1942) 237; Backer & Bakhuizen f., FJ 2 (1965) 10; Soepadmo, TFM 2 (1973)
413, FM 1, 8 (1977) 31; Anderson, CLTS (1980) 340; Corner, WSTM 3rd ed. 2 (1988) 735; Whitmore, Tantra & Sutisna,
CLK 2, 1 (1990) 361; Kessler & Sidiyasa, Trees of the Balikpapan-Samarinda Area (1994) 230.
Monoecious, unarmed shrubs or trees, without milky sap, mucilage cells or canals often present.
Indumentum of simple, bulbous-based, unicellular hairs and/or multicellular glandular hairs. Stipules
extrapetiolar or intrapetiolar (Parasponia, not in Borneo), caducous or rarely persistent, basally or
peltately (several species of Celtis) attached at the nodes. Leaves simple, alternate, pinnately veined
or 3-veined at base. Inflorescences male, female, or mixed (bearing functionally male and female or
bisextual flowers), 1many-flowered, lax or condensed panicles, racemes, or thyrses, axillary,
subterminal, or borne on leafless older twigs, or on short, lateral leafless new shoots; bracts caducous
or rarely persistent. Flowers unisexual, bisexual, or functionally unisexual; male flowers solitary or in
condensed cymose clusters along the main axes, sessile or short-stalked, perianth (4)5(7)-lobed,
lobes free or variously connate, imbricate or induplicate-valvate in bud, stamens erect in bud, as
many as and opposite perianth lobes, anthers dorsifixed or subdorsifixed, dehiscing lengthwise,
introrse or extrorse, pistillodes if present either well-developed or rudimentary, densely whitish to
silvery soft- or hirsute-pubescent; female and bisexual flowers sessile or stalked, solitary in the axils
of upper new leaves or arranged in various types of inflorescences, perianth herbaceous or thincoriaceous, (4)5(7)-lobed, lobes imbricate, connate at base, caducous or persistent, staminodes or
stamens as many as perianth lobes or absent, ovary superior, 2-carpellate, 1-locular, sessile, style 1,
tubular or absent, stigmatic arms 2, slender, often bifid to deeply lobed at the tip, stigmas papillose,
ovule 1, anatropous or hemi-anatropous, subapical and pendulous. Fruit a samara (Ulmus, not in
Borneo) or drupe, rounded, angular or compressed and lens-shaped in cross-section. Seeds with
scanty or copious or mostly without endosperm; embryo large, curved or straight (not in Borneo),
cotyledons plano-convex, fleshy, variously curved or folded; germination epigeal.
Distribution. Fifteen genera with about 200 species, distributed in the tropics, subtropics, and
temperate regions of Europe, Africa, continental Asia, Malesia, Australia, Pacific Is., and N, C and S
America. In Sabah and Sarawak, 4 genera with 10 species.
Ecology. In primary and secondary forests, on various types of soil, from sea level to 2000 m.
Taxonomy. In most recent classifications, it is widely accepted that Ulmaceae, together with
Moraceae, Urticaceae, and several other smaller related families, belong to the order Urticales.
383
The Ulmaceae differ from the Moraceae by the lack of lactiferous ducts, non-fleshy
receptacle, erect stamens in bud, and drupaceous or samara-type fruits; and from the Urticaceae by
their pendulous anatropous or hemi-anatropous ovules, erect stamens in bud, and the lack of stinging
hairs/glands.
Key to genera
1. Leaves exclusively pinnately veined; stipules leaving circular scars around the nodes after
falling. Perianth lobes of male flowers imbricate in bud, anthers subdorsifixed. Fruits
compressed and lens-shaped in cross-section; seeds without endosperm.......3. Gironniera
Leaves 3-veined at base, or if pinnately veined then the stipules not leaving circular scars
around the nodes after falling. Perianth lobes of male flowers induplicate-valvate in bud,
or if imbricate, anthers dorsifixed. Fruits 35-angular or rounded in cross-section or if
compressed and lens-shaped, the seed with scanty or copious endosperm..........................2
2. Leaf margins serrate throughout. Perianth lobes of male flowers induplicate-valvate in
bud. Fruits compressed, lens-shaped in cross-section............................................4. Trema
Leaf margins entire and undulate or crenate throughout, or entire and recurved or rarely
distantly serrulate in the upper half. Perianth lobes of male flowers imbricate. Fruits
rounded or 35-angular in cross-section..............................................................................3
3. Leaf base exclusively 3-veined. Male flowers always with pistillodes; anthers extrorse.
Staminodes always present in female flowers. Cotyledons foliaceous, broad, variously
folded or curved.......................................................................................................2. Celtis
Leaf base not 3-veined (Bornean taxa). Male flowers without pistillodes; anthers introrse.
Staminodes absent in female flowers. Cotyledons narrow, involute............1. Aphananthe
1. APHANANTHE Planch., nom. cons.

(Greek, aphanes = concealed, invisible, anthe = flower; small flowers)
Ann. Sc. Nat. 3, 10 (1848) 265, non Link, 1821; Soepadmo, FM 1, 8 (1977) 66; Whitmore, Tantra & Sutisna, CLK 2, 1
(1990) 361. Synonyms: Homoioceltis Blume, Mus. Bot. Lugd. Bat. 2 (1856) 64; Galumpita Blume l.c. 73; Gironniera
subgenus Galumpita (Blume) Hook. f. in Bentham & Hooker, Gen. Pl. 3 (1880) 356.
Deciduous or semi-deciduous trees, often buttressed. Stipules basally attached, not leaving circular
scars after falling. Leaves glabrous, coriaceous, 3-veined at base or pinnately veined. Inflorescences
axillary; bracts caducous; male inflorescence a condensed, many-flowered raceme, borne on the lower
parts of new shoots; mixed inflorescence 23-flowered. Flowers: males 45-merous, perianth lobes
membranous, imbricate in bud, filaments subulate, anthers dorsifixed, ovoid-subreniform, nonapiculate, introrse, pistillodes absent, replaced by a cluster of whitish to silvery, erect, soft, simple
hairs; females solitary in the axils of upper leaves of new shoot, or in clusters of 23, perianth lobes
45, persistent, staminodes absent, ovary sessile, ovoid-ellipsoid, rounded to angular in cross-section,
stigmatic arms tubular, ovule anatropous. Fruit a drupe, ovoid-globose, faintly 35-angular or
almost rounded in cross-section; endocarp hard and persistent. Seeds with scanty endosperm; testa
membranous, several-celled thick; cotyledons narrow, more or less equal, involute.
384
ULMACEAE (SOEPADMO & EDI HAMLI)
Distribution. About 45 species, distributed in Mexico, Madagascar, Sri Lanka, India, Myanmar,
China, Korea, Japan, Taiwan, Hong Kong, Indo-China, Thailand, Andamans, Malesia, Australia
(Queensland and New South Wales) and the Solomons. In Borneo, 1 species, so far known from
Sabah only.
Aphananthe cuspidata (Blume) Planch.
Fig. 1.
(Latin, cuspidatus = with a sharp, rigid point; the leaf apex)

In A. DC., Prod. 17 (1873) 209; Soepadmo l.c. (1977) 69; Whitmore, Tantra & Sutisna l.c. 361. Basionym: Cyclostemon
cuspidatum Blume, Bijdr. Fl. Ned. Ind. (1825) 599. Type: Blume, s.n., Java, Mt. Parang (holotype L). Synonyms:
Galumpita cuspidata (Blume) Blume l.c. (1856) 73; Gironniera nitida Benth., Fl. Hongk. (1861) 324; Gironniera reticulata
Thwaite, En. Pl. Zeyl. 1 (1861) 268; Gironniera lucida Kurz, For. Fl. Burma 2 (1877) 470; Gironniera cuspidata (Blume)
Kurz l.c. 470; Gironniera curranii Merr., Philip. J. Sc. 4 (1909) Bot. 251.
Tree to 30 m tall, 60 cm diameter; buttresses to 1 m high, and 2 m wide. Bark surface rough, greybrown, often flaky. Young twigs sparsely, minutely appressed-pubescent, glabrescent; older twigs
sparsely lenticellate. Stipules narrow, ovate-acute, 23x1 mm. Leaves pinnately veined; ovate-elliptic
to elliptic-oblong, (5)1014(20) x(2)36(8) cm; base rounded, subcordate or attenuate, symmetric
or occasionally slightly asymmetric, margin entire, recurved, or rarely distantly serrulate in the upper
half, apex cuspidate or acuminate, acumen to 2 cm long, pointed; midrib strongly raised beneath,
impressed to flattish above, lateral veins (5)78(10) pairs, slightly raised beneath, flattish above,
subparallel, arching and forming an angle of more than 60 with the midrib, weakly joining near the
margin, intercostal veins finely reticulate to subscalariform, indistinct on both surfaces; petioles
glabrous, (5)812(15) mm long, 12 mm thick, grooved on the upper side. Inflorescences male or
mixed; male inflorescences to 4 cm long, 1030-flowered; mixed inflorescences 23-flowered; bracts
ovate-acute, 0.250.5 x0.25 mm. Flowers: males globose, 1.52 mm diameter, short-stalked or rarely
sessile, perianth lobes obovate-lanceolate, c.2 x1 mm, anthers c. 1 mm diameter; females solitary or
23 together, ovoid-ellipsoid, c. 2x1 mm, perianth lobes coriaceous, ovate-acute, c.2x1 mm; ovary
sessile, ovoid, glabrous, c.1.5x1 mm, stigmatic arms 23 mm long. Fruits fleshy, ovoid, glabrous,
including the beak 1320 x 612 mm; stalk to 3 cm long, c. 1 mm thick.
Distribution. India, Sri Lanka, Myanmar, Andamans, Hong Kong, Indo-China, Thailand, NE
Sumatra, Borneo (Sabah), Java, Lesser Sunda Islands, Philippines (Mindanao, Palawan), and
Sulawesi.
Ecology. In Sabah, Aphananthe cuspidata, is extremely uncommon and so far recorded from a single
collection (SAN 64311) from the Ranau district. The deciduous or semideciduous habit, episodic
growth, structure, size, colour of the inflorescence and flowers suggest that pollination is effected by
wind. The drupes which turn to deep red when ripe are possibly dispersed by frugivorous birds.
2. CELTIS L.
(an old Latin plant-name)
Gen. Pl. ed. 5 (1754) 467; Ridley, FMP 3 (1924) 321; Backer & Bakhuizen f., FJ 2 (1965) 11; Soepadmo,
385
Fig. 1. Aphananthe cuspidata. A, leafy twig; BC, fruits; D, flowering leafy twig; E, male flower bud; F, male flower bud in
longitudinal section; G, female flower; H, female flower in longitudinal section. (AC from SAN 64311, DF from Koorders
30071b, GH from Koorders 27583b.)
386
TFM 2 (1973) 414, FM 1, 8 (1977) 55; Anderson, CLTS (1980) 340; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 361.
Synonym: Solenostigma Endl., Prod. Fl. Norf. (1833) 41.
Small to large, monoecious or polygamo-monoecious trees; often buttressed. Bark smooth or finely
fissured, often conspicuously warty lenticellate. Twigs initially densely yellow-brown or rufous-hairy,
glabrescent. Buds enclosed by the overlapping stipules or exposed. Stipules thick and tough, peltately
or basally attached at the nodes, caducous or permanent (not in Borneo). Leaves 3-veined at base.
Inflorescences male, female, or mixed, branched racemes or panicles, few- to many-flowered,
axillary or subterminal on new shoots. Flowers: males globular, stalked or sessile, perianth lobes (4)5, imbricate in bud, membranous, boat-shaped, at anthesis recurved, caducous, stamens inserted on
the densely pilose receptacle, filaments subulate, incurved in bud and then spreading, exserted at
anthesis, anthers ovoid to subreniform, dorsifixed, extrorse, pistillodes always present; functionally
female and bisexual flowers ovoid, stalked, perianth lobes (4)5, imbricate in bud, connate at base,
membranous, sparsely pubescent outside, boat-shaped, at anthesis recurved, caducous, stamens or
staminodes well-developed and functional or rarely rudimentary, other characters as in male flowers,
ovary ovoid-ellipsoid, sessile, style short or absent, stigmatic arms elongate, divergent, the tips entire
to deeply bifid, ovule anatropous. Fruit a drupe, fleshy, ovoid, ellipsoid or globose, 35-angular to
almost rounded in cross-section; exocarp thick and firm; mesocarp thin and fleshy, containing slimy
substances; endocarp hard and persistent, smooth or variously ridged or pitted. Seeds: testa
membranous, chalazal area broad, dark-coloured and close to the minute hilum; endosperm scanty,
oily or gelatinous; cotyledons broad, foliaceous, equal or unequal in thickness, flat or conduplicate,
variously folded.
Distribution. About 5060 species, widely distributed in the tropical and temperate regions of the
world, with the majority in the Old and New World tropics. In the Malesian region, 9 species are
known, 2 from Sabah and Sarawak.
Ecology. In primary and secondary lowland and hill forests, up to about 900 m, including exposed
and stunted forest stands on rocky shores and limestone hills. Pollination may be effected by wind or
insects, and fruit dispersal by frugivorous birds or sea current.
Key to Celtis species

Leaves thin-coriaceous, brittle on drying, ovate-elliptic, base strongly asymmetric, margins
crenate at least in the upper half, lateral veins 2(3) pairs, basal pair running up to half to
three-quarters the length of the blade. Stipules basally attached. Fruit more-or-less 4-angular
in cross-section..................................................................................................2. C. timorensis
Leaves thick-coriaceous, tough and not brittle on drying, elliptic-oblong, lanceolate-oblong,
to ovate-suborbicular, base mostly symmetric, margin entire and undulate, lateral veins 1 (to
rarely 2) pairs, the basal veins running throughout the length of the blade or very rarely
running up to about three-quarters the length of the blade. Stipules peltately attached. Fruits
more-or-less rounded in cross-section............................................................1. C. philippensis
1. Celtis philippensis Blanco
Fig. 2AD.

Fl. Filip. (1837) 197; Merrill, Spec. Blanc. (1918) 122; Soepadmo l.c. (1973) 416, l.c. (1977) 62;
387
Anderson l.c. 340; Whitmore, Tantra & Sutisna l.c. 361. Type: Merrill Spec. Blanc. 52, Phillippines, Luzon, Rizal Prov.,
Dec. 1912 (A, L). Synonyms: Celtis wightii Planch., Ann. Sc. Nat. 3, 10 (1848) 307, Backer & Bahkuizen f. l.c. 11;
Solenostigma philippinensis (Blanco) Miq., Fl. Ned. Ind. 1, 2 (1859) 220; Solenostigma wightii (Planch.) Miq. l.c. 220;
Celtis mindanaensis Elmer, Leafl. Philip. Bot. 8 (1915) 2842; Celtis collinsae Craib, Kew Bull. (1918) 370; Ridley l.c.
(1924) 322.
Low stunted shrub or treelet to large tree to 40 m tall, 80 cm diameter; buttresses, if present, to 4.5 m
tall. Bark smooth to finely fissured, pale grey to grey-brown. Twigs sparsely lenticellate, subsulcate,
glabrous or sparsely appressed-hairy. Stipules thick and tough, peltately attached, ovate-acute, 3
6 x13 mm, overlapping, completely enclosing the bud, sparsely appressed-hairy or glabrous. Leaves
thick-coriaceous, tough and not brittle on drying, glabrous, grey to dark brown; elliptic-oblong,
lanceolate-oblong to ovate-suborbicular, (6)814(16) x39 cm; base rounded or attenuate-rounded,
rarely truncate, mostly symmetric, margin entire and slightly undulate, apex rounded to acute; midrib
and lateral veins raised beneath, impressed to flattish above, lateral veins 1(2) pairs, basal veins
ascending, arching and running throughout the length of the blade or rarely to about three-quarters
the length of the blade, intercostal veins finely and densely reticulate or subscalariform or tessellate,
usually distinct beneath; petioles (3)513(17) mm long, (1)23(4) mm thick, somewhat grooved
on the adaxial side, glabrous or sparsely appressed-hairy. Inflorescences male or mixed, muchbranched panicles, many-flowered, including the bracts densely yellow-brown to rufous soft-hairy;
bracts ovate-acute, c. 3x1 mm; male inflorescences borne on the lower parts of new shoots, 24 cm
long, to 40-flowered; mixed inflorescences borne on the upper parts of new shoots, to 5 cm long, to
50-flowered, with the bisexual flowers on the distal parts of the rachis. Flowers: males c. 2 mm
across, perianth lobes ovate-elliptic, 1.52 x1 mm, filaments 11.5 mm long, anthers subreniform,
0.51x0.5 mm, pistillodes ovoid-cylindrical, compressed, 11.5 x0.5 mm; functionally female and
bisexual flowers c. 22.5 x2 mm, perianth lobes ovate-elliptic, c. 22.5 x1 mm, filaments 12 mm
long, anthers 0.51 mm diameter, ovary ovoid-cylindrical, 23(4)x(1)1.52 mm, glabrous except
at the base, stigmatic arms spreading, c. 11.5 mm long, bilobed to bifid at the tip. Fruits ovoidellipsoid, more-or-less rounded in cross section, only beaked when young, 813(15) x510 mm,
glabrous; exocarp less than 1 mm thick, sometimes lenticellate, turning orange to red when ripe;
endocarp more or less smooth. Seeds with unequal cotyledons.
Distribution. Tropical Africa to Madagascar, islands of the Indian Ocean (Reunion, Mauritius), India,
Myanmar, SE China, Hong Kong, Taiwan, Indo-China, Thailand, throughout Malesia, NE Australia
and the Solomons. In Sabah, scattered in coastal hill forests in the Semporna (including Sipadan
island), Sandakan, Lahad Datu (including Timbun Mata island) and Tawau districts in the east coast
and in Beaufort district (including Pulau Tiga) in the west coast. In Sarawak, twice collected from Mt.
Subis, Niah Caves, Miri (S. 16035 and S. 16041). Also known in Brunei.
Ecology. Understorey or main storey trees in primary and secondary forests, on various types of soil,
from sea level to about 700 m.
Uses. The timber, though not durable, is locally used in house-building.
Taxonomy. A rather variable species in terms of leaf and fruit size. Specimens collected from
limestone and rocky shores usually have smaller and narrower leaves and smaller fruits than those
from more inland coastal hills.
388
Fig. 2. Celtis philippensis (AD); Celtis timorensis (EF). A, fruiting leafy twig; B, flowering leafy twig; C, functionally
male flower; D, functionally female flower; E, fruiting leafy twig; F, fruit seen from the base. (A from SAN 66966, BD
from SAN 16496, EF from Clemens 30351.)
389
2. Celtis timorensis Span.
Fig. 2EF.
(of Timor Island)

Linnaea 15 (1841) 343; Soepadmo l.c. (1977) 65; Whitmore, Tantra & Sutisna l.c. 362. Type: Spanoghe, s.n., Timor (L;
n.v.). Synonyms: Celtis cinnamomea Lindl. ex Planch. l.c. 303; Celtis reticulosa Miq., Pl. Jungh. (1851) 69; Celtis hamata
Blume l.c. (1856) 72; Celtis waitzii Blume l.c. (1856) 71; Celtis crenato-serrata Merr., Philip. J. Sc. 5 (1910) Bot. 174.
Tree to 25 m tall, 30 cm diameter. Bark smooth, brownish grey, lenticellate; inner bark pale brown,
fibrous. Twigs initially rather densely rufous-pubescent, glabrescent; older twigs glabrous, rather
densely lenticellate. Stipules basally attached, free and scarious, linear-lanceolate, 510 x 12 mm.
Leaves thin-coriaceous, brittle on drying, except for the midrib and lateral veins glabrous, upper
surface dull grey to dark brown, lower surface chocolate-brown; ovate-elliptic, (6)1012(17) x(2
)45(8) cm; base rounded to subcordate, asymmetric, margin undulate and distantly crenate at least
in the upper half, apex acute to acuminate, acumen to 2 cm long; midrib and lateral veins flattish to
impressed above, raised beneath, lateral veins 2(3) pairs, arching and ascending, the basal pair
running to half to three-quarters the length of the blade, joining along the margin, intercostal veins
subscalariform, sparse, rather distinct beneath; petioles 515 mm long, 12 mm thick, adaxially
grooved. Inflorescences male or mixed, racemose, lax; axes slender, always longer than the petioles,
including the bracts sparsely rufous-hairy; bracts narrowly ovate-acute, c. 35 x12 mm; male
inflorescences borne on the lower leafless parts of new shoots, much-branched, 1020-flowered, to 3
cm long; mixed inflorescences borne in the axils of new leaves, 47-flowered, to 23 cm long.
Flowers: males c. 2 mm diameter, perianth lobes 1.52 x1 mm, filaments 11.5 mm long, anthers
reniform, c. 1 x0.5 mm, pistillodes present but strongly reduced in size; bisexual flowers (functionally
female) 23 mm diameter, perianth lobes 1.52 x1 mm, filaments 0.51 mm long, anthers
subreniform, 0.50.75 x0.5 mm, ovary ovoid-ellipsoid, c. 2 x 1 mm, stigmatic arms 11.5 mm long,
spreading. Fruits ovoid, more-or-less 4-angular in cross-section, strongly beaked, (4)58(10) x 6
mm, glabrous. Seeds with equal cotyledons.
Distribution. India, Bangladesh, Sri Lanka, Myanmar, Thailand, Indo-China, C Sumatra, Java,
Borneo, Philippines, and Lesser Sunda Islands (Flores, Timor). In Borneo, uncommon and known
from 6 specimens collected from the Mt. Kinabalu area, in Ranau district (Chew & Corner RSNB
5772, Clemens 30351, SAN 56520, SAN 92527, SAN 116594) and from Apin Apin near Laing Caves
in Keningau district (SAN 44568), and a few others from Kalimantan; so far no record from Sarawak
and Brunei.
Ecology. In hilly mixed dipterocarp forest and lower montane forest, at 6001500 m.
3. GIRONNIERA Gaudich.
(Proust de la Gironnier, prominent French citizen of Manila, c. 1840)
entabuloh, untoh bulu (Iban), medang kasap (Malay)
Voy. Bonite (1844) t. 85; Merrill, EB (1921) 216; Ridley, FMP 3 (1924) 317; Masamune, EPB (1942) 237; Brown, FTSB
(1955) 356; Phuphathanaphong, Thai For. Bull. 6 (1972) 49; Backer & Bakhuizen f., FJ 2 (1965) 12; Soepadmo, TFM 2
(1973) 417, FM1, 8 (1977) 70; Anderson, CLTS (1980) 340; Corner,
390
WSTM 3rd ed. 2 (1988) 735; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 362. Synonyms: Nemostigma Planch., Ann. Sc.
Nat. 3, 10 (1848) 265; Helminthospermum Thwaite in Hooker, J. Bot. Kew Misc. 6 (1854) 301.
Shrubs to large trees, very rarely buttressed. Bark smooth to finely fissured, grey-brown, often rugose
with pustular lenticels; inner bark usually pinkish, fibrous inwards, granular outwards. Sapwood hard,
pale fawn. Young parts densely or sparsely covered with golden yellow or yellowish brown
indumentum. Stipules basally attached, overlapping and completely enclosing the ellipsoidcylindrical terminal buds, after falling leaving distinct circular scars around the nodes. Leaves
pinnately veined, lateral veins parallel and regularly spaced. Inflorescences male or female, very
rarely mixed, axillary or borne on older, leafless twigs, 1many-flowered, paniculate, racemose or
thyrsoid; bracts caducous in male inflorescences, rather persistent in female and mixed inflorescences.
Flowers: males globular, sessile or short-stalked, perianth lobes 46, imbricate in buds, stamens (4
)5(6), glabrous or hairy, filaments subulate, inflexed, anthers introrse, ovoid-reniform, apiculate,
sub-dorsifixed, pistillodes present, rather well-developed or strongly rudimentary, densely covered
with long whitish or silky erect hairs; females ovoid-ellipsoid, compressed, perianth lobes 45,
usually unequal in size, persistent, staminodes absent, ovary ovoid-ellipsoid, strongly compressed,
sessile, densely or sparsely appressed-hairy, glabrescent, stigmatic arms to 1.5 cm long, curled in bud,
later spreading, rather persistent; ovules anatropous. Fruit a drupe, compressed ovoid-globose, lensshaped in cross-section, subtended by persistent perianth lobes; exocarp thin, strongly adnate to the
hard and persistent endocarp. Seeds mostly sterile; testa membranous, several-celled thick; endosperm
absent; cotyledons fleshy, narrow, equal, curved.
Distribution. Five species, distributed in E and SE Asia extending from Sri Lanka to Andaman Is.,
Thailand, Indo-China, China, Malesia, Micronesia, Melanesia, and Polynesia. In Malesia, throughtout
the region except for the eastern parts of Java, Lesser Sunda Islands, and SE Maluku. In Sabah and
Sarawak, represented by 3 species.
Ecology. In primary and secondary forests, from sea level to 1600 m, on different types of soil,
including those derived from limestone. Flowering and fruiting throughout the year. Pollination is
most likely by wind. The drupes which turn bright yellow or orange when ripe are mostly sterile, and
dispersed by frugivorous birds.
Key to Gironniera species

1. Leaves densely covered with yellowish brown, soft, slender hairs beneath; lateral veins
(10)1216(18) pairs, adjacent veins 58 mm apart, straight and parallel, not joining
near the margin. Male inflorescence a much-branched panicle; male flowers in condensed
cyme-like clusters of 510.............................................................................. 1. G. nervosa
Leaves glabrous or only sparsely covered with yellowish-brown, appressed short-hairs on
the midrib and lateral veins beneath; lateral veins (5)68(10) pairs, adjacent veins (7
)1015 mm apart, joining near the margin. Male inflorescence an unbranched or littlebranched raceme or panicle; male flowers solitary or in clusters of 35 on short,
condensed secondary branches of the inflorescence............................................................2
2. Leaves thick-coriaceous, sparsely covered with yellowish brown, short hairs beneath,
especially on the midrib and lateral veins; length 22.5 times the width, base more-or-less
391
asymmetric; intercostal veins laxly subscalariform. Male inflorescences little-branched

panicles, 40100-flowered. Female inflorescences (2)510(15)-flowered. Mature fruits
1012 x 89 mm, 56 mm thick................................................................3. G. subaequalis
Leaves chartaceous to thin-coriaceous, glabrous on both surfaces; length 33.5 times the
width, base more-or-less symmetric; intercostal veins densely, finely tessellate. Male
inflorescences unbranched racemes, 1530-flowered. Female inflorescences 13flowered. Mature fruits 78 x56 mm, 34 mm thick.................................2. G. parvifolia
1. Gironniera nervosa Planch.
(Latin, nervosus = conspicuously veined; the leaves)
l.c. 338; Merrill, EB (1921) 216; Ridley l.c. 320; Masamune l.c. 237; Soepadmo l.c. (1973) 419, l.c. (1977) 74; Corner l.c.
736. Type: Porter ex Wallich Cat. 7289, Penang (K). Synonyms: Gironniera penangiana Gandog., Bull. Soc. Bot. Fr. 66
(1919) 289; Gironniera sponioides Gandog. l.c. 289; Gironniera hirta Ridl., J. Str. Br. Roy. As. Soc. 82 (1920) 194, l.c.
(1924) 321, Soepadmo l.c. (1973) 417, l.c. (1977) 74, syn. nov.
Medium-sized to large tree to 40 m tall, 60 cm diameter; buttresses sometimes present. Bark smooth
or finely fissured, grey-green to dark grey-brown, often hoop-marked and lenticellate; inner bark pale
yellow. Young twigs, stipules, terminal buds, petioles, inflorescences and fruits densely covered with
long or short silky, soft, erect or appressed, yellowish-brown hairs. Terminal buds ovoid-ellipsoid or
conical 13 x0.20.6 cm. Stipules 1.53.5x0.250.5 cm. Leaves thick-coriaceous, rigid, upper surface
glabrous except for the midrib and lateral veins, lower surface densely covered with long, yellowish
brown, soft, silky, slender hairs; elliptic-lanceolate to elliptic-oblong, (6)1020(29)x(2.5)410(
13) cm, broadest at the middle, length 23 times the width; base rounded, subcordate, or attenuate,
often asymmetric, margin entire and recurved or rarely distantly serrulate in the upper half, apex
rounded-acute or acuminate; midrib and lateral veins strongly raised beneath, flattish or impressed
above, lateral veins (10)1216(18) pairs, adjacent veins 58 mm apart, straight and parallel for the
most part, arching but not joining near the margin, intercostal veins densely scalariform or
subscalariform, distinctly raised beneath, obscure to faintly visible above; petioles (3)510(15) mm
long, 12(3) mm thick, terete, subterete or flat on the upper side near the base. Inflorescences male
or female, axillary, borne on separate twigs; bracts ovate-acute, 12 x0.51 mm; male inflorescences
slender, lax, pendent, much-branched panicles of 510-flowered, condensed cyme-like clusters, 78
cm long, up to 100-flowered; female inflorescences unbranched or branched panicles, 2.56 cm long,
(2)520(25)-flowered. Flowers: males 12 mm diameter, sessile or short-stalked, perianth lobes
broadly ovate-acute, (1)1.52x11.5 mm, densely short, appressed-hairy outside, stamens (4)5(6),
sparsely covered with short, appressed hairs, filaments 11.5 mm long, anthers ovoid-reniform, 0.51
mm diameter, glabrous or hairy, pistillodes strongly rudimentary; females sessile or short-stalked, 2
3x1.52 mm, perianth lobes narrowly ovate-acute, 1.52x1 mm, densely appressed short-hairy
outside, ovary densely appressed-hairy, 1.53x12 mm, stigmatic arms 510 mm long. Fruits
subglobose or ovoid, dull orange, 510x36 mm, 35 mm thick, short-beaked.
Vernacular names. Sabahhugot hugot (Dusun), kiga (Dusun Sungei), luazon (Kadazan), sosopon
bonging (Dusun Putatan). Sarawakmedang kasap (Malay), puloh (Iban).
392
Distribution. Thailand, Sumatra, Peninsular Malaysia, Borneo, Maluku, and New Guinea. Common
in Sabah and Sarawak; also known for Brunei and Kalimantan.
Ecology. In primary and secondary forests, from sea level to 1400 m, mostly below 700 m; often
locally common as understorey trees in lowland mixed dipterocarps forest and hill and lower montane
forest or in forest on ultramafic soil.
Uses. The timber is locally used in house-building.
Taxonomy. In previous accounts, Soepadmo (l.c. 1973 & 1977), based on a limited number of
specimens, recognised G. hirta Ridl. as a species distinct from G. nervosa. However, recent
collections from Sabah and Sarawak indicate that these two species should be combined, due to the
presence of intermediate specimens integrating morphological characters of the two species.
2. Gironniera parvifolia Planch.

(Latin, parvum = small, folium = leaf)
l.c. 338; Ridley l.c. (1924) 321; Soepadmo l.c. (1973) 419, l.c. (1977) 75; Corner l.c. 689. Type: Griffith, s.n. (= Herb.
Lemann), Malacca (K). Synonyms: Gironniera subaequalis var. ceylanica Planch l.c. (1848) 339; Helminthospermum
scabridum Thwaite in Hooker, J. Bot. Kew Misc. 6 (1854) 303, t. 9c; Gironniera paucinervia Merr., J. Str. Br. R. As. Soc.
77 (1917) 189, l.c. (1921) 217, Masamune l.c. 238; Gironniera zeylanica Gandog., Bull. Soc. Bot. Fr. 66 (1919) 288;
Gironniera scabrida (Thwaite) Alston in Trimen, Fl. Ceyl. 6 (1931) 267.
Small to medium-sized tree to 20 m tall, 20 cm diameter. Bark smooth to finely fissured, lenticellate,
grey-green or grey-brown; inner bark pale yellow or pinkish. Terminal buds (5)815(18) x13 mm.
Stipules linear-acute, (6)820(23) x 25 mm. Leaves chartaceous to thin-coriaceous, glabrous or
rarely sparsely appressed-hairy; elliptic-lanceolate, very rarely elliptic-obovate, (4)715(19) x(1.5
)35(6) cm, length 33.5 times the width, broadest at or above the middle; base attenuate or rounded,
more or less symmetric, margin entire or sometimes minutely and distantly serrulate in the upper half
especially in young specimens, apex attenuate-acute or rounded-acuminate, acumen to 2.5 cm; midrib
and lateral veins slightly raised beneath, flattish above, lateral veins (4)68(10) pairs, adjacent
veins 710 mm apart, subparallel for the most part and forming an angle of up to 60 with the midrib,
arching and joining along the margin, intercostal veins densely finely tessellate, visible beneath,
obscure above; petioles (4)57(10) mm long, 13 mm thick, adaxially grooved, sparsely appressedpubescent, glabrescent. Inflorescences male or female, axillary, borne on separate twigs, including
bracts sparsely appressed-pubescent, glabrescent; bracts ovate-acute, membranous, c. 1x0.5 mm; male
inflorescence a slender, pendulous, unbranched or little-branched raceme, 10 30-flowered, axes 58
cm long, 0.30.7 mm thick; female inflorescence a slender, 13(5)-flowered raceme, 4 10 cm long,
axes 0.20.3 mm thick. Flowers: males solitary or in clusters of 35 along the axes, short-stalked or
sessile, 13 mm diameter, perianth lobes broadly ovate, 12.5x12 mm, stamens (4)5(6), filaments
11.5 mm long, anthers ovoid-reniform, 0.71 mm diameter, glabrous, pistillodes strongly
rudimentary or cylindrical, to 0.5 mm across, densely covered with long, yellowish white wavy hairs;
females 23 x 2 mm, short-stalked, perianth lobes mostly 4, 1.52.5 x 12 mm, ovary sparsely
appressed-pubescent, 23 x 2 mm, stigmatic arms to 1.5 mm long. Fruits 78
393
x 56 mm, 34 mm thick, ripening orange, sparsely appressed-pubescent, glabrescent; beak 510 mm

long.
Vernacular names. Sabahampas tebu, hampas tebu, medang kasap (Malay), rozan, ruazon (Dusun
Kinabatangan). Sarawakbelian landak (Iban), tepada (Kelabit).
Distribution. Sri Lanka, Sumatra, Peninsular Malaysia, Borneo. In Sabah, rather common in the
western districts (Kota Merudu, Tuaran, Kota Kinabalu, Penampang, Ranau, Papar, Tambunan,
Beaufort and Sipitang); but in Sarawak, very rare and so far known from a single collection from the
Kelabit Highlands (S. 35414).
Ecology. In primary as well as secondary forest, from sea level to 1600 m, mostly below 1000 m.
Occasionally in kerangas forest.
Uses. The timber is heavier and more compact than that of G. nervosa but not durable. It is locally
used in house-building.
3. Gironniera subaequalis Planch.
Fig. 3.
(Latin, subaequalis = not completely equal; the asymmetric leaf base)

l.c. (1848) 339, p.p., excl. var. ceylanica; Merrill l.c. (1921) 217; Ridley l.c. (1924) 320; Masamune l.c. 238; Backer &
Bakhuizen f. l.c. 12; Soepadmo l.c. (1973) 419, l.c. (1977) 75; Corner l.c. 690. Type: Spanoghe, s.n. (= Herb. Hooker), Java
(K). Synonym: Gironniera nervosa var. subaequalis (Planch.) Kurz, For. Fl. Burma 2 (1877) 470.
Small to large tree to 40 m tall, 60 cm diameter; buttresses up to 1.5 m high. Bark smooth to finely
fissured, pustular or lenticellate, grey brown; inner bark granular outwards, fibrous inwards, yellowbrown. Sapwood pale yellow. Young twigs, stipules, petioles, and inflorescences sparsely appressedhairy, glabrescent. Terminal buds ellipsoid, 23 x 0.250.5 cm, densely appressed, yellowish
pubescent. Stipules linear-lanceolate, 1.52.5 x 0.250.5 cm. Leaves thick-coriaceous, except for
midrib and lateral veins glabrous; broadly ovate-elliptic to elliptic-oblong, (6) 1020(27) x (3)5
9(13) cm, length 2 2.5 times the width, broadest below or at the middle; base attenuate or rounded,
more or less asymmetric, margin entire or occasionally (especially when young) distantly serrulate in
the upper half, apex rounded or attenuate-acute; midrib and lateral veins prominent beneath, flattish
above, sparsely yellowish-brown, appressed short-hairy or glabrous beneath, lateral veins (5) 68 (
10) pairs, adjacent veins 10 15 mm apart, arching and joining along the margin, intercostal veins
laxly subscalariform, clearly visible beneath, flattish and faintly visible or obscure above; petioles (5-)
1015 (26) mm long, 13 (5) mm thick, terete or adaxially flat near the base. Inflorescences male
or female, axillary, borne on separate or rarely on the same twigs, including bracts sparsely to densely
yellowish-brown, appressed short-hairy, glabrescent; bracts ovate-acute, 1 2 x 0.5 1 mm; male
inflorescences pendent, little-branched panicles, 40 100-flowered, axes 38 cm long, 0.5 1 mm
thick; female inflorescences racemose or more commonly paniculate (branched), axes 3 6 cm long,
1 1.5 mm thick, (2)5 10(15)-flowered. Flowers: males 1.5 2.5 mm diameter, sessile or shortstalked, solitary along the axes or in clusters of 3 5 on short, condensed secondary branches of the
panicle, perianth lobes sparsely appressed short-hairy, glabrescent, broadly ovate-rounded, 22.5 x 1
2 mm, stamens 5, filaments 12 mm long, anthers ovoid-subreniform, c. 1 mm long,
394
Fig. 3. Gironniera subaequalis. A, fruiting leafy twig; B, young fruit; C, stipules; D, male inflorescence; E, male flower and
dorsal view of a stamen. (AC from SAN 117970, D from SAN 92016, E from FRI 19281.)
395
glabrous, pistillodes cylindrical, to 1 mm long, or strongly rudimentary, densely covered with long,
yellowish, wavy or erect hairs; females solitary and short-stalked along the axes, 24 x mm, perianth
lobes broadly ovate-acute, 1.52 x11.5 mm, sparsely appressed-pubescent outside, ovary 23 x2
mm, densely appressed-pubescent, glabrescent, stigmatic arms to 2 cm long. Fruits 1012 x89 mm,
56 mm thick, sparsely appressed-pubescent, glabrescent; beak 25 mm long; ripening yellow-orange.
Vernacular names. Sabahkuayun, ruwayon (Dusun), luazon, russen (Dusun Kinabatangan).
Sarawakmedang kasap (Malay), untoh bulu (Iban).
Distribution. A rather variable species, widely distributed in the Andaman Is., Myanmar, China,
Hong Kong, Indo-China, Thailand, and Malesia (except the Lesser Sunda Is.). Common in Sabah and
Sarawak; also known in Brunei and Kalimantan.
Ecology. Understorey or main canopy trees in primary and secondary forests, from sea level to 1600
m, on various types of soil; more common in the lowlands between 100 and 1000 m. Flowering and
Uses. The timber, which is heavier and darker than that of G. nervosa, is locally used in housebuilding, flooring and making planks.
4. TREMA Lour.
(Latinized Greek word, trema = hole, gap, pit; the pitted endocarp of the fruit)
halindagong, randagong (Dusun, Sabah); kereneong (Iban, Sarawak); marong (Malay, Sarawak); towi
(Bidayuh, Sarawak)
Fl. Coch. 2 (1790) 562; Merrill, EB (1921) 217, PEB (1929) 45; Ridley, FMP 3 (1924) 319; Masamune, EPB (1942) 238;
Brown, FTSB (1955) 359; Backer & Bakhuizen f., FJ 2 (1965) 11; Elias, J. Arn. Arb. 51 (1970) 37; Soepadmo, TFM 2
(1973) 420, FM 1, 8 (1977) 47; Anderson, CLTS (1980) 341; Wong, DMT (1982) 247; Corner, WSTM 3rd ed. 2 (1988)
738; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 362; Kessler & Sidiyasa, Trees of the Balikpapan-Samarinda Area
(1994) 231. Synonym: Sponia Commers. ex Lamk, Dict. 4 (1795) 138.
Trees or shrubs, often buttressed and with spreading and drooping branches. Stipules basally attached,
overlapping but free, caducous. Leaves pinnately veined and 3-veined at base. Inflorescences
axillary, paniculate or thyrsoid, many-flowered, condensed or lax, male, female or mixed (with
structurally bisexual but functionally male and female flowers in the same inflorescence), variously
densely pubescent; bracts caducous. Flowers: males globose, perianth lobes (4)5, induplicatevalvate in bud, anthers dorsifixed, introrse, pistillodes present, hirsute; females ovoid, perianth lobes
(4)5, glabrous or sparsely short-hairy, staminodes rarely present, ovary ovoid, glabrous, slightly
compressed, sessile, style short, ovules anatropous or hemi-anatropous. Fruit a drupe, compressed,
lens-shaped in cross-section; exocarp fleshy and fibrous; endocarp stony and very hard, pitted. Seeds
with scanty or copious endosperm; cotyledons equal, curved.
396
Distribution. About 1015 species, widely distributed throughout the tropics and subtropics. In Asia
(67 species), ranging from the warmer parts of the Himalayas, extending northeastwards to China, S
Japan and southeastwards through India, Myanmar, Thailand, Indo-China, and Malesia to the tropical
and subtropical parts of Australia and Pacific Islands as far east as Tahiti. In Africa (34 species), it
occurs south of the Sahara to S Africa and Madagascar. In America (45 species), known from C and
S Florida and Mexico, extending southeastwards through Central America, Bermuda, and the
Bahamas, to the Greater Antilles and southwards to South America as far south as the northern parts
of Argentina. In Sabah and Sarawak, 4 species are known.
Ecology. Throughout its range of distribution, the genus seems to thrive and often grow gregariously
in newly cleared areas on various types of soil ranging from heavy laterite to limestone and soils
derived from volcanic ash, from sea level to 2000 m. Flowering and fruiting prolifically throughout
the year. Pollination is by wind or insects. The fruits, which turn orange, red or black when ripe, are
dispersed by frugivorous birds, especially bulbuls.
Uses. The timber of Trema orientalis and T. tomentosa is soft and light, with density ranging from
415465 kg/m3 air dry. The sapwood is straw-coloured and is sharply differentiated from the heartwood, which is light brown with a grey tinge. The wood is very easy to saw and to work, but not
durable; it has been used locally for making tea-chests, matches, and wooden clogs, and according to
Brown (l.c.) has the potential for use as pulp-wood. The fibrous inner bark is locally used as rough
cordage.
Key to Trema species

1. Lower leaf surface densely rough tomentose with short, matted, rufous or glaucous,
simple hairs and multicellular glandular hairs. Petioles (2)36(8) mm long. Mature
fruits subglobose, 1.52 mm across...........................................................1. T. angustifolia
Lower leaf surface glabrous or sparsely appressed hairy, or densely velvety tomentose
with silvery or glaucous, long simple hairs or combination of long simple hairs and short
multicellular glandular hairs. Petioles (7)1015(25) mm long. Mature fruits ovoid, (2
)34(5)x23(4) mm..........................................................................................................2
2. Leaves chartaceous to thin-coriaceous, smaller, (3)510(13)x(1.5)24(5.5) cm, lower
surface glabrous or rarely sparsely short-pubescent; base rounded to attenuate, mostly
symmetric, lateral veins (2)34(5) pairs. Inflorescences axes slender, less than 1 mm
thick. Fruits smaller, 23x2 mm...................................................................2. T. cannabina
Leaves mostly thick-coriaceous, larger, (5)1015(19)x(2)37(10) cm, lower surface
densely velvety tomentose with long simple hairs or densely tomentose with long simple
hairs and short, multicellular glandular hairs; base cordate, subcordate or rounded, mostly
asymmetric; lateral veins 58 pairs. Inflorescence axes stout, thicker than 1 mm. Fruits
larger, (2)35x24 mm.......................................................................................................3
3. Lower leaf surface densely tomentose with combination of silvery, long, appressed hairs
and short glandular hairs (lamina surface hidden, even under high magnification). Fruits
larger, 35x24 mm.......................................................................................3. T. orientalis
397
Lower leaf surface densely or sparsely velvety tomentose with glaucous, erect, simple hairs
(lamina surface clearly visible between the hairs even under low magnification). Fruits smaller,
23x2 mm.................................................................................................................4. T. tomentosa
1. Trema angustifolia (Planch.) Blume

Mus. Bot. Lugd. Bat. 2 (1856) 58; Merrill l.c. (1921) 217; Ridley l.c. (1924) 219; Masamune l.c. 238; Soepadmo l.c. (1973)
421, l.c. (1977) 54; Whitmore, Tantra & Sutisna l.c. 362. Basionym: Sponia angustifolia Planch., Ann. Sc. Nat. 3, 10 (1848)
326. Type: Wallich 3691, Penang (holotype K; isotype CGE). Synonyms: Sponia acuminatissima Miq., Fl. Ned. Ind. Suppl.
(1873) 410; Trema acuminatissima (Miq.) Boerl., Handl. 3 (1900) 358; Trema orientalis var. bicolor De Wit, Bull. Bot.
Gard. Btzg. 3, 18 (1949) 190.
Small tree to 7 m tall, 15 cm diameter. Bark smooth, grey-brown; inner bark greenish. Twigs slender,
spreading and drooping, densely covered with rufous, multicellular glandular hairs and glaucous short
and matted simple hairs, subglabrescent. Stipules linear-lanceolate, 12(3) x1 mm. Leaves
chartaceous to thin-coriaceous, discolorous, upper surface strongly scabrate, dark chocolate-brown to
blackish-brown, lower surface densely rough tomentose with short, matted, rufous to glaucous simple
and multicellular glandular hairs; narrowly ovate-lanceolate to lanceolate, (3)46(8) x (1)23(4)
cm, length 33.5 times the width, broadest below or at the middle; base symmetric, rounded to
attenuate, margin finely serrate throughout, apex acute to acuminate; midrib and lateral veins slightly
raised beneath, impressed above, lateral veins 35 pairs, straight, ascending, intercostal veins fine,
subscalariform to tessellate, obscure above, faintly visible beneath; petioles (2)36(8) long, 0.51
mm thick, densely short-hairy. Inflorescences male or mixed, rufous tomentose, much-branched, (5
)1015(30)-flowered, axillary, very condensed, shorter than or as long as the petioles; bracts
narrowly ovate-acute, 0.51 x0.250.5 mm. Flowers: males 12 mm diameter, perianth lobes 5,
subelliptic, (0.75)11.5x1 mm, filaments to 1 mm long, pistillodes obovoid-ellipsoid, 0.751 x 0.5
mm, strongly compressed; females c. 2 x1 mm, perianth lobes 5, narrowly lanceolate, acute, 0.5
1x0.250.5 mm, staminodes absent, ovary 12 x1 mm, stigmatic arms c. 0.51 mm long, incurved or
spreading. Fruits subglobose, 1.52 mm across, turning to orange or red when ripe. Seeds with
copious endosperm.
Vernacular name. Sabahrandagong (Dusun Ranau).
Distribution. China, Thailand, Indo-China, Sumatra, Peninsular Malaysia, Borneo, and Sulawesi. In
Sabah and Sarawak, apparently little-collected, known only from 3 collections (SAN 99290 from
Beluran in Sabah, and Native collector E. 284 and Haviland s.n. from Mt. Matang, Sarawak).
Ecology. Occasional in secondary forest and newly cleared areas in the lowlands and lower montane
areas, from sea level to 1200 m.
2. Trema cannabina Lour.

(Latinized Greek word, kannabinos = resembling the hemp or cannabis plant)
398
Fl. Coch. 2 (1790) 563; Backer & Bakhuizen f. l.c. 12; Soepadmo l.c. (1973) 421, l.c. (1977) 50; Anderson l.c. 341; Corner
l.c. 738; Whitmore, Tantra & Sutisna l.c. 363. Type: Loureiro, s.n., Indo-China (BM). Synonyms: Celtis amboinensis
Willd., Sp. Pl. 4, 2 (1805) 997; Sponia amboinensis (Willd.) Decne, Nouv. Ann. Mus. Hist. Nat. 3, 3 (1834) 498; Trema
amboinensis (Willd.) Blume l.c. (1856) 61, Merrill l.c. (1921) 217, l.c. (1929) 45, Ridley l.c. (1924) 319; Trema orientalis
var. amboinensis (Willd.) Kurz, For. Fl. Burma 2 (1877) 469; Sponia virgata Planch. l.c. 316; Trema virgata (Planch.)
Blume l.c. (1856) 59, Ridley l.c. (1924) 319.
Shrub or much-branched treelet to 6 m tall, 10 cm diameter. Bark smooth, grey-brown, lenticellate.

Twigs slender, spreading, often drooping, sparsely covered with short, appressed silvery hairs,
glabrescent. Stipules linear-lanceolate, 57 x12 mm. Leaves chartaceous to thin-coriaceous, upper
surface glabrous and scabrate, lower surface glabrous or occasionally sparsely appressed hairy on
the midrib and veins; narrowly to broadly ovate or elliptic-lanceolate, (3)510(13) x(1.5)24(
5.5) cm, length 23(4) times the width, broadest below or at the middle; base rounded to attenuate,
very rarely subcordate, mostly symmetric, margin serrulate to denticulate for its entire length, apex
acute to caudate with a sharp tip; midrib and lateral veins slightly raised beneath, impressed above,
lateral veins (2)34(5) pairs, subparallel for the most part and arching towards margin, intercostal
veins fine, subscalariform, obscure to visible beneath; petioles (7)812(15) long, 12 mm thick,
glabrescent. Inflorescences male or mixed; axes slender, less than 1 mm thick, (5)1015(20)flowered, at anthesis lax, 12.5 cm long, sparsely greyish appressed-hairy; bracts ovate-acute, 23 x1
mm. Flowers: males 12 mm diameter, sparsely hairy outside, glabrescent, perianth lobes (4)5,
membranous, oblong-lanceolate, c. 11.5 x0.51 mm, filaments c. 1 mm long, anthers c. 0.51 x0.5
mm, pistillodes obovoid, compressed, 0.51 x0.5 mm; females 1.52 x11.5 mm, perianth lobes
mostly 5, membranous, glabrous, ovate-acute, 11.5 x0.5 mm, staminodes absent, ovary 0.51 x0.5
mm, stigmatic arms spreading or incurved. Fruits ovoid, 23 x2 mm, turning to deep-orange or red
when ripe. Seeds with copious endosperm.
Vernacular names. Sabahrandagong (Dusun Ranau). Sarawakkereneong (Iban).
Distribution. Myanmar, China, Indo-China, Thailand, throughout Malesia to Australia, Melanesia, W
Polynesia, and Micronesia. Common in Sabah and Sarawak; also known for Brunei and Kalimantan.
Ecology. Common as a pioneer plant in newly cleared areas along road sides, edges of forests,
hillsides, thickets, and in young secondary vegetation, from sea level to 1500 mm.
Uses. In Sabah and Sarawak, the roots or leaves are boiled and the water is taken as a drink or used to
bathe for curing rheumatism and measles.
3. Trema orientalis (L.) Blume

(Latin, orientalis = from the East)
Mus. Bot. Lugd. Bat. 2 (1856) 62; Bentham, Fl. Austr. 6 (1873) 158; Backer & Bakhuizen f. l.c. 12; Soepadmo l.c. (1973)
421, l.c. (1977) 51; Corner l.c. 739; Whitmore, Tantra & Sutisna l.c. 363. Basionym: Celtis orientalis L., Sp. Pl. 2 (1753)
1044. Type: Herb. Hermann, Ceylon (BM). Synonyms: Celtis rigida Blume, Bijdr. Fl. Ned. Ind. (1825) 486; Sponia rigida
(Blume) Decne, Nouv. Ann. Mus.
399
Hist. Nat. 3, 3 (1834) 498; Trema rigida (Blume) Blume l.c. (1856) 61, Merrill l.c. (1921) 217, Masamune l.c. 238; Trema
orientalis var. rigida (Blume) Lauterb, Bot. Jahrb. 50 (1913) 320; Trema orientalis var. bicornis De Wit, Bull. Bot. Gard.
Btzg. 3, 18 (1949) 189.
Small to large tree to 36 m tall, 90 cm diameter; buttresses, if present, to 2 m high. Bark smooth to
finely fissured, lenticellate, grey-brown or whitish grey; inner bark greenish to reddish-brown. Twigs,
stipules, petioles, and inflorescences densely tomentose with appressed and matted or erect silvery
simple hairs and short multicellular glandular hairs. Stipules linear-lanceolate to ovate-acute, 26 x1
2 mm. Leaves thick-coriaceous, or rarely thin-coriaceous, often rigid and brittle, usually discolorous,
upper surface dull grey-brown or grey-green (in dried specimens), scabrate and sparsely covered with
bulbous-based simple hairs, lower surface densely tomentose with a combination of silvery, glaucous
or grey-brown, long appressed simple hairs and shorter crowded multicellular glandular or crisped
hairs which completely cover the lamina surface; ovate, ovate-lanceolate to narrowly ellipticlanceolate, (6)1015(20) x(1.5)47(10) cm, length (2)34(5.5) times the width, broadest at or
mostly below the middle; base cordate, rounded or sometimes truncate, mostly asymmetric, margin
serrate to denticulate for its entire length, apex acute-acuminate; midrib and lateral veins strongly
raised beneath, impressed above, lateral veins 58 pairs, intercostal veins subscalariform to tessellate,
sometimes strongly raised and distinct beneath; petioles (7)1015(20) mm long, 12 mm thick,
densely short-pubescent. Inflorescences male and female on separate twigs, much-branched panicles
or thyrses, lax or condensed at anthesis, axes stout, 12 mm thick; bracts ovate-acute, 23 x1 mm;
male inflorescences 35 cm long, 20100-flowered; female inflorescences to 4 cm long, (3)515(
20)-flowered. Flowers: males 1.52 mm diameter, perianth lobes ciliate, 1.52 x1 mm, filaments 1
1.5 mm, anthers c. 1 x0.5 mm, pistillodes obovoid-conical, compressed, 0.51 x0.5 mm; females 2
3 x12 mm, perianth lobes ovate-acute, 11.5 x0.5 mm, ciliate and densely short-pubescent,
glabrescent, staminodes absent, ovary ovoid-conical, c. 2 x1 mm, stigmatic arms slender, 11.5 mm
long, spreading. Fruits ovoid, 35 x24 mm, turning to deep purple or black when ripe, glabrous;
stalks to 3 mm long. Seeds with scanty to copious endosperm.
Vernacular names. Sabahhalindagong, randagong (Dusun Ranau).
Distribution. Widely distributed from India (W Himalayas to Bombay and Malabar) to Myanmar,
Thailand, Indo-China, China, S Japan, through Malesia to Queensland, Melanesia, Micronesia, and
Polynesia. In Malesia, the species is found in Sumatra, Peninsular Malaysia, Java, Borneo,
Philippines, Sulawesi, Maluku, Lesser Sunda Islands, and New Guinea. In Borneo, so far known from
a few collections from Keningau (SAN 85756), Kunak (SAN 89965), Ranau (Mt. Kinabalu, Chew,
Corner & Stainton RSNB 2586; SAN 116596), Sandakan (Kadir A. 932), and Tambunan (SAN 59469)
districts, in Sabah.
Ecology. In Sabah, scattered in secondary forest, road sides, and other newly cleared areas in the hills
and lower montane areas, from sea level to 1500 m.
4. Trema tomentosa (Roxb.) Hara
Fig. 4.
(Latin, tomentosus = covered with matted woolly or short hairs; the young parts of the plants, inflorescences and
fruits)
400
Fig. 4. Trema tomentosa. A, fruiting leafy twig; B, lower leaf surface; C, upper leaf surface; D, mature fruit; E, basal view of
mature fruit; F, mature fruit with the pericarp halfway removed to show the endocarp; G, mature fruit in longitudinal section;
H, female flower in longitudinal section; IK, male flower buds; L, open male flower. (AG from SAN 35146, H from
Cuadra A 259, IL from SAN 107230.)
401

Fl. E. Himal. 2 (1971) 19; Soepadmo I.e. (1973) 423, I.e. (1977) 53; Corner I.e. 739; Whitmore, Tantra & Sutisna
I.e. 363; Kessler & Sidiyasa I.e. 231. Basionym: Celtis tomentosa Roxb., Fl. Ind. ed. Carey 2 (1832) 66. Type:
Roxburgh, s.n., "Native to Chittagong" (n.v.; no drawing at BM or K). Synonyms: Sponia tomentosa (Roxb.)
Planch. I.e. 336; Sponia velutina Planch. I.e. 327, p.p., excl. specim. Cuming 1232 ex Luzon; Trema velutina
(Planch.) Blume I.e. (1856) 58.
Shrub to medium-sized tree, (3-)5-15(-24) m tall, (5-)10-30(-50) cm diameter. Bark greybrown, smooth to finely fissured, lenticellate; inner bark yellowish green. Young parts,
inflorescences, petioles, and stipules densely and thickly tomentose with grey-brown, erect,
velvety simple hairs. Stipules linear-lanceolate, 2-6 x 1-1.5 mm. Leaves thick-coriaceous, rarely
thin-coriaceous, more or less concolorous, drying dark-chocolate brown to blackish brown,
upper surface strongly scabrate, lower surface densely velvety tomentose with grey-brown to
glaucous simple hairs (lamina surface between hairs visible even under low magnification);
broadly ovate to ovate-elliptic, (5-)8-15(-20) x (2-)4-7(-9) cm, length 2.5-3 times the width,
broadest mostly below the middle; base cordate, subcordate, rarely rounded or truncate, mostly
strongly asymmetric, rarely symmetric, margin serrate throughout, apex acute to acuminatecaudate, acumen sharp, 0.5-3 cm long; midrib and lateral veins strongly raised beneath,
impressed above, lateral veins 5-6 pairs, subparallel, ascending, intercostal veins
subscalariform to tessellate, often rather distinct beneath; petioles 10-15 mm long, 1-2 mm thick.
Inflorescences male, female, or mixed, axes 1-2 mm thick; bracts ovate-acute, c. 1 x 0.5 mm;
male and mixed inflorescences 3-4.5 (-6) cm long, lax, 10-100-flowered; female inflorescences
to 5 cm long, axes stout, 1-2 mm thick, 5-15-fTowered. Flowers: males 1-2 mm diameter,
perianth lobes mostly 5, elliptic, c. 1.5 x 1 mm, filaments c. 1 mm long, flat, glabrous, anthers
c. 1 x 0.5 mm, pistillodes obovoid-ellipsoid, compressed, 1-1.5 x 0.5 mm; females c. 2 x 1 mm,
perianth lobes 4-5, ovate-acute, c. 1 x 0.5 mm, staminodes absent or strongly rudimentary,
ovary c. 1.5 x 0.5-1 mm, stigmatic arms slender, c. 1 mm long, spreading. Fruits ovoid, c . 3 x 2
mm, turning to black when ripe, glabrous. Seeds with copious endosperm.
Vernacular names. Sabahbintanong (Murut), balik-balik angin (Bajau Tuaran), lindagong
(Malay), lundagong (Dusun), nasi-nasi (Malay), ondogon (Dusun Kimanis), randagong
(Dusun Kinabatangan, Dusun Labuk, Dusun Ranau), rondongon (Dusun Kinabatangan),
rondogong (Dusun Bundu Tuhan), tikam (Dusun Sungei). Sarawakkereneong (Iban),
lindagong (Kedayan), marong (Malay), murieng (Bidayuh), tuku baroh (Bidayuh).
Distribution. East tropical Africa, Madagascar, Pakistan, India, Bangladesh, Myanmar,
Thailand, Indo-China, China, Ryu Kyu Is., throughout Malesia to Queensland, New
Caledonia, Micronesia, and Polynesia. Common in Sabah and Sarawak; also known for Brunei and
Kalimantan.
Ecology. Common pioneer shrubs or trees in the lowlands and hills, from sea level to 1000 m, in
newly cleared areas on all types of soil, including limestone.
Taxonomy. In the past, this species has been wrongly identified and described as T. orientalis
(Merrill I.e., 1921, I.e., 1929; Masamune I.e.; Brown I.e.; Anderson I.e.). However, though T.
tomentosa and T. orientalis are closely allied, they can be distinguished easily by the
characters given in the key.
402
WINTERACEAE
K.M. Wong
Sandakan, Malaysia
A.C. Smith, J. Arn. Arb. 24 (1943) 1, 119; Vink, Blumea 18 (1970) 225; Anderson, CLTS (1980) 346,
excl. Illicium.
Shrubs and trees, rarely epiphytes. Wood without true xylem vessels. Leaves simple, entire,
spirally arranged, typically with pale translucent dots on the lower side, often with aromatic
tissue; stipules none. Inflorescence axillary or terminal, of a solitary flower or several flowers
spirally and closely arranged on a very short flowering shoot, or a condensed shoot with 12
orders of very short branches bearing the flowers; developing in the axil of a scale-like bract
or leaf. Flowers bisexual, unisexual (and then plants dioecious), or unisexual and bisexual
together (plants polygamo-dioecious); sepals 26, valvate to spirally inserted, overlapping and
slightly twisted, or basally fused and cup-like or nearly completely fused and hood-like and
rupturing into 2 or more parts; petals none, or 2many, in whorls of (2)3 or in spirals, free;
stamens 5numerous (more than 100) in 1several whorls, filaments club-shaped and slightly
flattened or tapering, anthers opening by slits; ovary superior, of 1many carpels in 1 whorl,
carpels free or fused or sometimes initially connate and later separating, each a conduplicate
(lengthwise folded) structure with 2 stigmatic crests forming a ventral pair of ridges or a
completely closed structure with a terminal style and stigma; ovules 1 or several to many
(and then in 2 rows (laminar placentation) within the carpel), anatropous. Fruit berry-like
or a cluster of follicles developed from free carpels, or a multilocular capsule or syncarp
developed from fused carpels. Seeds with copious endosperm; embryo minute.
Distribution. About 5 genera, c. 60 species, Old and New World but principally southern
hemisphere. Drimys (represented by section Drimys) is the only genus extending to the New
World (S Mexico to Cape Horn) and Takhtajania is a monotypic genus of Madagascar. Borneo,
the Philippines, Flores, Sulawesi, Maluku, New Guinea, the Solomons, New Caledonia; also
eastern Australia, Tasmania and New Zealand. In Borneo (all territories), Drimys is the sole
genus.
Taxonomy. In some older classifications, the Winteraceae (often together with Illicium)
were regarded as a part of the Magnoliaceae, but differ in having oil-glands in the leaf
tissue, its comparatively small floral axis and the lack of stipules. Illicium is now removed
to its own family, Illiciaceae (Volume 1), on account of, among others, its differing wood
structure, nodal anatomy, carpellary structure and cytology (Smith l.c.). Notwithstanding, the
Winteraceae, Annonaceae, Degeneriaceae, Magnoliaceae and Myristicaceae are generally
accepted as a closely related alliance of families thatalong with other families such as
the Illiciaceae and Schisandraceaeretain some of the most primitive (often also termed
403
ranalean) characteristics among flowering plants. The lack of xylem vessels in the wood and
presence of conduplicate carpels among members of the Winteraceae make this family one of
the most primitive in this sense. Indeed, Bailey & Swamy (Amer. J. Bot. 38 (1951) 373) have
remarked that the least modified [= advanced] form of surviving carpel occurs in species of
Drimys, including D. piperita.
DRIMYS J.R. Forst. & G. Forst.
(Greek, drimus = acrid; the bitter taste of the bark)

Char. Gen. 42 (1776) 83; Merrill, EB (1921) 252 (as part of Magnoliaceae); Masamune, EPB (1942) 277
(as part of Magnoliaceae); A.C. Smith l.c. 122; Vink l.c. 302; Anderson l.c. 346. Synonym: Tasmannia
R. Br. ex DC., Syst. 1 (1817) 445.
Shrubs, trees or rarely epiphytes. Leaves glabrous. Inflorescence axillary, with (in section
Drimys) or without distinct peduncles, with 1many flowers. Flowers usually unisexual, very
rarely bisexual and then together with unisexual flowers on the same plant, or (in section
Drimys) always bisexual; calyx a hood-like structure enclosing the flower bud, rupturing and
falling as the flower opens; petals none or 225, free; stamens 7numerous (more than 100),
filaments thick and tapering; ovary of 124 carpels, carpels free, globular to ovoid, slightly
laterally flattened, sessile or short-stalked, each a conduplicate structure with 2 stigmatic
crests forming a ventral pair of ridges along the carpel length; ovules several to many, in 2
rows within the carpel. Fruit berry-like or a cluster of follicles developed from free carpels.
Seeds straight to strongly curved.
Distribution. Nine species, of which four (in section Drimys) in S America and five (in
section Tasmannia) in Malesia, Australia, New Zealand and the SW Pacific. In Borneo
(including Sabah and Sarawak), one species.
Uses. The most famous product of the genus is Winters bark, from the S American D. winteri,
used in 1578 by Capt. William Winter against scurvy and as a spice. In Brazil, bark of this
species is used as an astringent and a stimulant. No recorded uses in Sabah and Sarawak.
Taxonomy. Only two sections are recognised for the genus. Section Drimys (S American)
has distinct inflorescence peduncles, bisexual flowers and a base chromosomal number of 43,
whereas section Tasmannia (Old World) has indistinct peduncles, typically unisexual flowers
and a base chromosomal number of 13. Very many species recognised by A.C. Smith l.c. 122,
especially from New Guinea, have been reduced to synonymy under D. piperita by Vink l.c.,
who considers the species as a widespread Malesian-Australasian species with at least 39 local
entities often without any sharply defined taxonomic boundaries.
Drimys piperita Hook. f.
Fig. 1
(Latin, piperita = pepper-flavoured; the aromatic bark and leaf tissue)

Icon. Pl. 9 (1852) pl. 896; Beccari, Malesia 1 (1877) 185; Stapf, FMK (1894) 128; Merrill l.c. 252;
Masamune l.c. 277; A.C. Smith l.c. 137; Vink l.c. 314; Anderson l.c. 346. Type: Low, s.n., Borneo, Mt.
404
WINTERACEAE (WONG)
Fig. 1. Drimys piperita. A, leafy twig with male flowers; B, young male flower; C, mature male flower;
D, leafy twig with female flowers; E, mature female flower; F, leafy twig with fruits; G, twig with mature
fruits; H, lateral view of fruit (left) and section through carpel showing seeds; J, detail of lower leaf
surface. (A, B & J from Chew et al. 1056, C from Ding Hou 249, D & E from SAN 29058, F from SAN
57540, G from SAN 123345, H from SAN 41271.)
405
Kinabalu (holotype K; isotype UC). Synonym: Tasmannia piperita (Hook. f.) Miers, Ann. Mag. Nat.
Hist. 3, 2 (1858) 110.
Shrub less than 1 m high or tree to 12 m high and 25 cm diameter, usually a small tree 24
m high, rarely an epiphyte. Bark smooth, dark brown; inner bark pale brown, thin, aromatic
when bruised. Sapwood pale yellow to white. Leaves reddish when young, chartaceous to
coriaceous; glabrous on both sides; occasionally very slightly to conspicuously glaucous,
with scattered dark glands on the lower side; aromatic when crushed; lanceolate to elliptic
or obovate, 215 14.5 cm, base usually cuneate and slightly decurrent on the petiole to
obtuse, apex acute to acuminate; midrib slightly grooved to flat or slightly prominent on
the upper side, prominent on the lower side, often reddish in fresh specimens, lateral veins
(6)1014 pairs, immersed in the blade in fresh material, flat to slightly prominent on both
sides in dried specimens, branching conspicuously towards the margin, sometimes reddish
in fresh material, intercostal veins loosely to densely reticulate, immersed in the blade in
fresh material, slightly prominent on both sides in dried specimens; petiole (3)725 mm
long, shallowly grooved on the upper side; Inflorescence of 13(5) flowers on a very short
inconspicuous axis and resembling a fascicle of flowers, in the axils of reduced leaves (scaleleaves) at the proximal part of a new episode of shoot extension or a new axillary shoot, very
rarely in the axils of normal leaves, scale-leaves triangular to ovate, papery, 312 16 mm,
caducous. Flowers unisexual, very rarely bisexual and then occurring with other unisexual
flowers; pedicels (5)1040(60) mm long; calyx hood-like, 212 mm long, rupturing into 2
(sometimes more) parts as the flower opens, pale green with reddish apex, caducous; petals
(2)512(15), narrowly blunt-elliptic to oblanceolate, 418 1.56 mm, white, spreading
to strongly folded back in the open flower; stamens in male flower (7)2560 or more in
several series, filaments slightly flattened, tapering, 15 mm long, white and pink-tinged,
anthers 0.51 mm long, yellow (in female flowers: stamens none); carpels in female flowers
(2)312(15), obovoid, 22.5 mm long, short-stalked, pale green, stigmatic crest pale brown
and extending nearly the entire carpel length (in male flowers: carpels none or 13 and sterile,
1.52 mm long); ovules (2)1230(46) in fertile carpels, none or degenerate in sterile carpels.
Fruit of (1)312(15) follicles; follicles globular to blunt-ellipsoid to obovoid, 515 38
mm, each marked by a dark brown longitudinal thickening (the remains of the stigmatic crest),
short-stalked, pale green ripening reddish and then purple-black. Seeds many per follicle,
narrowly blunt-ellipsoid, slightly curved, 24 13 mm, dark brown to black.
Distribution. Philippines, Borneo, Flores, Sulawesi, Maluku, New Guinea, eastern Australia.
In Borneo in all territories (Sabah, Sarawak, Brunei, Kalimantan). This is the only Asiatic
species of the genus known outside of New Guinea and Australia.
Ecology. Montane forests, mostly at 10003000 m, from around 700 m on coastal mountains
(e.g., Mt. Silam in Sabah) and up to 3800 m on Mt. Kinabalu in Sabah; locally frequent, also
on exposed ridges. Occurring on sandstone, granite, ultramafic rocks (in Sabah), dacite and
limestone (in Sarawak).
Notes. The inflorescences are produced in the axils of reduced (scale) leaves that mark the
beginning of each new episode of stem elongation or a new axillary branch shoot; these scale
leaves are arranged closely together and contrast with the normal leaves produced higher up
together with internodal elongation, which are more distantly spaced. It is not known whether
each episode of new shoot growth corresponds to a definite growth season during a year, but
406
WINTERACEAE (WONG)
the presence of some exceedingly short leafy sections on some stems, and the occurrence of
both flowering and non-flowering individuals in most populations seen at various times of the
year, suggest that both episodic shoot growth and flowering occur intermittently throughout
the year in Borneo.
407
408
ABBREVIATIONS OF
Anderson, J.A.R. 1980. A Checklist of the Trees of Sarawak. Forestry Dept., Sarawak.
CLTS
Ashton, P.S. 1988. Manual of the Non-Dipterocarp Trees of Sarawak. Forestry Dept.,
Sarawak.
MNDTS
the Netherlands.
FJ
Browne, F.G. 1955. Forest Trees of Sarawak and Brunei. Govt. Press, Sarawak.
FTSB
TBS
Lumpur.
EPMP
TS
Corner, E.J.H. 1988. Wayside Trees of Malaya. 2 Vols. Malayan Nature Society, Kuala
Lumpur.
WSTM
Flora Malesiana
FM
TNB
Kuala Lumpur.
OFMSP
PB
EPB
409
Merrill, E.D. 1921. A Bibliographic Enumeration of Bornean Plants. J. Str. Br. Roy. As. Soc.,
Sp. No.
EB
Merrill, E.D. 1929. Plantae Elmerianae Borneenses. University of California Publications in
Botany, Vol. 15. University of California Press, Berkeley.
PEB
Ng, F.S.P. (ed.). 1978 & 1989. Tree Flora of Malaya, Vols. 3 & 4. Longman, Malaysia.
TFM
Ng, F.S.P. 1991 & 1992. Manual of Forest Fruits, Seeds and Seedlings. Malaysian Forest
Records, No. 34. Vols. 1 & 2. FRIM, Malaysia.
MFFSS
MPESA
Plant Resources of South East Asia. Wageningen.(Various authors and volumes).
(PROSEA, to be listed without editor or author names, but to include volume and page
numbers).
Ridley, H.N. 1923-1925. The Flora of the Malay Peninsula, Vols. 1-5. Reeves, London.
FMP
CST
Stapf, O. 1894. On the Flora of Mount Kinabalu, in North Borneo. Transactions of the
Linnean Society, London, Ser. 2, 4, pp. 69-263.
FMK
Whitmore, T.C. (ed.). 1972 & 1973. Tree Flora of Malaya, Vols. 1 & 2. Longman,
Malaysia.
TFM
CLS
Whitmore, T.C., I.G.M. Tantra & U. Sutisna. 1990. Tree Flora of Indonesia. Check List for
Kalimantan, Part 1 & Parts 2.1-2.2. Forest Research & Development Centre, Bogor.
CLK
Willis, J.C. 1973. A Dictionary of the Flowering Plants & Ferns, revised by H.K. Airy Shaw.
Cambridge University Press, Cambridge.
DFPF
Wong, T.M. 1982. A Dictionary of Malaysian Timbers. Malayan Forest Records No. 30.
Forest Research Institute, Kepong, Kuala Lumpur.
DMT
410
COMMONLY USED
Malay
English
Word
Abbreviation
Word
Abbreviation
Central
Bukit
Bt.
Division
Div.
Gunung
G.
East
Kampung
Kg.
Forest Reserve
FR
Sungai
Sg.
Island
Is.
Tanjung
Tg.
Mount
Mt.
National Park
NP
North-East
NE
North-West
NW
River
R.
South
South-East
SE
South-West
SW
West
411
412

(compiled by R.C.K. Chung)
Names in italics here refer to synonyms; family names given in upper case are those of families revised
in this volume. Pages with illustrations are listed in italics.
Abelmoschus 231
Abelmoschus esculentus 233, 234
Abelmoschus moschatus 234
Abutilon 235
Abutilon indicum subsp. indicum 235
Aglaia 3
Alectryon 264, 267, 270, 271
Alectryon glaber 271, 272
Allantospermum 175
Allocasuarina 119
Allophylus 264, 266, 268, 273
Allophylus cobbe 274, 275, 276
Allophylus glaber 274
Allophylus javensis 274
Allophylus obliquus 274
Allophylus racemosus 274
Allophylus scandens 274
Allophylus sumatranus 274
Allophylus ternatus 274
Allophylus timorensis 274
Alzatea 136
Alzateaceae 136
Ammannia 225
ANACARDIACEAE 1, 2, 3
Anacardium 2
Anacardium occidentale 2
Anauxanopetalum schwenkii 92
Androtium 1, 3, 4
Androtium astylum 5, 6
Aneulophus 167
Annonaceae 403
Anthusa 93
Antonia griffithii 221
Aphananthe 384
Aphananthe cuspidata 385, 386
Aphania 309
Aphania dasypetala 324
Argusia 94, 95
Argusia argentea 95, 96
Argusia gnaphalodes 95
Argusia sibirica 95
Arytera 263, 264, 268, 270, 276
413
Arytera litoralis 277, 278

Ascarina 129, 130, 131
Ascarina philippinensis 129, 132, 133
Ascarina reticulata 133
Axinandra 136, 137
Axinandra sect. Naxiandra 136
Axinandra alata 137
Axinandra beccariana 137, 138
Axinandra borneensis 138
Axinandra coriacea 137, 138, 139
Axinandra maingayi 138
Barringtonia 239, 243
Barringtonia conoidea 29
Blattiaceae 136
Boerhavia 251, 252
Boerhavia chinensis 252
Boerhavia diffusa 252
Boerhavia erecta 252
Bombacaceae 234
BORAGINACEAE 93
Borago 93
Bouea 1, 2, 3, 5, 7
Bouea angustifolia
7
Bouea macrophylla 7
Bouea microphylla 7
Bouea oppositifolia 7, 8
Bougainvillea 251, 252
Bougainvillea glabra 252
Bougainvillea spectabilis 252
Buchanania 1, 3, 5, 9
Buchanania arborescens 9, 10, 11
Buchanania auriculata 14
Buchanania florida var. lucida 10
Buchanania fragrans 13
Buchanania glaberrima
10
Buchanania insignis 10
Buchanania lucida 10
Buchanania macrophylla 16
Buchanania sessifolia 9, 10, 11, 12
Buchanania splendens 9, 13
Buddleja 189
Buddlejaceae 189
Burseraceae 2, 3
Camelliaceae 379
Campnosperma 2, 3, 13, 14
Campnosperma auriculatum 14, 15
Campnosperma coriaceum 2, 14
Campnosperma macrophylla 16
414
Campnosperma squamatum 14, 15, 16

Canarium 3
Caniogeton arborescens 10
CAPRIFOLIACEAE 107
Cardiospermum 264, 265, 268
Cardiospermum halicacabum 265
Casuarina 119, 120
Casuarina equisetifolia 119, 120, 121, 122
Casuarina nobilis 125
Casuarina sumatrana 127
CASUARINACEAE 119
Celtis 383, 384, 385, 387
Celtis amboinensis 399
Celtis cinnamomea 390
Celtis collinsae 388
Celtis crenato-serrata 390
Celtis hamata 390
Celtis mindanaensis 388
Celtis orientalis 399
Celtis philippensis 387, 389
Celtis reticulosa 390
Celtis rigida 399
Celtis timorensis 387, 389, 390
Celtis tomentosa 402
Celtis waitzii 390
Celtis wightii 388
Ceodes umbellifera 255
Cerinthe 93
Ceuthostoma 119, 120, 121
Ceuthostoma terminale 123, 124
CHLORANTHACEAE 129
Chloranthales 129
Chloranthus 129, 130
Chloranthus erectus 131
Clusiaceae 7
Cnemidiscus thorelii 293
Coelopyrum coriaceum 14
Cordia 93, 94, 97, 98
Cordia cylindristachya 97, 98
Cordia dichotoma 97, 98, 99
Cordia fragrantissima 98
Cordia gerascanthus 97
Cordia griffithii 98
Cordia premnifolia 98
Cordia rumphii 100
Cordia suaveolens 98
Cordia subcordata
97, 98, 100
Crossonephelis 291
415
Crossonephelis palawanicus 293

Crossonephelis philippinensis 292
Crossonephelis thorelii 293
Crypteronia 135, 136, 140, 141
Crypteronia borneensis 141
Crypteronia elegans 141, 142
Crypteronia glabriflora 141, 143, 144
Crypteronia griffithii 141, 143
Crypteronia macrophylla 141, 145
Crypteronia paniculata var. paniculata 141, 146
CRYPTERONIACEAE 135, 136, 225
Ctenolophon 151
Ctenolophon grandifolius 153
Ctenolophon parvifolius 151, 152
CTENOLOPHONACEAE 151, 175
Cubilia 264, 266, 267, 269, 279
Cubilia cubili 279, 280
Cupania acuta 363
Cupania diplopetala 298
Cupania pleuropteris 296, 300
Cyclostemon cuspidatum 385
Cynoctonum 190
Cynoglossum 93, 94
Cynoglossum lanceolatum 95
Cyrillopsis 175
Cyrtophyllum peregrinum 206
Cyrtophyllum speciosum 205
Dactylocladus 135, 136, 147
Dactylocladus stenostachys 147, 148
DAPHNIPHYLLACEAE 155
Daphniphyllales 155
Daphniphyllum 155, 156
Daphniphyllum bancanum 156
Daphniphyllum borneense 158
Daphniphyllum glaucescens 156
Daphniphyllum glaucescens subsp. borneense 157, 158
Daphniphyllum glaucescens subsp. dichotomum 158
Daphniphyllum laurinum 156
Degeneriaceae 403
Dendromyza 258
Dendromyza reinwardtiana 258
Dendrotrophe 258
Dictyoneura 263, 264, 266, 270, 281
Dictyoneura acuminata subsp. acuminata 282, 283
Dimocarpus 264, 266, 266, 269, 270, 282, 284, 357
Dimocarpus dentatus 284, 285
Dimocarpus fumatus subsp. fumatus 286
Dimocarpus fumatus 284, 285, 286
416
Dimocarpus longan 264, 284, 287

Dimocarpus longan subsp. longan var. longan 285, 288
Dimocarpus longan subsp. malesianus var. echinatus 285, 288, 350
Dimocarpus longan subsp. malesianus var. malesianus 285, 288
Dipsacales 107
Dodonaea 264, 265, 268
Dodonaea ?lamponga 103
Dodonaea viscosa 265
Dracontomelon 2, 4, 17
Dracontomelon brachyphyllum 18
Dracontomelon costatum 17
Dracontomelon dao 17, 18, 19
Dracontomelon edule 18
Dracontomelon edulis 18
Dracontomelon mangiferum 18
Dracontomelon sylvestre 18
Drimycarpus 1, 2, 3, 20
Drimycarpus luridus 20, 21
Drimycarpus maximus 20, 21, 22
Drimys 403, 404
Drimys sect. Drimys 404
Drimys sect. Tasmannia 404
Drimys piperita 404, 405
Drimys winteri 404
Echium 93
Ehretia 94, 101
Ehretia laevis var. timorensis 101
Ehretia timorensis 101, 102
EPACRIDACEAE 159
Equisetum 121
Ericaceae 159
Erioglossum 309
Erioglossum rubiginosum 322
ERYTHROXYLACEAE 167
Erythroxylum 167, 168, 169
Erythroxylum bancanum 170
Erythroxylum borneense 170
Erythroxylum burmanicum 170
Erythroxylum coca 168
Erythroxylum cuneatum 169, 171
Erythroxylum cuneatum forma cuneatum 170, 172
Erythroxylum cuneatum forma sumatranum 170, 172
Erythroxylum cuneatum var. bancanum 170
Erythroxylum densinerve 170
Erythroxylum iwahigense forma calcicola 169, 173
Erythroxylum iwahigense forma iwahigense 169
Erythroxylum latifolium 169, 171, 172, 174
Erythroxylum latifolium var. angustatum 171
417
Erythroxylum latifolium var. longipetiolatum 171

Erythroxylum longistipulatum 170
Erythroxylum novogranatense 168
Erythroxylum platyphyllum 170
Erythroxylum sarawakanum 169, 171, 173, 174
Erythroxylum sumatranum 170
Eugenia 7
Euodia 78
Euphorbiaceae 155
Euphoria cinerea 288
Euphoria cubili 279
Euphoria echinulata 288
Euphoria elongata 288
Euphoria malaiensis 288
Euphoria microcarpa 288
Euphoria nephelioides 288
Euphoria noronhiana 371
Euphoria sclerocarpa 288
Euphoria stellata 288
Euphoria succulenta 288
Exocarpos 258
Exocarpos latifolius
259, 260
Fagraea 189, 190, 191, 192
Fagraea sect. Cyrtophyllum
Fagraea sect. Cyrtophyllum 191, 192
Fagraea sect. Fagraea 191, 192
Fagraea sect. Racemosae 191, 192
Fagraea acuminatissima 196
Fagraea acutibracteata 195
Fagraea appendiculata 217
Fagraea aurantiodora 205
Fagraea auriculata 192, 194
Fagraea auriculata subsp. borneensis 202
Fagraea belukar 199, 200, 201
Fagraea blumei 192, 195, 197, 202, 206
Fagraea borneensis 194, 197, 202
Fagraea carnosa 196
Fagraea caudata 199, 203
Fagraea ceilanica 192, 196, 197, 209
Fagraea coarctata 217
Fagraea cochinchinensis 206
Fagraea collina 200, 203
Fagraea congestiflora 214
Fagraea cordifolia 217
Fagraea crassipes 204
Fagraea crenulata 191, 192
Fagraea cuspidata 198, 204
Fagraea cymosa 204
418
Fagraea dulitensis 196, 205

Fagraea elliptica 192, 200, 203, 205, 212
Fagraea eucalyptifolia 214
Fagraea floribunda 192, 194, 206, 207
Fagraea fragrans 192, 199, 203, 206, 208
Fagraea gardenioides 192
Fagraea gardenioides subsp. borneensis 197
Fagraea gigantea 199, 208
Fagraea gracilis 217
Fagraea grandifolia 217
Fagraea havilandii 197
Fagraea heterophylla 196
Fagraea iliasii 194
Fagraea involucrata 193, 194
Fagraea involucrata var. longipetiolata 193, 194
Fagraea javensis 205
Fagraea kalimantanensis 192
Fagraea kimangu 205
Fagraea kinabaluensis 196, 209, 210
Fagraea kuminii 195
Fagraea latifolia 217
Fagraea ligustrina 217
Fagraea ligustrina var. disparifolia 217
Fagraea littoralis 198
Fagraea littoralis var. amboinensis 198
Fagraea littoralis var. borneensis 198
Fagraea littoralis var. forstenii 198
Fagraea longipetiolata 198
Fagraea macroscypha 193, 194
Fagraea megalantha 193
Fagraea minor 209
Fagraea montana 199, 209
Fagraea morindaefolia 217
Fagraea nonok 202
Fagraea oblonga 198
Fagraea obovata 202
Fagraea obovata var. brevicalyx 202
Fagraea obovato-javana 202
Fagraea obovato-javana var. bebeak 202
Fagraea oreophila 197
Fagraea pauciflora 217
Fagraea pendula 204
Fagraea peregrina 206
Fagraea philippinensis 199, 211
Fagraea picrophloea 205
Fagraea pseudoelliptica 205
Fagraea pusilliflora 205
Fagraea racemosa 192, 204, 209, 211, 214, 215, 217
419
Fagraea racemosa var. pauciflora 217

Fagraea rarissima 197
Fagraea renae 195
Fagraea resinosa 193, 195, 202, 211
Fagraea ridleyi 195
Fagraea robusta 204
Fagraea rodatzii 217
Fagraea rostrata 196
Fagraea rugulosa 200, 212, 213
Fagraea scholaris 217
Fagraea sororia 208
Fagraea speciosa 205, 208
Fagraea spicata 198, 214
Fagraea splendens 196
Fagraea stenophylla 199, 214
Fagraea stonei 196
Fagraea subreticulata 217
Fagraea sumatrana 205
Fagraea teysmannii 199, 215
Fagraea tuyukii 197
Fagraea uniflora 194, 196
Fagraea vaginata 202
Fagraea valida 205
Fagraea volubilis 199, 215
Fagraea volubilis var microcalyx 217, 216
Fagraea volubilis var. volubilis 217
Fagraea sp. A 193
Ficus cuneata 169
Flacourtia rukam 378
Flacourtiaceae 375, 378
Galumpita 384
Galumpita cuspidata 385
Ganophyllum 264, 266, 269, 289
Ganophyllum falcatum 289, 290
Garcinia 7
Garuga 3
Gelsemium 189, 190, 191
Gelsemium elegans 191
Geniostoma 189, 190, 191, 218
Geniostoma ancilimbum 218
Geniostoma avene 218
Geniostoma batanense 218
Geniostoma brevipes 218
Geniostoma celebicum 218
Geniostoma cumingianum 218
Geniostoma fasciculata 218
Geniostoma lasiostemon 218
Geniostoma laxa 218
420
Geniostoma longipes 218

Geniostoma micranthum 218
Geniostoma mindanaense 218
Geniostoma miquelianum 218
Geniostoma moluccanum 218
Geniostoma montanum 218
Geniostoma oblongifolium 218
Geniostoma pachyphyllum 218
Geniostoma philippinensis 218
Geniostoma ramosii 218
Geniostoma rupestre 218, 219
Geniostoma stenophyllum 218
Gentianales 190
Gentianaceae 190
Gironniera 384, 390, 391
Gironniera subgen. Galumpita 384
Gironniera curranii 385
Gironniera cuspidata 385
Gironniera hirta 392
Gironniera lucida 385
Gironniera nervosa 391, 392, 393, 394, 396
Gironniera nervosa var. subaequalis 394
Gironniera nitida 385
Gironniera parvifolia 392, 393
Gironniera paucinervia 393
Gironniera penangiana 392
Gironniera reticulata 385
Gironniera scabrida 393
Gironniera sponioides 392
Gironniera subaequalis 392, 394, 395
Gironniera subaequalis var. ceylanica 393
Gironniera zeylanica 393
Glenniea 264, 266, 270, 291, 292
Glenniea philippinensis 292
Glenniea thorelii
293, 292, 294
Gluta 1, 2, 3, 22, 23
Gluta aptera 23, 24, 25, 26
Gluta beccarii 23, 24, 26
Gluta laccifera 2
Gluta laxiflora 23, 24, 26
Gluta macrocarpa 24, 26
Gluta oba 23, 24, 27
Gluta rugulosa 23, 24, 27
Gluta sabahana 22, 23, 24, 25, 28
Gluta speciosa 23, 24, 28
Gluta usitata 2
Gluta velutina 23, 24, 29
Gluta wallichii 23, 24, 25, 29
421
Gossypium 233, 235

Gossypium barbadense var. acuminatum 235
Gouphia laurina 156
Guioa 264, 267, 270, 295
Guioa bijuga 295, 296, 297
Guioa bullata 298
Guioa diplopetala 296, 298, 299
Guioa diplopetala forma borneensis 298
Guioa diplopetala var. borneensis 298
Guioa koelreuteria 296, 300
Guioa pleuropteris 296, 300, 301
Guioa pleuropteris var. bijuga 296
Guioa pterorhachis 296, 303
Guioa pubescens 296, 297, 303
Guioa regularis 298
Guioa rubrofusca 296
Gymnostoma 119, 120, 123, 125
Gymnostoma nobile 125
Gymnostoma sumatranum 125, 126, 127
Hamamelidaceae 155
Harpullia 263, 264, 266, 270, 304, 305
Harpullia arborea 305, 306
Harpullia confusa 307
Harpullia cupanioides 305, 306, 307
Harpullia tomentosa 307
Havilandia 94
Hebecoccus falcatus 317
Hedyachras 291
Hedyachras philippinensis 292
Hedyosmum 129, 130
Hedyosmum orientale 130
Heliotropium 94
Heliotropium indicum
94
Helminthospermum 391
Helminthospermum scabridum 393
Henslowia 258
Hibiscus 233, 234, 236
Hibiscus archeri 233
Hibiscus borneensis 236, 237
Hibiscus decaspermus 238
Hibiscus floccosus 238
Hibiscus macrophyllus 236, 237, 238
Hibiscus populneus 241
Hibiscus radiatus 237
Hibiscus rosa-sinensis 233
Hibiscus surattensis 236
Hibiscus tiliaceus 236, 237, 240, 241, 243
Hibiscus tiliaceus subsp. tiliaceus 239
422
Homoioceltis 384
Icica timorensis
69
Illiciaceae 403
Illicium 403
Irina 356
Irina alnifolia 357
Irina tomentosa 357
IXONANTHACEAE
151, 175
Ixonanthes 175, 176
Ixonanthes sect. Brewstera 176
Ixonanthes sect. Ixonanthes 176
Ixonanthes beccarii 177
Ixonanthes crassifolia 177
Ixonanthes grandiflora 177
Ixonanthes icosandra 176
Ixonanthes multiflora 177
Ixonanthes petiolaris 176, 177, 178
Ixonanthes philippinensis 177
Ixonanthes reticulata 176, 177, 179
Juglandales 245
Koordersiodendron 2, 4, 30
Koordersiodendron pinnatum 30, 31
Lagerstroemia 226, 227
Lagerstroemia borneensis 226
Lagerstroemia crassifolia 228
Lagerstroemia floridunda 226, 227
Lagerstroemia flos-reginae 230
Lagerstroemia indica 226, 227
Lagerstroemia piriformis 227, 228
Lagerstroemia pustulata 227, 228, 229
Lagerstroemia speciosa 227, 230
Lawsonia 226
Lawsonia inermis 226
Leea 181, 182
Leea aculeata 182, 183, 184
Leea aequata 182, 183
Leea amabilis 182
Leea angulata 182, 183, 185
Leea guineensis 182, 186
Leea indica 183, 186
Leea javanica 186
Leea robusta 186
Leea rubra 182
Leea sandakanensis 183
Leea sundaica 186
LEEACEAE 181
Lepisanthes 264, 265, 266, 267, 268, 269, 270, 308, 309, 310
Lepisanthes subgen. Aphania 309
423
Lepisanthes subgen. Erioglossum 309

Lepisanthes subgen. Lepisanthes 309
Lepisanthes subgen. Lepisanthes sect. Hebecoccus 309
Lepisanthes subgen. Lepisanthes sect. Lepisanthes 309
Lepisanthes subgen. Otophora 309
Lepisanthes subgen. Otophora sect. Anomotophora 309
Lepisanthes subgen. Otophora sect. Otophora 309
Lepisanthes subgen. Otophora sect. Pseudotophora 309
Lepisanthes alata 309, 310, 311, 312, 313
Lepisanthes amoena 309, 310, 312, 313, 314
Lepisanthes amoena race amoena 314
Lepisanthes amoena race imbricata 314
Lepisanthes amoena race pubescens 314
Lepisanthes borneensis 317
Lepisanthes divaricata 309, 310, 314
Lepisanthes divaricata forma divaricata
315, 316, 317
Lepisanthes divaricata forma lunduensis 316, 317
Lepisanthes falcata 310, 317
Lepisanthes falcata subsp. borneensis 267, 309, 312, 317, 318
Lepisanthes falcata subsp. celebica 317, 318
Lepisanthes falcata subsp. falcata 317, 318
Lepisanthes fruticosa 265, 309, 310, 312, 318
Lepisanthes fruticosa race acuminata 319
Lepisanthes fruticosa race fruticosa 319
Lepisanthes fruticosa race glandulosa 265, 319
Lepisanthes heterolepis 325
Lepisanthes kinabaluensis 309, 310, 319, 320
Lepisanthes kunstleri 326
Lepisanthes multijuga 309, 310, 312, 321
Lepisanthes palawanica 293
Lepisanthes petiolaris 325
Lepisanthes ramiflora 309, 310, 322
Lepisanthes rubiginosa 309, 310, 312, 322
Lepisanthes senegalensis 309, 310, 312, 324
Lepisanthes senegalensis race dasypetala 325
Lepisanthes senegalensis race montana 325
Lepisanthes tetraphylla 309, 310, 312, 325
Leucopogon abnormis 161
Leucopogon acuminatus 161
Leucopogon lancifolius 161
Leucopogon malayanus 161
Leucopogon malayanus var. moluccanus 161
Leucopogon moluccanus 161
Leucopogon philippinensis 163
Leucopogon suaveolens 163
Linaceae 151, 167, 175
Litchi 264, 267, 269, 326, 329
Litchi chinensis 264, 265
424
Litchi chinensis subsp. chinensis 327, 328

Litchi ramboutan-ake 348
Litsea 348
Logania 190
LOGANIACEAE 189, 190
Loranthaceae 257
LYTHRACEAE
135, 225
Magnolia 158
Magnoliaceae
155, 403, 404
Malpighiaceae 167
MALVACEAE 233, 234
Mangifera 1, 2, 3, 32, 33, 35
Mangifera sect. Limus 33
Mangifera sect. Mangifera 33
Mangifera applanata 39, 40
Mangifera beccarii 39
Mangifera blommesteinii 33, 35, 36
Mangifera bompardii 39, 40
Mangifera caesia 34, 35, 37
Mangifera decandra 34, 35, 37
Mangifera foetida 34, 35, 38, 43
Mangifera griffithii 34, 36, 38
Mangifera havilandii 43
Mangifera indica 33, 34, 36, 39, 40
Mangifera khoonmengiana 34, 36, 40, 41
Mangifera laurina 39, 40
Mangifera longipes 39, 40
Mangifera macrocarpa 33, 36, 42
Mangifera magnifica 34, 36, 42
Mangifera odorata 33, 35, 36
Mangifera oppositifolia 7
Mangifera orophila 39, 40
Mangifera pajang 34, 35, 43
Mangifera parvifolia 34, 36, 43
Mangifera pentandra 33, 36
Mangifera quadrifida 34, 36, 44
Mangifera rubropetala 39, 40
Mangifera rufocostata 34, 44
Mangifera sclerophylla 39
Mangifera similis 46
Mangifera subsessilifolia 33, 35, 45
Mangifera swintonioides 34, 36, 45
Mangifera torquenda 34, 36, 46
Marcgraviaceae 379
Melanochyla 1, 2, 3, 47
Melanochyla angustifolia 48, 49
Melanochyla auriculata 47, 49
Melanochyla axillaris 47, 50
425
Melanochyla beccariana 49, 50, 51

Melanochyla borneensis 48, 51, 52
Melanochyla bullata 49, 52, 57
Melanochyla caesia 48, 53
Melanochyla castaneifolia 48, 53
Melanochyla condensata 49, 54
Melanochyla elmeri 48, 55
Melanochyla fulvinervis 48, 51, 55
Melanochyla minutiflora 48, 56
Melanochyla montana 48, 56
Melanochyla scalarinervis 48, 57
Melanochyla semecarpioides 48, 57
Melanochyla tomentosa 49, 58
Melanochyla woodiana 48, 58
Melanocommia 72
Melanocommia borneensis 83
Melanorrhoea 22
Melanorrhoea aptera 24
Melanorrhoea beccarii
26
Melanorrhoea macrocarpa 27
Melanorrhoea maingayi 29
Melanorrhoea oba 27
Melanorrhoea speciosa 28
Melanorrhoea tricolor 24
Melanorrhoea wallichii 29
Melastomataceae 136
Meliaceae 2, 3
Melicoccus amoenus 313
Messerschmidia 95
Messerschmidia argentea 95
Mesua 7
Mirabilis 251, 252
Mirabilis jalapa 252
Mischocarpus 263, 264, 267, 268, 270, 329, 330
Mischocarpus fuscescens 330
Mischocarpus pentapetalus 330, 331
Mischocarpus sumatranus 330
Mischocarpus sundaicus 330, 331, 332
Mitrasacme 189, 190
Mitrasacme neglecta 190
Mitrasacme pygmaea 190
Mitreola 189, 190
Monimiaceae 375
Moraceae
375, 383, 384
Morella 245
Mostuea 189
Munchausia speciosa 230
Myrica 245, 246
426
Myrica esculenta 246

Myrica esculenta var. farquhariana 246
Myrica esculenta var. lobbii 246
Myrica esculenta var. missionis 246
Myrica esculenta var. sapida 246
Myrica farquhariana 246
Myrica javanica 246, 247, 248
Myrica lobbii 246
Myrica missionis 246
Myrica sapida 246
MYRICACEAE 245
Myricales 245
Myristicaceae 403
Myrtaceae 7
Myrtales 135
Naxinandra 136
Nectaropetalum 167
Nemostigma 391
Nephelium 264, 267, 269, 270, 333, 334, 349, 352
Nephelium aculeatum 334, 336
Nephelium acuminatum 357
Nephelium beccarianum 340
Nephelium bengalense 288
Nephelium chryseum 343
Nephelium compressum 334, 336, 337, 338
Nephelium cuspidatum 335, 337, 338
Nephelium cuspidatum var. cuspidatum subvar. cuspidatum
Nephelium cuspidatum var. eriopetalum 338, 339
Nephelium cuspidatum var. multinerve 340
Nephelium cuspidatum var. ophioides subvar. beccarianum
Nephelium cuspidatum var. robustum 339
Nephelium daedaleum 334, 337, 341
Nephelium dasyneurum 338
Nephelium echinulatum 288
Nephelium fumatum 286
Nephelium glabrum 343, 345
Nephelium havilandii 335, 341
Nephelium lappaceum 264, 334, 335, 342
Nephelium lappaceum var. lappaceum 343
Nephelium lappaceum var. pallens 343, 350
Nephelium lappaceum var. xanthioides 344, 350
Nephelium longan 287
Nephelium long-yan 287
Nephelium macrophyllum 344, 345, 350
Nephelium maingayi 334, 337, 345
Nephelium malaiensis 288
Nephelium meduseum 334, 345, 346
Nephelium melanomiscum 334, 345, 347
427
340
340
Nephelium multijuga 321

Nephelium multinerve 340
Nephelium mutabile 348
Nephelium mutabile var. pallens 343, 348
Nephelium obliquinerve 338
Nephelium obovatum 343
Nephelium ophelioides 340
Nephelium papillatum 335, 346, 348
Nephelium pupillum 288
Nephelium ramboutan-ake 334, 335, 342, 345, 346, 348, 350
Nephelium reticulatum 334, 335, 350
Nephelium subfalcatum 335, 351
Nephelium subferrugineum 343
Nephelium uncinatum 335, 346, 351
Nephelium xanthioides 344
Nephelium xerospermoides 347
Norrisia 189, 190, 191, 220
Norrisia maior 220, 221
Norrisia malaccensis 221, 222
Norrisia malaccensis var. major 221
Norrisia philippinensis 221
Nothopegia borneensis 52
NYCTAGINACEAE 251
Ochnaceae 379
Ochthocosmus 175
Olacaceae 257
Opiliaceae 257
Otophora 309
Otophora alata 311
Otophora amoena 313
Otophora bijuga 318
Otophora cordigera 313
Otophora divaricata 314
Otophora edulis 311, 313
Otophora fruticosa 318
Otophora glandulosa 318
Otophora imbricata 313
Otophora latifolia 318
Otophora lunduensis 316
Otophora macrocarpa 316
Otophora multijuga 321
Otophora pubescens 313
Otophora pulchella 318
Otophora pyramidalis 316, 317
Otophora ramiflora 322
Otophora spectabilis 313
Otophora tricocca 321
Paliurus dao 18
428
Paliurus edulis 18
Pandanus helicopus 29
Paranephelium 263, 264, 265, 268, 269, 352, 353
Paranephelium joannis 353, 354
Paranephelium nitidum 355
Paranephelium xestophyllum 353, 354, 355
Parasponia 383
Parishia 1, 2, 4, 59, 60
Parishia borneensis 60
Parishia dinghouiana
60
Parishia elmeri 63
Parishia insignis 60
Parishia maingayi 60, 61, 62
Parishia maingayi var. maingayi 63
Parishia maingayi var. minor 63
Parishia paucijuga 60, 63
Parishia polycarpa 63
Parishia sericea 60, 64
Parishia trifoliolata 60, 64
Pavonia rigida 233
Pegia 1, 4
Pegia philippinensis 87
Pegia sarmentosa 4
Pemphis 225, 230
Pemphis acidula 231, 232
Pentamerista 379, 380
Pentaspadon 2, 4, 65
Pentaspadon motleyi 66, 67
Pentaspadon telianthera 87
Peplis 225
Phacellaria 258
Phacellaria malayana 258
Phlebochiton philippinensis 87
Picrophloeus javanensis 205
Pinacopodium 167
Piperales 129
Pisonia 251, 252, 253
Pisonia aculeata 252, 253
Pisonia alba 253
Pisonia excelsa 255
Pisonia grandis 251, 253
Pisonia umbellifera 253, 254, 255
Pleiogynium 4, 67
Pleiogynium timorense 68, 69
Pometia
264, 266, 267, 270, 356, 357
Pometia acuminata 357
Pometia alnifolia 357
Pometia pinnata 357, 358
429
Pometia ridleyi 357, 359

Pometia tomentosa 357
Populus 241
Pseudonephelium fumatum 286
Ptelea arborea 307
Pteleocarpa 94, 103
Pteleocarpa lamponga 103, 104
Pteleocarpa longistyla 103
Pteleocarpa malaccensis 103
Quilamum 140
Rhamnales 181
Rhus 2, 3, 4, 69, 70
Rhus borneensis 70
Rhus chinensis 70
Rhus cobbe 274
Rhus coriaria
69
Rhus nodosa
70, 71
Rhus perakensis 70
Rhus verniciflua 2, 70
Rhynchocalycaceae 136
Rhynchocalyx 136
Rhysotoechia 263, 264, 267, 270, 360
Rhysotoechia koordersii 360, 361
Rotala 225
Rutaceae 78
Rutales 245
Sambucaceae 107
Sambucus 108
Sambucus javanicus 108
SANTALACEAE 257
Santalales 257
Santalum album 257
SAPINDACEAE
2, 263, 264
Sapindaceae (Dyssapindaceae) 264
Sapindaceae (Eusapindaceae) 264
Sapindaceae (Eusapindaceae), Anomophyllae 264
Sapindaceae (Eusapindaceae), Nomophyllae 264
Sapindaceae subfam. Dodonaeoideae 264
Sapindaceae subfam. Dodonaeoideae tribe Cossinieae 264
Sapindaceae subfam. Dodonaeoideae tribe Dodonaeeae 264
Sapindaceae subfam. Dodonaeoideae tribe Doratoxyleae 264
Sapindaceae subfam. Dodonaeoideae tribe Harpullieae 264
Sapindaceae subfam. Dodonaeoideae tribe Koelreuterieae 264
Sapindaceae subfam. Sapindoideae 264
Sapindaceae subfam. Sapindoideae tribe Aphanieae 264
Sapindaceae subfam. Sapindoideae tribe Cupanieae 264
Sapindaceae subfam. Sapindoideae tribe Lepisantheae 264
Sapindaceae subfam. Sapindoideae tribe Melicocceae 264
430
Sapindaceae subfam. Sapindoideae tribe Nephelieae 264

Sapindaceae subfam. Sapindoideae tribe Paullinieeae 264
Sapindaceae subfam. Sapindoideae tribe Sapindeae 264
Sapindaceae subfam. Sapindoideae tribe Schleichereae 264
Sapindaceae subfam. Sapindoideae tribe Thouinieae 264
Sapindi 263
Sapindus cinereus 288
Sapindus fruticosa 318
Sapindus koelreuteria 300
Sapindus longifoloius 288
Sapindus pubescens 303
Sapindus rubiginosa 322
Sapindus senegalensis 324
Sapindus stellatus 288
Sapindus tetraphylla 325
Sarcandra 129, 131
Sarcandra glabra subsp. brachystachys var. brachystachys 131
Schinopsis 2
Schisandraceae 403
Schleichera pentapetala 330
Scleropyrum 258, 260
Scleropyrum maingayi 262
Scleropyrum wallichianum 261, 262
Scrophulariaceae 189
Scrophulariales 189
Scyphostegia 375
Scyphostegia borneensis 376, 377, 378
SCYPHOSTEGIACEAE 375
Semecarpus 1, 2, 3, 72
Semecarpus anacardium 72
Semecarpus angulatus 73, 74
Semecarpus angustifolius 73, 75
Semecarpus borneensis 74, 75
Semecarpus bunburyanus 73, 74, 75
Semecarpus caesia 53
Semecarpus calcicolus 73, 76
Semecarpus cuneiformis 73, 74, 77
Semecarpus cupularis 73, 77
Semecarpus euodiifolius 73, 78
Semecarpus fulvinervis 55
Semecarpus glaucus 73, 74, 78, 79, 84
Semecarpus heterophylla var. caesia
53
Semecarpus heterophyllus 73, 80
Semecarpus impressicostatus 74, 80
Semecarpus kinabaluensis 74, 81
Semecarpus lanceolatus 77
Semecarpus lineatus 73, 81
Semecarpus lurida 20
431
Semecarpus minutipetalus 73, 82

Semecarpus paucinervius 73, 82
Semecarpus pulvinatus 73, 83
Semecarpus rufovelutinus 74, 80, 83
Semecarpus sandakanus 73, 84
Semecarpus scaberulus 76
Semecarpus trengganuensis 74, 84
Sida 233, 235
Sida acuta subsp. acuta 235
Sida rhombifolia subsp. retusa 235
Sida rhombifolia subsp. rhombifolia 235
Simaroubaceae 175
Solenocarpus 4, 85
Solenocarpus indica 85
Solenocarpus philippinensis 85, 86
Solenostigma 387
Solenostigma philippinensis 388
Solenostigma wightii 388
Sonneratiaceae 136, 225
Spanogheaglabra 271
Sphaerocarya wallichianum
262
Spondias cytherea 2
Spondias indica 85
Spondias philippinensis 85, 87
Sponia 396
Sponia acuminatissima 398
Sponia amboinensis 399
Sponia angustifolia 398
Sponia rigida 399
Sponia tomentosa 402
Sponia velutina 402
Sponia virgata 399
Staphylea indica 186
Strychnaceae 190
Strychnos 189, 190
Styphelia 159, 160
Styphelia abnormis 161, 162
Styphelia celebica 164
Styphelia lancifolia 161
Styphelia learmonthiana 166
Styphelia malayana 160, 161
Styphelia malayana var. malayana 161
Styphelia moluccana 161
Styphelia philippinensis 163
Styphelia suaveolens 160, 163
Swintonia 1, 2, 3, 87, 88
Swintonia acuta 88, 89
Swintonia elmeri 29
432
Swintonia foxworthyi 88, 90

Swintonia glauca 88, 90
Swintonia lurida
20
Swintonia minutalata 88, 89, 91
Swintonia sarawakana 88, 91
Swintonia schwenkii 87, 88, 92
Swintonia schwenkii var. beccarii 88
Symphytum 93
Symplocos 158
Takhtajania 403
Tasmannia 404
Tasmannia piperita 406
Terminalia 13
Tetramerista 379, 380
Tetramerista crassifolia 380
Tetramerista glabra 380, 381
Tetramerista montana 380
TETRAMERISTACEAE 379
Tetramista crassifolia 380
Tetramista montana 380
Theaceae 379
Thespesia 235, 241
Thespesia populnea 241, 242
Thymelaeaceae 155
Tournefortia arborea 95
Tournefortia argentea
95
Toxicodendron borneense 70
Toxicodendron nodosum 70
Trema 384, 396, 397
Trema acuminatissima 398
Trema amboinensis 399
Trema angustifolia 397, 398
Trema cannabina 397, 398
Trema orientalis 397, 399, 402
Trema orientalis var. amboinensis.399
Trema orientalis var. bicolor 398
Trema orientalis var. bicornis 400
Trema orientalis var. rigida 400
Trema rigida 400
Trema tomentosa 397, 398, 400, 401, 402
Trema velutina 402
Trema virgata 399
Trigonachras 264, 266, 268, 269, 362, 363
Trigonachras acuta 363, 364
Trigonachras sp. A 363, 365
Trigonachras sp. B 363, 365
Trigonotis 94
Trigonotis borneensis 94
433
Tristira 366
Tristiropsis 264, 265, 268, 366
Tristiropsis ferruginea 366, 367
Trochocarpa 160, 164
Trochocarpa celebica 164, 165
Trochocarpa learmonthiana 166
ULMACEAE 383, 384
Ulmus 383
Urena 234
Urena lobata 234
Urostigma ?cuneatum 169
Urticaceae 383, 384
Urticales 383
Valerianaceae 107
Viburnaceae 107
Viburnum 108, 109
Viburnum amplificatum 109
Viburnum beccarii 109, 110, 111
Viburnum clemensae 109, 110, 112
Viburnum hispidulum 109, 113, 114
Viburnum hispidum 113
Viburnum lutescens 109, 113
Viburnum sambucinum 109, 115
Viburnum vernicosum 109, 115, 116
Vitaceae 181
Willughbeia elliptica 205
WINTERACEAE 403, 404
Xerospermum 264, 266, 267, 269, 368, 369
Xerospermum acuminatissimum 371
Xerospermum acuminatum 371
Xerospermum echinulatum 371
Xerospermum intermedium 371
Xerospermum laevigatum 369
Xerospermum laevigatum subsp. acuminatum 370, 371
Xerospermum laevigatum subsp. laevigatum 370, 371
Xerospermum lanceolatum 371
Xerospermum muricatum 371
Xerospermum noronhianum 369, 370, 371
Xerospermum unijugum 371
Zollingeria 264, 267, 268, 372
Zollingeria borneensis 373, 374
434

(compiled by R.C.K. Chung)
Scientific names in brackets are the correct names corresponding to vernacular names listed here.
amat (Tetramerista glabra Miq.)

379, 380
ambok gobang (Styphelia malayana (Jack) Spreng. var. malayana)
ambuakat (Paranephelium xestophyllum Miq.) 355
ampas tebu (Gironniera parvifolia Planch.) 394
ampungit (Arytera litoralis Blume) 277
ampungot (Paranephelium xestophyllum Miq.) 355
andipatan (Guioa pleuropteris (Blume) Radlk.) 302
angil manuk (Guioa pterorhachis Welzen) 303
anging manuk (Arytera litoralis Blume) 277
angir manuk (Guioa pubescens (Zoll. & Moritzi) Radlk.) 304
antuyut (Tetramerista glabra Miq.) 380
apoh (Harpullia arborea (Blanco) Radlk.) 307
apong keliau (Dimocarpus longan Lour.) 287
arip (Harpullia arborea (Blanco) Radlk.) 307
arit-arit (Lepisanthes tetraphylla (Vahl) Radlk.) 326
asam (Mangifera spp.) 32
asam kumbang (Mangifera quadrifida Jack) 44
asam raba (Mangifera griffithii Hook. f.) 39
asem pajang (Mangifera pajang Kosterm.) 43
bachang (Mangifera foetida Lour.) 38
badut (Mangifera foetida Lour.) 38
balaro (Allophylus cobbe (L.) Raeuschel) 274
balik-balik angin (Trema tomentosa (Roxb.) Hara) 402
bambakat (Paranephelium xestophyllum Miq.) 355
bambangan (Mangifera pajang Kosterm.) 43
bambo (Dimocarpus longan Lour.) 287
bambuakat (Harpullia arborea (Blanco) Radlk.) 307
banati (Fagraea fragrans Roxb.) 208
bannang (Norrisia maior Soler.) 221
bansisi (Lepisanthes tetraphylla (Vahl) Radlk.) 326
baru (Thespesia populnea (L.) Sol. ex Corra) 241
baru-baru (Hibiscus tiliaceus L. subsp. tiliaceus) 239
bawang (Mangifera pajang Kosterm.) 43
bedulang semut (Crypteronia macrophylla Beus.-Osinga) 146
belet (Norrisia maior Soler.) 221
belian landak (Gironniera parvifolia Planch.) 394
belimbing talun (Guioa diplopetala (Hassk.) Radlk.) 298
belingasan (Lepisanthes fruticosa (Roxb.) Leenh.) 319
beluno (Mangifera caesia Jack) 37
besi-besi (Ctenolophon parvifolius Oliv.) 153
biis (Norrisia maior Soler.) 221
binjai (Mangifera caesia Jack) 37
bintanong (Trema tomentosa (Roxb.) Hara) 402
435
163
bira parak (Fagraea montana Wong & Sugau) 211

bonbongan (Allophylus cobbe (L.) Raeuschel) 274
bongor biru (Lagerstroemia speciosa (L.) Pers.) 230
borobogan (Lepisanthes rubiginosa (Roxb.) Leenh.) 323
buah abong (Nephelium lappaceum L.) 342
buah kebuau (Nephelium cuspidatum Blume) 339
buah keregan (Dimocarpus longan Lour.) 287
buah maha (Nephelium lappaceum L.) 342
buah mersilat (Nephelium cuspidatum Blume) 339
buah mua (Nephelium ramboutan-ake (Labill.) Leenh.) 349
buah penancang (Allophylus cobbe (L.) Raeuschel) 274
buah pusit (Pentaspadon motleyi Hook. f.) 67
buah sibau (Nephelium cuspidatum Blume) 339
buah sungket (Guioa bijuga (Hiern) Radlk.) 297
buah sungkit (Nephelium maingayi Hiern) 345
bunga raya (Hibiscus rosa-sinensis L.) 233
bungor (Lagerstroemia spp.) 226
buniton (Mangifera torquenda Kosterm.) 46
bunyitan (Mangifera torquenda Kosterm.) 46
chiru (Dimocarpus longan Lour.) 287
chuchur atap (Styphelia malayana (Jack) Spreng. var. malayana)
damai (Lepisanthes rubiginosa (Roxb.) Leenh.) 323
datanut (Pometia pinnata J.R. Forst. & G. Forst.) 359
delako (Scyphostegia borneensis Stapf) 376
dipangah (Pometia pinnata J.R. Forst. & G. Forst.) 359
embang (Mangifera foetida Lour.) 38
embang (Mangifera pajang Kosterm.) 43
embawang (Mangifera pajang Kosterm.) 43
embuakat (Allophylus cobbe (L.) Raeuschel) 274
empaling (Norrisia maior Soler.) 221
empelam (Mangifera indica L.) 39, 40
emplangau (Pentaspadon motleyi Hook. f.) 67
enkelili (Lepisanthes alata (Blume) Leenh.) 311
enkiong (Dimocarpus longan Lour.) 287
enselan (Pometia pinnata J.R. Forst. & G. Forst.) 359
ensiru (Harpullia arborea (Blanco) Radlk.) 307
entabuloh (Gironniera spp.) 390
entuyut (Tetramerista glabra Miq.)
379, 380
galid (Guioa pterorhachis Welzen) 303
gamperut (Allophylus cobbe (L.) Raeuschel) 274
garong (Guioa pterorhachis Welzen) 303
gatal (Dactylocladus stenostachys Oliv.) 149
geronok (Guioa pubescens (Zoll. & Moritzi) Radlk.) 304
gulambir ayam (Guioa diplopetala (Hassk.) Radlk.) 298
gulambir ayam (Guioa pleuropteris (Blume) Radlk.) 302
gunnong (Norrisia maior Soler.) 221
gurin (Dimocarpus longan Lour.) 287
gurujod (Guioa pleuropteris (Blume) Radlk.) 302
436
163
guruyod (Guioa pterorhachis Welzen) 303

halindagong (Trema spp.) 396, 400
hampas tebu (Gironniera parvifolia Planch.) 394
harut (Nephelium melanomiscum Radlk.) 348
huani (Mangifera odorata Griff.) 35
hugot hugot (Gironniera nervosa Planch.) 392
ilat (Guioa bijuga (Hiern) Radlk.) 297
ilat (Xerospermum noronhianum (Blume) Blume) 372
indulus (Lepisanthes fruticosa (Roxb.) Leenh.) 319
inggir burung (Ixonanthes spp.) 176, 180
intotobokon (Pisonia umbellifera (J. R. Forst.) v. Seemen) 255
ising (Allophylus cobbe (L.) Raeuschel) 274
jabing (Mangifera foetida Lour.) 38
japing (Pentaspadon motleyi Hook. f.) 67
jatem (Fagraea dulitensis Wong & Sugau) 205
jongkong (Dactylocladus stenostachys Oliv.) 147, 149
jung (Lepisanthes tetraphylla (Vahl) Radlk.) 326
kalambuko (Nephelium cuspidatum Blume) 339
kalas (Nephelium daedaleum Radlk.) 341
kalas (Xerospermum noronhianum (Blume) Blume) 372
kalasi (Allophylus cobbe (L.) Raeuschel) 274
kali bang-bang (Hibiscus tiliaceus L. subsp. tiliaceus) 239
kamali (Leea aculeata Blume ex Spreng.) 183
kanawit (Guioa pleuropteris (Blume) Radlk.) 302
kangi runok (Guioa pleuropteris (Blume) Radlk.) 302
kapan (Fagraea teysmannii Cammerl.) 215
kapusong (Nephelium cuspidatum Blume) 339
kasai (Pometia spp.) 356, 359
kasai daun besar (Pometia pinnata J.R. Forst. & G. Forst.) 359
kasat (Buchanania spp.) 9
kata manok (Scyphostegia borneensis Stapf) 376
kayo bilong (Dimocarpus longan Lour.) 287
kayo mupa (Ixonanthes spp.) 176
kayu chelega (Tetramerista glabra Miq.) 380
kayu hujan (Tetramerista glabra Miq.) 380
kayu kenupan (Leea indica (Burm. f.) Merr.) 187
kayu lulus (Guioa diplopetala (Hassk.) Radlk.) 298
kayu mup (Ixonanthes spp.) 176
kayu soman (Dimocarpus longan Lour.) 287
kayu tangiran (Tetramerista glabra Miq.) 380
kejangan (Guioa pleuropteris (Blume) Radlk.) 302
kelampu (Allophylus cobbe (L.) Raeuschel) 274
kelinga (Lepisanthes fruticosa (Roxb.) Leenh.) 319
kelinga gaba (Lepisanthes senegalensis (Juss. ex Poir.) Leenh.) 325
kemali (Leea aculeata Blume ex Spreng.) 183
kemali (Leea indica (Burm. f.) Merr.) 187
kemantan (Mangifera foetida Lour.) 38
kemantan (Mangifera torquenda Kosterm.) 46
437
kepala tundang (Buchanania spp.) 9

kereneong (Trema spp.) 396, 399, 402
kerimbau (Allophylus cobbe (L.) Raeuschel) 274
ketang (Campnosperma spp.) 13
kiah (Paranephelium joannis Davids) 355
kiga (Gironniera nervosa Planch.) 392
kinkidon mantok (Crypteronia paniculata Blume var. paniculata) 146
kolomundui (Nephelium maingayi Hiern) 345
krian (Ctenolophon parvifolius Oliv.) 153
kuayun (Gironniera subaequalis Planch.) 396
kubili (Cubilia cubili (Blanco) Adelbert) 279
kundang (Bouea spp.) 2, 5, 7
kundang rumenia (Bouea spp.) 5
kwini (Mangifera odorata Griff.) 35
lakacho (Pentaspadon motleyi Hook. f.) 67
langkubing (Hibiscus tiliaceus L. subsp. tiliaceus) 239
lanyat (Mangifera caesia Jack) 37
lapak lapak (Guioa bijuga (Hiern) Radlk.) 297
layang-layang (Parishia spp.) 59
lemag (Nephelium cuspidatum Blume) 339
lemantan (Mangifera torquenda Kosterm.) 46
limau antu (Scleropyrum wallichianum (Wight & Arn.) Arn.) 262
lindagong (Trema tomentosa (Roxb.) Hara) 402
lipang lipang (Guioa pterorhachis Welzen) 303
lipupudsu (Lepisanthes rubiginosa (Roxb.) Leenh.) 323
litchi (Litchi chinensis Sonn.) 264
litoh (Ctenolophon parvifolius Oliv.) 153
lokud (Harpullia cupanioides Roxb.) 308
lokud-lokud (Guioa spp.) 295
longan (Dimocarpus longan Lour.) 264
longitam (Nephelium ramboutan-ake (Labill.) Leenh.) 349
luazon (Gironniera nervosa Planch.) 392
luazon (Gironniera subaequalis Planch.) 396
ludang-ludang (Styphelia malayana (Jack) Spreng. var. malayana) 163
lulupitan (Lepisanthes fruticosa (Roxb.) Leenh.) 319
lundagong (Trema tomentosa (Roxb.) Hara) 402
lung bipuyoh (Nephelium lappaceum L.) 342
lychee (Litchi chinensis Sonn.) 264, 327
ma (Nephelium ramboutan-ake (Labill.) Leenh.) 349
mabo (Dimocarpus longan Lour.) 287
machang (Mangifera spp.) 32, 33, 38
mahau (Dimocarpus longan Lour.) 287
maki china (Styphelia malayana (Jack) Spreng. var. malayana) 163
mali-mali (Leea spp.) 181, 183, 185, 187
mambahino (Allophylus cobbe (L.) Raeuschel) 274
mambu (Dimocarpus longan Lour.) 287
mambuakat (Dimocarpus fumatus (Blume) Leenh. subsp. fumatus) 287
mampongoh (Harpullia cupanioides Roxb.) 308
438
mangga ayer (Mangifera indica L.) 40

manggalum (Nephelium cuspidatum Blume) 339
mantan (Mangifera torquenda Kosterm.) 46
marakiang (Dimocarpus fumatus (Blume) Leenh. subsp. fumatus) 287
marong (Trema spp.) 396, 402
mata kucing (Dimocarpus longan Lour.) 287
mata pait (Guioa pleuropteris (Blume) Radlk.) 302
mawang (Mangifera pajang Kosterm.) 43
medang jongkong (Dactylocladus stenostachys Oliv.) 149
medang kasap (Gironniera spp.) 390, 392, 394, 396
medang miang (Dactylocladus stenostachys Oliv.) 149
medang tabak (Dactylocladus stenostachys Oliv) 149
melamut (Styphelia malayana (Jack) Spreng. var. malayana) 163
melenjau (Nephelium ramboutan-ake (Labill.) Leenh.) 349
melingkat (Lepisanthes senegalensis (Juss. ex Poir.) Leenh.) 325
mempelam (Mangifera indica L.) 39, 40
mengalim (Nephelium cuspidatum Blume) 339
mengkuang (Dracontomelon spp.) 17
menoa (Hibiscus tiliaceus L. subsp. tiliaceus) 239
mentigi (Pemphis acidula J.R. Forst. & G. Forst.) 232
mepa (Norrisia maior Soler.) 221
merabong (Pometia pinnata J.R. Forst. & G. Forst.) 359
merbuta gunung (Myrica esculenta Buch.-Ham.) 247
merebong (Dactylocladus stenostachys Oliv.) 149
meridian (Ixonanthes spp.) 176
meritam (Nephelium ramboutan-ake (Labill.) Leenh.) 349
merkiang (Dimocarpus longan Lour.) 287
merpauh (Swintonia spp.) 87, 88
mujau (Nephelium maingayi Hiern) 345
mujau (Nephelium ramboutan-ake (Labill.) Leenh.) 349
mumboh (Dimocarpus longan Lour.) 287
munggulan ayam (Guioa pleuropteris (Blume) Radlk.) 302
murieng (Trema tomentosa (Roxb.) Hara) 402
nasi-nasi (Trema tomentosa (Roxb.) Hara) 402
nunuk-nunuk (Arytera litoralis Blume) 277
nyaletang (Campnosperma spp.) 13
nyvang (Norrisia maior Soler.) 221
obah (Ctenolophon parvifolius Oliv.) 153
ombinaton (Fagraea fragrans Roxb.) 208
ondogon (Trema tomentosa (Roxb.) Hara) 402
otak udang (Buchanania spp.) 9
pachu (Mangifera foetida Lour.) 38
pagar anak (Ixonanthes spp.) 176, 177
panapok ayer (Ganophyllum falcatum Blume) 291
pangin (Mangifera foetida Lour.) 38
pangiz (Thespesia populnea (L.) Sol. ex Corra) 241
pantan pantan (Leea indica (Burm. f.) Merr.) 187
pareh (Litchi chinensis Sonn. subsp. chinensis) 327
439
pawuk (Mangifera foetida Lour.) 38

pelajau (Pentaspadon motleyi Hook. f.) 65, 67
pelasih (Pentaspadon motleyi Hook. f.) 67
pelong (Pentaspadon spp.) 65
pengkul (Guioa pleuropteris (Blume) Radlk.) 302
perajau (Pentaspadon motleyi Hook. f.) 67
peraju (Lepisanthes alata (Blume) Leenh.) 311
perepat burong (Erythroxylum spp.) 170
perlimpong hutan (Lagerstroemia pustulata Furtado & Montien) 228
petinag (Arytera litoralis Blume) 277
petudang (Cordia dichotoma Forster) 100
piri manok (Guioa pleuropteris (Blume) Radlk.) 302
pitoh (Swintonia spp.) 87
pitoh ayer (Swintonia acuta Engl.) 88
pitoh paya (Swintonia glauca Engl.) 91
pokok seriawan burung (Guioa pleuropteris (Blume) Radlk.) 302
potudung (Cordia dichotoma Forster) 100
pudun (Nephelium ramboutan-ake (Labill.) Leenh.) 349
puloh (Gironniera nervosa Planch.) 392
punah (Tetramerista glabra Miq.) 380
raba (Mangifera parvifolia Boerl. & Koord.) 44
rade (Mangifera torquenda Kosterm.) 46
rafag (Dimocarpus longan Lour.) 287
rambai-rambai (Crypteronia griffithii C.B. Clarke) 145
rambutan (Nephelium spp.) 264, 333, 339, 341, 342
rambutan ake (Nephelium ramboutan-ake (Labill.) Leenh.) 348
rambutan hutan (Nephelium meduseum Leenh.) 347
rambutan hutan (Nephelium ramboutan-ake (Labill.) Leenh.) 349
rambutan kunyit (Dimocarpus dentatus Meyer ex Leenh.) 286
randagong (Trema spp.) 396, 398, 399, 400, 402
randog (Hibiscus macrophyllus Roxb. ex Hornem.) 238
rangalau (Nephelium lappaceum L.) 342
ranggu (Koordersiodendron spp.) 30
rapanah (Pometia pinnata J.R. Forst. & G. Forst.) 359
rengas (Gluta spp.) 22
rengas (Semecarpus spp.) 72
rengas padang (Androtium spp.) 4
resat (Xerospermum noronhianum (Blume) Blume) 372
rhu bukit (Gymnostoma sumatranum (Jungh. ex de Vriese) L.A.S. Johnson)
rhu laut (Casuarina spp.) 120
rhu ronang (Gymnostoma nobile (Whitmore) L.A.S. Johnson) 125
ribau balabau (Dimocarpus fumatus (Blume) Leenh. subsp. fumatus) 287
rondogong (Trema tomentosa (Roxb.) Hara) 402
rondongon (Trema tomentosa (Roxb.) Hara) 402
rozan (Gironniera parvifolia Planch.) 394
ruazon (Gironniera parvifolia Planch.) 394
rukam hantu (Scyphostegia borneensis Stapf) 376
rupag (Guioa bijuga (Hiern) Radlk.) 297
440
127
rusak (Styphelia malayana (Jack) Spreng. var. malayana) 163

russen (Gironniera subaequalis Planch.) 396
rutan berangat (Guioa pubescens (Zoll. & Moritzi) Radlk.) 304
ruwayon (Gironniera subaequalis Planch.) 396
saasa (Guioa koelreuteria (Blanco) Merr.) 300
saasa (Guioa pleuropteris (Blume) Radlk.) 302
saasa (Guioa pubescens (Zoll. & Moritzi) Radlk.) 304
sabon-sabon (Dimocarpus longan Lour.) 287
sabong ambok (Scleropyrum wallichianum (Wight & Arn.) Arn.) 262
saksah (Guioa koelreuteria (Blanco) Merr.) 300
saksah (Guioa pubescens (Zoll. & Moritzi) Radlk.) 304
sakubong (Harpullia arborea (Blanco) Radlk.) 307
samondu (Tetramerista glabra Miq.) 380
sangan (Nephelium lappaceum L.) 342
sangkurat (Scyphostegia borneensis Stapf) 376
sasah (Allophylus cobbe (L.) Raeuschel) 274
sasah (Guioa koelreuteria (Blanco) Merr.) 300
segera (Lepisanthes falcata (Radlk.) Leenh. subsp. borneensis (Leenh.) Leenh.)
selan (Pometia pinnata J.R. Forst. & G. Forst.) 359
sengkuang (Dracontomelon spp.) 17
serait (Nephelium maingayi Hiern) 345
sikip sikip (Guioa pterorhachis Welzen) 303
silak (Pometia pinnata J.R. Forst. & G. Forst.) 359
simpapait (Norrisia maior Soler.) 221
sintatanod (Lepisanthes alata (Blume) Leenh.) 311
siopnama (Cordia dichotoma Forster) 100
sipaju (Lepisanthes alata (Blume) Leenh.) 311
sira (Fagraea spicata Baker) 214
sisipen pelandok (Allophylus cobbe (L.) Raeuschel) 274
situgog (Nephelium lappaceum L.) 342
sokong ranyai (Fagraea volubilis Wall.) 217
sokungu (Lepisanthes alata (Blume) Leenh.) 311
sosopon bonging (Gironniera nervosa Planch.) 392
suang rason (Lepisanthes rubiginosa (Roxb.) Leenh.) 323
sukang ranyai (Fagraea cuspidata Blume) 204
sukong ganyai (Fagraea cuspidata Blume) 204
suyut (Tetramerista glabra Miq.) 380
tamasuk hutan (Fagraea belukar Wong & Sugau) 200
tamasuk jantan (Fagraea rugulosa Wong & Sugau) 212
tambiaton (Fagraea fragrans Roxb.) 208
tambinaton (Fagraea fragrans Roxb.) 208
tambuakat (Harpullia arborea (Blanco) Radlk.) 307
tampasu (Crypteronia borneensis J.T. Pereira & Wong) 142
tamud tamud (Lepisanthes fruticosa (Roxb.) Leenh.) 319
tanggianggi (Guioa pterorhachis Welzen) 303
tanggianuk (Guioa pleuropteris (Blume) Radlk.) 302
tangking manok (Guioa pleuropteris (Blume) Radlk.) 302
tanjiong awan (Dactylocladus stenostachys Oliv.) 149
441
317
tanjsanuk (Guioa pleuropteris (Blume) Radlk.) 302

tarukakang (Scyphostegia borneensis Stapf) 376
telatang (Campnosperma spp.) 13
teletang (Crypteronia macrophylla Beus.-Osinga) 146
teloyang (Leea aculeata Blume ex Spreng.) 183
telur cicak (Myrica esculenta Buch.-Ham.) 247
temasuk (Fagraea fragrans Roxb.) 208
temasuk (Fagraea gigantea Ridl.) 209
temasuk hutan (Fagraea gigantea Ridl.) 209
temasuk laut (Fagraea fragrans Roxb.) 208
temasuk pasir (Fagraea fragrans Roxb.) 208
tembusu (Fagraea fragrans Roxb.) 208
tembusu (Fagraea volubilis Wall.) 217
tembusu bukit (Fagraea belukar Wong & Sugau) 200
tembusu gajah (Fagraea cuspidata Blume) 204
tembusu tagai (Fagraea belukar Wong & Sugau) 200
tengaranuk (Guioa pterorhachis Welzen) 303
tepada (Gironniera parvifolia Planch.) 394
teradam bukit (Daphniphyllum glaucescens Blume) 158
terentang (Campnosperma spp.) 13
terentang (Campnosperma auriculatum (Blume) Hook. f.) 14
terentang paya (Campnosperma coriaceum (Jack) Hall. f. ex Steenis) 16
terentang puteh (Campnosperma squamatum Ridl.) 16
terepit (Tetramerista glabra Miq.) 380
terindak ati (Styphelia malayana (Jack) Spreng. var. malayana) 163
terongau paya (Fagraea resinosa Leenh.) 212
tetang (Campnosperma spp.) 13
tibabah (Lagerstroemia speciosa (L.) Pers.) 230
tikam (Trema tomentosa (Roxb.) Hara) 402
tinggal asam (Guioa bijuga (Hiern) Radlk.) 297
tinggirang pirak (Fagraea cuspidata Blume) 204
tingir manok (Guioa pleuropteris (Blume) Radlk.) 302
tingir manuk (Guioa pterorhachis Welzen) 303
todopon puak (Fagraea cuspidata Blume) 204
todopon puak (Fagraea spicata Baker) 214
todopon puok (Fagraea volubilis Wall.) 217
togimamalid (Leea indica (Burm. f.) Merr.) 187
togunumali (Leea aculeata Blume ex Spreng.) 183
tombuakat (Paranephelium xestophyllum Miq.) 355
tomudong (Cordia dichotoma Forster) 100
tongianum (Guioa pleuropteris (Blume) Radlk.) 302
towi (Trema spp.) 396
tukil-tukil (Allophylus cobbe (L.) Raeuschel) 274
tuku baroh (Trema tomentosa (Roxb.) Hara) 402
tulau (Scyphostegia borneensis Stapf) 376
tundun biawak (Xerospermum noronhianum (Blume) Blume) 372
tutubokon (Pisonia umbellifera (J. R. Forst.) v. Seemen) 255
tuyot (Tetramerista glabra Miq.) 380
442
tuyut (Tetramerista glabra Miq.) 379, 380

ubah samak (Crypteronia macrophylla Beus.-Osinga) 146
ubah semut (Crypteronia griffithii C.B. Clarke) 145
ubah semut (Crypteronia macrophylla Beus.-Osinga) 146
ubah semut (Crypteronia paniculata Blume var. paniculata)
uchip bali (Fagraea teysmannii Cammerl.) 215
umit (Pentaspadon motleyi Hook. f.) 67
untoh bulu (Gironniera spp.) 390, 396
upi (Parishia spp.) 59
upi bunga (Parishia insignis Hook. f.) 61
upi paya (Parishia maingayi Hook. f.) 63
uping (Pentaspadon motleyi Hook. f.) 67
wahab (Mangifera griffithii Hook. f.) 39
wani (Mangifera odorata Griff.) 35
443
146
vi
TREE FLORA
of
SABAH
AND
SARAWAK
Tree Flora of Sabah and Sarawak Editorial Committee Chairman: E. Soepadmo (Forest Research
Institute Malaysia). Members: A. Latiff (Universiti Kebangsaan Malaysia), Ruth Kiew (Universiti
Pertanian Malaysia), H.S. Lee (Sarawak Forestry Department), L.G. Saw (Forest Research Institute
Malaysia), K.M. Wong (Sabah Forestry Department). Editorial assistant: R.C.K. Chung (Forest
Research Institute Malaysia). Advisors: P.S. Ashton (Harvard University, U.S.A.), J. Dransfield
(Royal Botanic Gardens, Kew, U.K.), C. Kalkman (Rijksherbarium, Leiden, the Netherlands), T.C.
vi
TREE FLORA
of
SABAH AND SARAWAK
Volume Three
edited by
E. Soepadmo and L.G. Saw
Sabah Forestry
Forest Research Institute

Malaysia
2000
L.G. Saw & B. Aslina (2007)
vi
Sarawak Forestry
Copyright Forest Research Institute Malaysia (FRIM)

Sabah Forestry Department, Malaysia
Sarawak Forestry Department, Malaysia
Correspondence:
Forest Research Institute Malaysia (FRIM), Kepong, 52109 Kuala Lumpur, Malaysia.
Perpustakaan Negara Malaysia Cataloguing-in-Publication Data

Tree Flora of Sabah and Sarawak / edited by E. Soepadmo,
L.G. Saw
Includes index
ISBN 983-2181-06-2 (Vol. 3)
1. BotanySabah. 2. BotanySarawak. 3. BotanyBorneo.
4. BotanyClassification. I. Soepadmo, E., II. Saw, L.G.
581.95952
Electronic version by L. G. Saw and B. Aslina, Forest Research Institute Malaysia, 2007.
CONTENTS
Page
Foreword
vii
Acknowledgements
ix
Dedication
xi
FAMILIES
1. Fagaceae (E. Soepadmo, S. Julia & Rusea Go)
2. Leguminosae-Caesalpinioideae (Ding Hou)
119
3. Moraceae (K.M. Kochummen & Rusea Go)
181
4. Myristicaceae (W.J.J.O. de Wilde)
335
475
479
481
501
FAGACEAE (SOEPADMO, JULIA & GO)
FOREWORD
nce again we are delighted to be able to welcome a new volume of the Tree
Flora of Sabah and Sarawak. This volume contains accounts of 4 families
(Fagaceae, Leguminosae-Caesalpinioideae, Moraceae, and Myristicaceae)
comprising 34 genera and 431 species. We learn that of the total 431 tree species treated
in this volume, 28 species or 9.8% are new to science, and 176 species or 40.8% are
endemic either to Borneo or to Sabah and Sarawak. Of the endemic species, 44 or
10.2% are hyper-endemic being rare and restricted in their distribution to a particular
type of forest habitat, and are known only from a single or a few localities in Sabah
and Sarawak. We also understand that 11 of the new species belong to the Fagaceae,
a family already revised 27 years ago under the Flora Malesiana project, and 17 species
belong to the Moraceae, a family yet to be revised under the Flora Malesiana project.
The family Leguminosae-Caesalpinioideae had fairly recently been revised for Flora
Malesiana (Ding Hou, K. Larsen & S.S. Larsen, 1996), while the Myristicaceae have
yet to be published in the same Flora. Thus, Dr. W.J.J.O. de Wildes account on the
Myristicaceae for Sabah and Sarawak represents the most up-to-date treatment of the
family for the region. It contains descriptions of 110 tree species in 5 genera, representing
almost three-fold the number of species listed by Merrill in 1921 and nearly two-fold
the number of species enumerated by Masamune in 1942 for Borneo. The results
demonstrate once again that continued exploration of the tree flora of Sabah and Sarawak,
followed by detailed scientific investigation and accurate documentation, will yield
valuable information pertaining to the botanical and conservation status of the tree
resources of Sabah and Sarawak. Such information is pivotal to our understanding of
the biological diversity of Borneo in general and that of Sabah and Sarawak in particular.
The availability of fundamental botanical information on the Tree Flora of Sabah and
Sarawak (TFSS) will further enhance our capability to protect, conserve, use and manage
tree resources on a sustainable basis. At the national level, the publication of TFSS
volume 3 symbolises the Malaysian Governments commitment towards achieving its
National Policy on Biological Diversity launched in April 1998. In the international
arena, the publication also proves that the Malaysian Government is fully committed
in fulfilling its obligations and responsibilities under the International Convention on
Biological Diversity.
Funding for a long-term project is a notorious problem, perennially faced by research
institutions that embark on such a project. Between 1996 and 1999, no additional
international funding was available for the Tree Flora of Sabah and Sarawak Project,
resulting in the drastic reduction of its activities and the delay of publication of TFSS
volume 3. However, we are extremely grateful for the timely approval of research grant
in 1996 by the Malaysian Government (through the IRPA grant for Project 0804
010024/1996) and further additional support in 1997 from the Malaysian Forestry
Research and Development Board (MFRDB). We also wish to acknowledge the generosity
of the Overseas Development Administration of the United Kingdom for allowing the
vii
Project to continue utilising the balance of funds approved in the 1996/1997 Financial
Year.
To continue the project, and thus to achieve its original objective of producing 8 volumes
of the Tree Flora of Sabah and Sarawak, we earnestly hope that in the new millennium
the Project would be able to attract enough funding from both national and international
agencies.
We wish to express our deepest gratitude to various national and international research
institutions for their help and collaboration in making the publication of the Tree Flora
of Sabah and Sarawak volume 3 possible. To the contributors of this volume, we wish
to congratulate them for their excellent achievement. Finally, our thanks and congratulations
are also due to the editors of this volume and to staff of the Forest Research Institute
Malaysia (FRIM), Sabah Forestry Department, and Sarawak Forestry Department for
their support and involvement in the project.
Dato Dr. Abdul Razak Mohd. Ali

Director-General
Mr. Daniel Khiong Kok Sin

Director
Sabah Forestry Department
Malaysia
Mr. Cheong Ek Choon

Director
Sarawak Forestry Department
Malaysia
viii
ACKNOWLEDGEMENTS
t gives us great pleasure once again to put on record our deepest gratitude and
appreciation to various institutions and individuals who have, in one way or another,
assisted in the production and publication of the Tree Flora of Sabah and Sarawak
volume 3. The publication of this volume would not have been possible without the
timely financial support of the Malaysian Government and the Malaysian Forestry
Research and Development Board (MFRDB), and the generous permission granted by
the Overseas Development Administration of the United Kingdom for allowing the Project
to utilise the balance of funds approved in the 1996/1997 Financial Year. Tan Sri G.
K. Rama Iyer (Chairman of the MFRDB), Dato Dr. Abdul Razak Mohd. Ali (DirectorGeneral of the Forest Research Institute Malaysia), Mr. Daniel Khiong Kok Sin (Director
of the Sabah Forestry Department), and Mr. Cheong Ek Choon (Director of the Sarawak
Forestry Department) have continuously given their full encouragement and support
for the project to succeed and achieve its objectives.
The Directors, Keepers, and Curators of the herbaria at Bogor (BO), the British Museum
(BM), Copenhagen (C), Geneva (G), Kew (K), Kepong (KEP), Universiti Malaya (KLU),
Leiden (L), Paris (P), Sandakan (SAN), Kuching (SAR), Singapore (SING), Stockholm
(S), Utrecht (U), the Universiti Kebangsaan Malaysia, Bangi (UKMB), and the Uppsala
University (UPS) have kindly allowed the project botanists to access information and
obtain services available at their institutions, and provided loans of specimens.
Prof. P.S. Ashton (Harvard University, U.S.A.), Dr. J. Dransfield (Royal Botanic Gardens,
Kew, U.K.), Dr. T.C. Whitmore (University of Cambridge, U.K.), and Dr. W.J.J.O. de
Wilde (Rijksherbarium, Leiden, the Netherlands) provided valuable advice and guidance
on editorial matters. Dr. C.C. Berg (University of Bergen, Norway) generously gave
constructive criticism and other valuable assistance in improving the scientific quality
of the manuscript of the Moraceae, particularly that of Ficus. Dr. J.F. Veldkamp
(Rijksherbarium, Leiden, the Netherlands) kindly provided Latin diagnoses for new
species described under the genera Artocarpus, Ficus (Moraceae), and Lithocarpus
(Fagaceae) published elsewhere. We are deeply indebted to Prof. Abdul Latiff Mohamad
(Universiti Kebangsaan Malaysia), Dr. H.S. Lee (Sarawak Forestry Department), Dr.
Ruth Kiew (Singapore Botanic Gardens), Dr. K.M. Wong (Universiti Malaya), and Mr.
R.C.K. Chung (Forest Research Institute Malaysia) for their help and cooperation as
members of the Editorial Committee.
We are extremely grateful to many others who have rendered help and assistance during
the visits of project botanists to their institutions. They are, at Bogor: Dr. E.A. Widjaya,
Dr. J.P. Mogea, Dr. R.E. Nasution, and Dr. S. Danimihardja; at Kew: Prof. G.T. Prance,
Mr. M.J.E. Coode, Dr. J. Dransfield, Dr. D. Kirkup, Dr. M. Cheek, Mr. P. Boyce, Mrs.
Diane Bridson, and Mr. N. Martland; at Leiden: Prof. P. Baas, Dr. Marco Roos, Dr.
J.F. Veldkamp, Dr. W.J.J.O. de Wilde, Dr. E.F. de Vogel, Dr. Ding Hou, Dr. P. van
ix
Welzen, Dr. M.M.J. van Balgooy, Dr. P. Kessler, Dr. F. Adema, Mrs. Stans Kofman,
and Mr. C.J.W. Lut; at Oxford: Dr. P. Bacon and Ms. Rosemary Wise; and at Singapore:
Dr. S.C. Chin and Dr. Ruth Kiew. We are also very much obliged to the staff of the
libraries and photography units at the British Museum, Kew, and Leiden.
At the home front, we are greatly indebted to many who have given help and assistance,
and they are, at Kepong: Dr. Wan Razali Wan Mohd., Dr. N. Manokaran, Dr. F.S.P.
Ng, Dr. H.T. Chan, Dr. Abd. Rashid A. Malek, Dr. Lillian S.L. Chua, the late Mr.
K.M. Kochummen, Mr. R.C.K. Chung, Ms. Noorsiha Ayop, Mr. Mat Asri Ngah Sanah,
Mr. Kamarudin Saleh, Mr. Damahuri Sabari, Mrs. Zainun Othman, Mrs. Asnah Hashim,
Mrs. Saripah Barom, and Mr. Samsul Bahrin Abdullah; at Sandakan: Dr. Sining Unchi,
Mr. R.C. Ong, Dr. Y.F. Lee, Ms. J.T. Pereira, Mr. J.B. Sugau, Mr. S.P. Lim, Mr. Leopold
Madani, and Mr. Dewol Sundaling; and at Kuching: Dr. H.S. Lee, Dr. Paul P.K. Chai,
Mr. Abang Abdul Hamid Karim, Mr. Abang Mohtar Abang Pawozan, Mrs. Runi S.
Pungga, Mr. Julaihi Abdullah, Mr. Stephen P. Teo, and Ms. Mohizah Mohamad.
Finally, we would like to express our deepest gratitude and appreciation to Ms. Rosemary
Wise (Oxford Forestry Institute), Mr. Joseph Pao (Sarawak Forestry Department), Mr.
Mohd. Nizam Isa (Universiti Kebangsaan Malaysia), Mr. Ubaldos Majawal (Sabah
Forestry Department), Mr. Wan Omarshah Wan Putera (Sarawak Forestry Department),
Mr. J.V. Os (Rijksherbarium, Leiden), Dr. Ding Hou (Rijksherbarium, Leiden), and the
late Mr. A. Hamzah and Mr. Md. Toha (Herbarium Bogoriense, Bogor) who have ably
and diligently prepared the illustrations included in this volume. To Dr. Marco Roos
(Rijksherbarium, Leiden) and the Flora Malesiana Editorial Committee, we are very
much indebted for their permission to reproduce figures 1, 3, 4, 6, 7, 12, 13, and 14
of the Leguminosae-Caesalpinioideae in this volume.
E. Soepadmo
L.G. Saw
December 1999
FAGACEAE
DEDICATION
(SOEPADMO,
(SOEPADMO
JULIA
& &SAW)
GO)
DEDICATION
Kizhakkedathu Mathai Kochummen
(6 July 1931, Kerala, India31 March 1999, Kuala Lumpur, Malaysia)
he birth of Tree Flora of Sabah and Sarawak volume 3 sadly coincided with the
untimely departure of K.M. Kochummen on 31 March 1999 at the age of 67,
following a brief spell of illness. With his demise we also lost the most experienced
and much respected Forest Botanist.
Born in Kerala on 6 July 1931, Koch, as his peers and friends affectionately knew
him, obtained his B.Sc. degree from the University of Travancore, Kerala in 1951 (van
Steenis-Kruseman, 1974). He came to Malaya in December 1952 and joined the Forest
Research Institute (FRI) at Kepong in January 1953 as Research Assistant. He worked
under the guidance and leadership of John Wyatt-Smith, who was then the Forest Botanist
of the Institute. In 1957 he was promoted to Assistant Forest Botanist, and subsequently
took charge of the Kepong herbarium during 19601963. In 1968, he went to the
University of Edinburgh, U.K., to pursue a Diploma Course in Plant Taxonomy under
the Colombo Plan Scholarship Award. Following his return to Kepong, he was promoted
first to Forest Botanist in 1971, and then to Senior Forest Botanist in 1980 (Wong,
1987). He retired from the Institute in 1986 after 33 years of service. Immediately
after his retirement, Koch took up a consultancy assignment as Senior Research Fellow
(19861991) with the Smithsonian Tropical Research Institute (STRI) to identify and
enumerate tree species in the 50-ha demographic plot in Pasoh Forest Reserve, Negri
Sembilan (Kiew, 1999). From 1992 to his demise in March 1999, the Tree Flora of
Sabah and Sarawak Project based at the Kepong herbarium engaged Koch as Senior
Botanist.
Kochs major botanical contribution hinged largely on his unrivalled natural ability in
recognising tree species by their leaf and other readily accessible field/vegetative
characters, such as those of the bark, bole, exudate, buttress, and root system. This
unsurpassed ability was acquired and sharpened during his first decade of service at
Kepong, spurred by the need for botanical and ecological data of all timber species
native to Malaya. This fundamental information was vital and essential to the success
of Wyatt-Smiths monumental undertaking to produce the Manual of Malayan Silviculture
for Inland Forests (Ng & Saw, 1999). Kochs hard work and relentless effort resulted
in the publication of his Keys for the botanical identification of non-dipterocarp timber
trees of Malaya, based on field characters (FRI Research Pamphlet No. 43, 1963).
After further testing and improvement of his approach and methodology of field
identification, Koch used his practical knowledge and experience to revise Wyatt-Smiths
Pocket Check List of Timber Trees (first published in 1952). The much-improved Pocket
Check List was published in 1964.
xi
The next major assignment in Kochs botanical career at FRI began when the Institute,
with the financial assistance of the Colombo Plan Technical Aid Programme, embarked
on another monumental project, the Tree Flora of Malaya. Under this project, T.C.
Whitmore was engaged as the leader and coordinator during 19651972. The first and
second volumes of the Tree Flora, edited by T.C. Whitmore, were published in 1972
and 1973 respectively. The third and fourth volumes, edited by F.S.P. Ng, appeared
in 1978 and 1989, respectively. A total of 100 families comprising 2677 tree species
were treated in these volumes. Koch contributed revisions of 26 families embracing
1089 species (about 41% of the total). Despite his heavy commitment with the project,
Koch was always mindful of his responsibility in ensuring the usefulness of WyattSmiths Pocket Check List of Timber Trees. Hence, in 1976 he began to revise the
book by incorporating more up-to-date botanical and other information generated by
the publication of the first two volumes of the Tree Flora of Malaya and other sources
such as monographic accounts of families published in the Flora Malesiana volumes.
The revised Pocket Check List was eventually published in 1979.
Being indefatigable in his botanical ambition, Koch, after retirement, continued to make
significant contributions towards the betterment of our knowledge of the tree flora of
Malaysia. Under the leadership of P.S. Ashton (Harvard University) and in collaboration
with other scientists involved in the STRI Project (19861991), Koch managed to establish
the species identities of a staggering 335,240 juvenile and mature trees (with dbh = 1 cm
and greater) enumerated in the 50-ha demographic plot at Pasoh Forest Reserve (Manokaran
et al.,1992). As a testimony of his unsurpassed knowledge on the usefulness of vegetative
characters for identifying forest trees, Koch, in 1997, published his book on the Tree Flora
of Pasoh Forest.
The final chapter of Kochs illustrious botanical achievements is epitomised by his
involvement in the Tree Flora of Sabah and Sarawak (TFSS) Project, jointly carried
out by the Forest Research Institute Malaysia (FRIM), Sabah Forestry Department, and
Sarawak Forestry Department. In 1992, he joined the Project as Senior Botanist with
a special task of revising large and difficult families. During his tenure, he contributed
revisions of the Burseraceae, Celastraceae and Ochnaceae published in the TFSS volume
1 (1995), and of the Anacardiaceae in volume 2 (1996). In December 1998, Koch
completed the manuscript of the Moraceae (Artocarpus and Ficus) published in this
volume. Apart from revising families for the TFSS volumes, Koch also actively participated
in the workshop and training courses organised by the project to upgrade the taxonomic
research capability of 25 young Malaysian botanists involved in the project. To them,
Koch was a good teacher and friendly instructor. Concurrent with his duty under the
TFSS Project, Koch also revised Wyatt-Smiths Pocket Check List for the third time.
The much extended and updated edition was published by FRIM in the middle of 1999,
just a few months after his departure. Beside his enormous contribution towards the
advancement of our knowledge of the Malaysian flora, his peers and friends in Malaysia
as well as in other parts of the world will remember the humble and soft-spoken Koch
for his unfailling courtesy and willingness to share knowledge with others.
References
Kiew, R. (1999). Gard. Bull. Sing. 51(1): 125126.
xii
FAGACEAE
DEDICATION
(SOEPADMO,
(SOEPADMO
JULIA
& &SAW)
GO)
Manokaran, N., J.V. LaFrankie, K.M. Kochummen, E.S. Quah, J.E. Klahn, P.S. Ashton &
S.P. Hubbell (1992). FRIM Research Data No. 1: 1454.
Ng., F.S.P. & L.G. Saw (1999). Fl. Males. Bull. 12(5): 251253.
Van Steenis-Kruseman, M.J. (1974). Fl. Males. 8(1): LV.
Wong, K.M. (1987). Gard. Bull. Sing. 40(1): 1530.
Publications
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
Kochummen, K.M. (1960). Manual of Malayan Timber Trees. No. 7. Dilleniaceae.

FRI Research Pamphlet No. 32.
Kochummen, K.M. (1960). List of tree species at Forest Research Institute, Kepong.
Malay. For. 23: 230235.
Kochummen, K.M. (1961). Precocious flowering (paedogenesis) in Dipterocarpaceae.
Malay. For. 24: 236.
Kochummen, K.M. (1962). Dipterocarpus sarawakensis v. Slooten A new record
for Malaya. Malay. For. 25: 163.
Kochummen, K.M. (1963). Keys for the botanical identification of non-dipterocarp
timber trees of Malaya, based on field characters. FRI Research Pamphlet No. 43:
1119.
Kochummen, K.M. (1964). Dr. Quacks tea leaves (Orthosiphon aristatus (Bl.) Miq.).
Malay. Nat. J. 18: 7374.
Kochummen, K.M. (1964). First revisionPocket Check List of Timber Trees by J.
Wyatt-Smith. Malay. For. Rec. No. 17.
Kochummen, K.M. (1966). Natural plant succession after farming in Sg. Kroh. Malay.
For. 29: 170181.
Kochummen, K.M. (1969). Fifty years of forest botany in Malaya. Malay. For. 32:
6769.
Kochummen, K.M., F.S.P. Ng & T.C. Whitmore (1970). Notes on the Systematy of
Malayan Phanerogams. VIX. Fed. Mus. J. n.s. 13: 133137.
Kochummen, K.M. (1972). Notes on the Systematy of Malayan Phanerogams. XI:
Annonaceae. Gard. Bull. Sing. 26: 49; XVII: Tiliaceae, ibid. 5961.
Kochummen, K.M. (1972). Alangiaceae. In T.C. Whitmore (ed.). Tree Flora of Malaya
1: 5660; Annonaceae, ibid.: 6199; Bombacaceae; ibid.: 100120; Burseraceae, ibid.:
121155; Combretaceae, ibid.: 172178; Dilleniaceae, ibid.: 183193; Malvaceae,
ibid.: 309314; Nyssaceae, ibid.: 346.
Kochummen, K.M. & T.C. Whitmore (1972). Celastraceae. In T.C. Whitmore (ed.).
Tree Flora of Malaya 1: 156171.
Kochummen, K.M. & T.C. Whitmore (1973). Notes on the Systematy of Malayan
Phanerogams. XVIIIXXII. Gard. Bull. Sing. 26: 269287.
Kochummen, K.M. (1973). Hypericaceae. In T.C. Whitmore (ed.). Tree Flora of
Malaya 2: 248252; Loganiaceae, ibid.: 267275; Proteaceae, ibid.: 311320;
Simaroubaceae, ibid.: 345352; Sterculiaceae, ibid.: 353382; Tiliaceae, ibid.: 392
412.
xiii
16. Kochummen, K.M. (1973). A note on the genus Eugenia in Malaya. Malay. For. 36:
6675.
17. Wong, T.M. & K.M. Kochummen (1973). The identification of common commercial
logs in Peninsular Malaysia. Malay. For. 36: 164184.
18. Kochummen, K.M. (1973). Lesser-known timber trees of Malaysia. In E. Soepadmo
& K.G. Singh (eds.). Proc. Symp. Biol. Res. & Nat. Dev., Malay. Nat. Soc., Kuala
Lumpur: 123129.
19. Kochummen, K.M. (1976). Notes on the Systematy of Malayan Phanerogams. XXIII:
Myrtaceae. Gard. Bull. Sing. 28: 227229.
20. Kochummen, K.M. & F.S.P. Ng (1977). Natural plant succession after farming in
Kepong. Malay. For. 40: 6078.
21. Kochummen, K.M. (1978). Notes on the Systematy of Malayan Phanerogams. XXVIII
XXIX. Moraceae and Verbenaceae. Malay. For. 41: 2931.
22. Kochummen, K.M. (1978). Bignoniaceae. In F.S.P. Ng (ed.). Tree Flora of Malaya
3: 3644; Connaraceae, ibid.: 4752; Cornaceae, ibid.: 5355; Icacinaceae, ibid.: 108
118; Moraceae, ibid.: 119168; Myrtaceae, ibid.: 167254; Symplocaceae, ibid.: 266
274; Verbenaceae, ibid.: 297313.
23. Kochummen, K.M. (1979). Second RevisionPocket Check List of Timber Trees
by J. Wyatt-Smith. Malay. For. Rec. No. 17.
24. Kochummen, K.M. (1980). Botanical name changes of some common Malayan timber
trees. Malay. For. 43: 117118.
25. Kochummen, K.M. (1980). The occurrence of Agathis borneensis Warburg (damar
minyak) in heath forest in Peninsular Malaysia. Malay. For. 43: 119123
26. Gianno, R. & K.M. Kochummen (1981). Notes on some minor forest products. Malay.
For. 44: 566568.
27. Kochummen, K.M. (1982). Effects of elevation on vegetation on Gunung Jerai, Kedah.
FRI Research Pamphlet No. 87: 1129.
28. Kochummen, K.M. (1983). Notes on the Systematy of Malayan Phanerogams. XXX:
Anacardiaceae. Gard. Bull. Sing. 36: 187196.
29. Kochummen, K.M. & K.M. Wong (1984). A new species of Alstonia (Apocynaceae)
from the Malay Peninsula and some comments on the genus. Blumea 29: 513522.
30. Manokaran, N. & K.M. Kochummen (1987). Recruitment, growth and mortality of
tree species in a lowland dipterocarp forest in Peninsular Malaysia. J. Trop. Ecol. 3:
315330.
31. Kochummen, K.M. (1989). Anacardiaceae. In F.S.P. Ng (ed.). Tree Flora of Malaya
4: 957; Lauraceae, ibid.: 98178; Rhizophoraceae, ibid.: 302323.
32. Manokaran, N., J.V. LaFrankie, K.M. Kochummen, E.S. Quah, J.E. Klahn, P.S. Ashton
& S.P. Hubbell (1990). Methodology for the fifty-hectare research plot at Pasoh Forest
Reserve. FRIM Research Pamphlet No. 104: 169.
33. Kochummen, K.M., J.V. LaFrankie & N. Manokaran (1990). Floristic composition of
Pasoh Forest Reserve, a lowland rain forest in Peninsular Malaysia. J. Trop. For. Sci.
3: 113.
34. Manokaran, N. & K.M. Kochummen (1990). A re-examination of data on structure
and floristic composition of hill and lowland dipterocarp forest in Peninsular Malaysia.
Malay. Nat. J. 44: 6175.
35. Kochummen, K.M. (1991). Notes on the Systematy of Malayan Phanerogams. XXXI:
Lauraceae. Gard. Bull. Sing. 43: 2326.
36. Saw, L.G., J.V. LaFrankie, K.M. Kochummen & S.K. Yap (1991). Trees bearing edible
xiv
FAGACEAE
DEDICATION
(SOEPADMO,
(SOEPADMO
JULIA
& &SAW)
GO)
fruits in a Malaysian rain forest. Econ. Bot. 45: 120136.

37. Kochummen, K.M., J.V. LaFrankie & N. Manokaran (1992). Floristic composition of
Pasoh Forest Reserve, a lowland rainforest in Peninsular Malaysia. Malay. Nat. J.
45: 545554.
38. Manokaran, N., J.V. LaFrankie, K.M. Kochummen, E.S. Quah, J.E. Klahn, P.S. Ashton
& S.P. Hubbell (1992). Stand table and distribution of species in the 50-ha research
plot at Pasoh Forest Reserve. FRIM Research Data No. 1: 1454.
39. Kochummen, K.M., W.C. Wong, S. Sudo & F.T. Frietema (1994). Hopea. In I.
Soerianegara & R.H.M.J. Lemmens (eds.). PROSEA 5(1). Timber trees: Major
commercial timbers: 238264; Shorea, ibid.: 384434.
40. Kochummen, K.M. (1994). Taxonomic notes on the Celastraceae of Sabah and
Sarawak. Sandakania No. 5: 5166.
41. Kochummen, K.M. (1994). New species of Burseraceae from Sabah and Sarawak.
Sandakania No. 5: 7391.
42. Kochummen, K.M., R.B. Miller & M.S.M. Sosef (1995). Canarium. In R.H.M.J.
Lemmens, I. Soerianegara & W.C. Wong (eds.). PROSEA 5(2). Timber trees: Minor
commercial timbers: 92108; Santiria, ibid.: 416423.
43. Kochummen, K.M., W.C. Wong & Ani Sulaiman (1995). Kokoona. In R.H.M.J.
Lemmens, I. Soerianegara & W.C. Wong (eds.). PROSEA 5(2). Timber trees: Minor
commercial timbers: 273277.
44. Kochummen, K.M. (1995). Burseraceae. In E. Soepadmo & K.M. Wong (eds.). Tree
Flora of Sabah and Sarawak 1: 45100; Celastraceae, ibid.: 107154; Ochnaceae,
ibid.: 257270.
45. Kochummen, K.M. (1996). New species of Anacardiaceae and a new variety of
Parishia maingayi from Sabah and Sarawak. Sandakania No. 7: 81111.
46. Kochummen, K.M. (1996). Anacardiaceae. In E. Soepadmo, K.M. Wong & L.G. Saw
(eds.). Tree Flora of Sabah and Sarawak 2: 192.
47. Kochummen, K.M. (1997). Tree Flora of Pasoh Forest. Malay. For. Rec. 44: 1
461.
48. Kochummen, K.M. (1998). New species and variety of Moraceae from Malaysia. Gard.
Bull. Sing. 50: 197219.
49. Kochummen, K.M. (1998). Dacryodes. In M.S.M. Sosef, L.T. Hong & S.
Prawirohatmodjo (eds.). PROSEA 5(3). Timber trees: Lesser-known timbers: 178
180; Euonymus, ibid.: 225226; Gomphia, ibid.: 261262; Melanochyla, ibid.: 362
437; Pimelodendron, ibid.: 446448.
50. Kochummen, K.M. & J. Ilic (1998). Brackenridgea. In M.S.M. Sosef, L.T. Hong & S.
Prawirohatmodjo (eds.). PROSEA 5(3). Timber trees: Lesser-known timbers: 116
117; Haplolobus, ibid.: 281282; Perrottetia, ibid.: 446448.
51. Kochummen, K.M. (1998). Timber trees. In E. Soepadmo (ed.). The Encyclopedia of
Malaysia 2 Plants: 7879.
52. Kochummen, K.M. (1999). Third RevisionPocket Check List of Timber Trees by
J. Wyatt-Smith. Malay. For. Rec. No. 17: 1367.
53. Kochummen, K.M. & Rusea Go (2000). Moraceae. In E. Soepadmo & L.G. Saw (eds.).
Tree Flora of Sabah and Sarawak 3: 177324.
Eponymy
Alseodaphne kochummenii Kosterm. (Lauraceae). Candollea 28 (1973) 115.
xv
Diospyros kochummenii Ng (Ebenaceae). Malay. For. 40 (1977) 227.

Erythroxylum kochummenii Ng (Erythroxylaceae). Gard. Bull. Sing. 28 (1976) 235.
Kochummenia K.M. Wong (Rubiaceae). Malay. Nat. J. 38 (1984) 31.
Lithocarpus kochummenii S. Julia & Soepadmo (Fagaceae). Gard. Bull. Sing. 50 (1998)
134.
Mesua kochummenii Whitmore (Clusiaceae). Gard. Bull. Sing. 26 (1973) 280.
E. Soepadmo
L.G. Saw
December 1999
xvi
FAGACEAE
E. Soepadmo, S. Julia1 & Rusea Go2
Forest Research Institute Malaysia (FRIM), Kepong, Malaysia
Dumortier, Ann. Fam. Pl. (1829) 59 (Fagineae); A. De Candolle, Prodr. 16, 2 (1864) 1 (Cupuliferae);
Oersted, Kngl. Danske Vid. Selsk. Skrift. 5, 9 (1871) 351 (Cupuliferae); Prantl in Engler & Prantl,
Nat. Pflz. Fam. 3, 1 (1889) 52 (Fagaceae); King, Ann. Roy. Bot. Gard. Calc. 2 (1889) 17; Gamble, J.
As. Soc. Beng. 75, 2 (1915) 406; Koidzumi, Bot. Mag. Tokyo 30 (1916) 186; Rehder, J. Arn. Arb. 1
(1919) 121, ibid. 10 (1929) 132; Merrill, EB (1921) 210; Ridley, FMP 3 (1924) 372; Masamune, EPB
(1942) 232; Barnett, Trans. Proc. Bot. Soc. Edinb. 34 (1942) 327, ibid. 34 (1944) 159; Backer &
Bakhuizen f., FJ 2 (1965) 2; Soepadmo, FM 1, 7 (1972) 265; Cockburn, TFM 1 (1972) 196, TS 1
(1976) 85; Anderson, CLTS (1980) 197; Corner, WSTM 3rd. edition 1 (1988) 327; Whitmore, Tantra
& Sutisna CLK 2, 1 (1990) 152; Kubitzki in Kubitzki et al. (eds.), Fam. & Gen. Vasc. Pl. 2 (1993)
301; Turner, Gard. Bull. Sing. 47 (1995) 234; Coode et al. (eds.), CLBD (1996) 123; Argent et al.
(eds.), MNDT-CK 1 (1997) 237.
Monoecious, evergreen trees or shrubs; buttresses or stilt-roots sometimes present; growth

mode flush-wise, with perular buds. Indumentum comprising simple, stellate or fascicled
hairs. Stipules caducous or rather long persistent, inter- or extra-petiolar. Leaves simple,
spirally arranged, in whorls of 3 (Trigonobalanus), distichous (a few species of
Castanopsis), or crowded near the top of twigs (species of Quercus), pinnately veined,
margin entire, denticulate, serrulate, crenate or lobed. Inflorescences unbranched or
branched, bracteate spikes; male, female, androgynous, or mixed. Male inflorescences
erect or pendulous, in the axils of lower (proximal) leaves or in the axils of reduced
leaves and crowded together in paniculate clustes on lateral or subterminal new shoots.
Male flowers solitary or in dichasial clusters of 37(30) along the rachis, sessile;
perianth campanulate, 6(9)-lobed; stamens (4)612(15), filaments filiform, exserted,
free; anthers reniform or ovoid, dorsi- or basifixed, longitudinally dehiscent; pistillode
absent or present, if present densely hairy. Female, androgynous and mixed inflorescences
erect, in the axils of upper (distal) leaves or on the lower parts of paniculate clusters
on lateral or subterminal new shoots. Female flowers sessile, solitary or in dichasial
clusters of 315 along the rachis; each flower surrounded by its own cupule (Lithocarpus
and Quercus) or the entire cluster of flowers surrounded by a dichasial cupule (Castanopsis
and Trigonobalanus); perianth 6-lobed; staminodes 612 or absent; ovary inferior, 3
6(9)-loculed, ovules anatropous, 2 per locule, apical and pendulous; styles as many
as ovary locules, terete, conical, or cylindrical, stigmas capitate (Quercus and
Trigonobalanus) or punctiform (Castanopsis and Lithocarpus). Cupules (woody or horny
structures covering or substending flowers/fruits) sessile or stalked, solitary or in
dichasial clusters along the rachis, saucer- to cup-shaped or completely enclosing the
Current address: 1Sarawak Biodiversity Centre, Kuching, Malaysia; 2Universiti Putra Malaysia, Serdang,
Malaysia
fruit, at maturity indehiscent or splitting into 28 (or more) equal or unequal segments;
external wall muricate, scaly, spiny, tubercled, or with concentric or spiral lamellae,
or rarely almost smooth. Fruits indehiscent nuts, 13-loculed, sometimes falsely
multiseptate, rounded to sharply 3-angular in cross-section, at base with flat, concave
or convex circular or rounded-triangular scar. Seeds without endosperm; embryo large,
cotyledons large, flat-convex, plicate or ruminate; germination epigeal or hypogeal.
Distribution. About eight genera with more than 800 species, distributed both in the
temperate (up to 62N and 33S) and tropical regions but mostly in the northern
hemisphere. In the Malesian region, there are 5 genera with about 180 species, distributed
from Peninsular Malaysia southeast- and eastwards to New Britain and the Louisiades
Islands in the Pacific. The family is poorly represented in the seasonally dry parts of
C and E Java, and absent in the Lesser Sunda Islands, northern parts of Maluku, New
Ireland and the Solomon Islands. In Sabah and Sarawak, the family is represented by
4 genera comprising 100 species. Of these 49 species are endemic to Borneo, and 17
species to Sabah and Sarawak.
Ecology. In Sabah and Sarawak, members of the Fagaceae are common constituents
of the understorey, main canopy and rarely emergent layers of the lowland to montane
forests, from sea level to about 3000 m altitude. Most species are however, found in
hill mixed dipterocarp to lower montane forests, at altitude ranging from 500 to 1800
m, on a variety of soils. A few species, e.g. Lithocarpus bullatus, L. tawaiensis, Q.
kinabaluensis, and Q. lowii are confined to ultrabasic soils, whereas others, e.g.
Castanopsis foxworthyi and L. andersonii may be found in kerangas and peatswamp
forests as well as in forests growing on well-drained, mineral-rich soils. A few species,
such as Lithocarpus bennettii, L. conocarpus, L. papillifer, L. rassa and L. sundaicus
occasionally occur in limestone habitats. Growth mode is flush-wise, and it is common
for a reproductively matured tree to undergo multiple flushing during a particular growing
period. Flowering is seasonal but in most species it is synchronous among reproducing
species and individuals. Inflorescence primordia usually appear at the end of the rainy
season which in Sabah and Sarawak takes place during the months of December through
March. Floral anthesis occurs during the following dry season (AprilJune), and mature
fruits are produced about 5 months later. In Castanopsis, Lithocarpus and Trigonobalanus
pollination is affected by insects, while in Quercus is by wind. Dispersal of fruits (acorns
or nuts) is carried out by arboreal vertebrates such as squirrels. Seeds sterility and
mortality is relatively high, with the former is due to unsuccessful fertilization and
the latter to predation and attack by weevils and fungi.
Uses. The medium to heavy hardwood timber of Castanopsis is traded as berangan,
and that of Lithocarpus and Quercus as mempening. The timber is suitable for medium
to heavy construction under cover, such as house post, beam joineries, and flooring.
Duly processed, it is also suitable for furniture, cabinet making, interior fittings, panelling,
and for manufacturing plywood and sliced veneers. Locally the wood is used for fence
post, mining props, shingle, boat building, and for making tool handles, rice pounder,
poles for carts, and for firewood and charcoal. Branches and stems are used as bed
logs in mushroom cultivation. The bark of most species contains a high percentage
of tannin and is occasionally used to tan leather and to dye rattan and cotton fabrics.
The nuts of some species of Castanopsis, e.g. C. costata, C. javanica, C. motleyana,
and C. oviformis are edible, and consumed after boiling or roasting.
2
Taxonomy and Classification. According to the widely accepted taxonomic concepts

based largely on gross morphological attributes, the family Fagaceae is classified into
three distinct subfamilies, namely the Castaneoideae comprising Castanea, Castanopsis,
Chrysolepis and Lithocarpus, the Fagoideae with Fagus and Nothofagus, and the
Quercoideae, consisting of Quercus and Trigonobalanus. In the recent years, however,
a number of authors have either proposed to split the family into two distinct ones,
i.e. the Fagaceae s.s. to include Castanea, Castanopsis, Chrysolepis, Fagus, Lithocarpus,
Quercus and Trigonobalanus, and Nothofagaceae s.s. comprising the genus Nothofagus,
or to split a few genera such as Quercus into two smaller genera, Quercus s.s. and
Cyclobalanopsis (Nixon, K.C. & W.L. Crepet, Am. J. Bot. 76 (1989) 842855), and
Trigonobalanus into three smaller genera, Trigonobalanus s.s., Formanodendron and
Colombobalanus (Nixon, K.C. & W.L. Crepet, Am. J. Bot. 76 (1989) 828841). At
subfamily level, proposals have also been made to rearrange the genera into different
subfamilies and tribes. For practical reasons, however, the current account follows
Kubitzkis classification (1993).
Key to genera
(based on vegetative and field characters)
1. Leaves in whorls of 3. Trees coppicing.............4. Trigonobalanus
Leaves spirally arranged or distichous. Trees not coppicing (except in Castanopsis
acuminatissima)..............2
2. Petiole slender, distinctly thickened at the base, often kneed. Inner bark not growing
into the sapwood. Wood with exclusively narrow-uniseriate rays..........1. Castanopsis
Petiole thickened throughout its length or slender and thickened at the base but
never kneed. Inner bark with ridges penetrating the sapwood. Wood with both narrowuniseriate and broad-multiseriate rays..............3
3. Leaves usually crowded near the ends of twigs; margin usually serrate or serrulate
at least in the apical half, rarely entire. Terminal buds crowded at the ends of twigs.
Petiole always thickened at the base..................3. Quercus
Leaves not crowded near the ends of twigs; margin always entire. Terminal buds
solitary. Petiole thickened throughout its length............2. Lithocarpus
Key to genera
(for flowering and/or fruiting specimens)
1. Male inflorescences pendent, usually unbranched. Male flowers without pistillode.
Female flowers usually without staminodes................3. Quercus
Male inflorescences erect or pendent (Trigonobalanus), usually branched. Male
flowers with pistillode. Female flowers with staminodes...........2
2. Male flowers with basifixed anthers. Stigmas capitate. Cupules open, 312-lobed.
Fruits trigonous in cross-section.....................4. Trigonobalanus
Male flowers with dorsifixed anthers. Stigmas minute, punctiform. Cupule open but
not lobed, or completely covering the fruits, indehiscent or dehiscent into a definite
number of segments. Fruits not trigonous in cross-section........................3
3
3. Cupule completely covering (1)3(7) fruits; external wall tubercled or covered with
rigid and woody spines, at maturity splitting into 24(8) segments or
irregularly....................1. Castanopsis
Cupule open cup- or saucer-shaped, or if completely covering the fruit then
indehiscent, enclosing or substending a solitary fruit; outer wall lamellate, scaly,
or at most covered with flexible, slender, spine-like appendages......2. Lithocarpus
1. CASTANOPSIS (D. Don) Spach

(resembling Castanea = the Spanish chestnut)
berangan (Malay)
Hist. Veg. Phan. 2 (1842) 185; Merrill, EB (1921) 210; Masamune, EPB (1942) 232; Burkill, EPMP
1 (1966) 492; Soepadmo, Reinwardtia 7 (1968) 384, FM 1, 7 (1972) 294; Cockburn, TFM 1 (1972)
199, TS 1 (1976) 85; Anderson, CLTS (1980) 197; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990)
152; Kubitzki in Kubitzki et al. (eds.), Fam. & Gen. Vasc. Pl. 2 (1993) 307; Kessler & Sidiyasa,
Trees of the Balikpapan-Samarinda Area, East Kalimantan (1994) 136; PROSEA 5, 2 (1995) 108;
Coode et al. (eds.), CLBD (1996) 123; Argent et al. (eds.), MNDT-CK 1 (1997) 237. Basionym:
Quercus L. subdiv. Castanopsis D. Don, Prod. Fl. Nep. (1825) 275. Synonym: Callaeocarpus
Miq., Pl. Jungh. (1851) 13.
Trees, rarely buttressed. Wood rays narrow and uniseriate. Bark smooth or fissured
or sometimes lenticellate; inner bark granular or fibrous, not growing into the sapwood.
Twigs glabrous or hairy, lenticellate or smooth. Stipules extrapetiolar, small, caducous.
Terminal bud solitary. Leaves spirally arranged or distichous, margin entire or rarely
toothed above the middle (only in Castanopsis acuminatissima); petiole thickened and
usually kneed at the base. Inflorescences erect, male, female, androgynous or mixed,
densely stellate-pubescent. Male inflorescences in lateral or subterminal dense paniculate
clusters on the new shoots or solitary in the axils of lower (proximal) leaves, branched
or unbranched; bracts and bracteoles small, persistent. Male flowers solitary or in
dichasial clusters of 37 along the rachis or in an androgynous inflorescences along
the upper parts of the rachis; perianth 6-lobed; stamens (7)12, filaments glabrous; anthers
dorsifixed, reniform, 2-loculed, 4-lobed; pistillode rudimentary or well developed, more
or less rounded, densely woolly pubescent. Female, androgynous or mixed inflorescences
unbranched, solitary in the axils of upper (distal) leaves or in the upper parts of paniculate
clusters on the new shoots; bracts and bracteoles usually larger and thicker than those
of male inflorescences, persistent. Female flowers solitary or in clusters of 35 along
the rachis or in an androgynous inflorescences along the basal parts of the rachis; perianth
6-lobed; staminodes 1012, rudimentary; styles 3, conical or cylindrical, straight or
recurved, densely hairy at the base; stigma minute, terminal and punctiform. Cupules
solitary along the rachis, completely enclosing 14(7) nuts, surface variously spiny
or set with short tubercles or almost smooth, at maturity spliting irregularly or into
a regular number of segments; spines branched or unbranched, woody and rigid, flat
to triangular, straight or recurved, glabrous or pubescent, regularly or irregularly arranged
on the cupule. Fruits (nuts) more or less angular in cross-section; wall woody, completely
or partly adnate to the cupule. Seed germination hypogeal.
Distribution. About 120 species, distributed in NE India (Nepal, Bhutan, Assam), Burma,
China (except N and NW parts), Korea, Japan, Formosa, Hainan, Indo-China, Thailand,
and Malesia. In Malesia, throughout most islands except the eastern half of Java and
in the Lesser Sunda Islands. The largest number of species are found in Borneo. In
Sabah and Sarawak, the genus is represented by 21 species, of which 10 species are
Bornean endemics.
Ecology. In primary and old secondary forests, from sea level up to 2500 m altitude,
on different types of soils except limestone.
Uses. The nuts of several species are edible. The bark yields tannin and dye. The timber
is used in medium to heavy construction work and in the manufacture of furniture,
packing cases and pallets.
Key to Castanopsis species

1.
Leaves narrowly ovate to lanceolate, margin toothed in the upper half. Trees with
coppicing habit.....................1. C. acuminatissima
Leaves not so, margin entire, plane or revolute. Trees not coppicing...2
2.
Leaves glabrous on both surfaces...3

Leaves sparsely to densely hairy below........10
3.
Intercostal venation lax, faintly prominent on both surfaces..4

Intercostal venation dense, distinctly prominent on both surfaces.....5
4.
Cupule subsessile, spines unbranched. Lateral veins disappearing near the leaf margin.
Twig smooth.....7. C. endertii
Cupule with stalk c. 0.5 cm long, spines much-branched. Lateral veins joining near
the leaf margin. Twig sparsely minute-lenticellate...8. C. evansii
5.
Leaves narrower, with a length:width ratio of 34:1. Cupule tubercled or almost

smooth........6
Leaves wider, with a length:width ratio of 23:1. Cupule variously spiny....7
6.
Cupule almost smooth; stalk 11.5 cm long. Twig sparsely to densely lenticellate.
Leaves elliptic to oblong; lateral veins joining near the leaf margin; petiole 15
20 mm long.........20. C. pedunculata
Cupule tubercled, subsessile or with stalk 0.51 cm long. Twig smooth. Leaves
ovate, lanceolate or elliptic; lateral veins disappearing near the leaf margin; petiole
813 mm long............21. C. psilophylla
7.
Cupule globose or subglobose. Petiole glabrous.......8

Cupule ovoid, obovoid or ellipsoid. Petiole hairy.....9
8.
Cupule densely spiny; spines unbranched, slender; nuts 1 in each cupule. Leaves
coriaceous, margin plane, lateral veins usually less than 7 pairs, joining near the
leaf margin.......2. C. borneensis

Cupule sparsely spiny; spines branched, flat to triangular; nuts 3 in each cupule.
Leaves thick-coriaceous, margin revolute, lateral veins usually more than 7 pairs,
disappearing near the leaf margin........9. C. foxworthyi
9. Stipules usually persistent, 79 mm long. Leaves elliptic, margin plane, acumen
810 mm long; lateral veins disappearing near the leaf margin.......4. C. clemensii
Stipules caducous, 35 mm long. Leaves elliptic, oblong or ovate, margin revolute,
acumen c. 5 mm long; lateral veins joining near the leaf margin..18. C. oviformis
10. Twig densely covered with large lenticels.......11
Twig sparsely covered with minute or large lenticels or smooth or finely fissured....12
11. Petiole 2035 mm long. Cupule depressed subglobose, 2.54 cm in diameter, shortly
spiny or tubercled; stalk thick and twisted; nut wall thick and woody. Leaf apex
obtuse ........13. C. lucida
Petiole (4)68(12) mm long. Cupule discoid, asymmetrical, 11.5 cm in diameter,
spiny; stalk slender and straight; nut wall thin. Leaf apex acuminate, acumen 10
15 mm..........12. C. javanica
12. Leaf apex acute or short-acuminate, acumen c. 5 mm long..13
Leaf apex long-acuminate, acumen 1020 mm long, rarely acute.......17
13. Cupule stalk 0.50.7 cm long.........................14
Cupule sessile or subsessile........15
14. Lateral veins closely spaced. Nuts 1 in each cupule, free part glabrous...................
................................................................................................................6. C. densinervia
Lateral veins distantly spaced. Nuts 23 in each cupule, free part densely hairy...
..........5. C. costata
15. Leaves ovate or elliptic, margin slightly revolute; midrib thin, lateral veins 57
pairs............17. C. oligoneura
Leaves elliptic-oblong or obovate, margin strongly revolute, midrib broad and
strongly raised below, lateral veins 1015 pairs...............16
16. Mature cupule 58 35.5 cm; nuts 1 in each cupule, free apical part glabrous.
Leaves elliptic-oblong, apex acute14. C. megacarpa
Mature cupule 23 1.52.5 cm; nuts 23 in each cupule, sparsely pilose at the
apex. Leaves obovate or rarely elliptic-oblong, apex short-acuminate....16. C. motleyana
17. Leaves 58 1.52.5 cm, linear to lanceolate. Nuts 23 in each cupule, equally
developed...........15. C. microphylla
Leaves (7)1117 (3)47 cm, ovate, elliptic, oblong, or ovate-lanceolate. Nuts
usually only 1 well-developed in each cupule................18
18. Cupule stalk 0.51 cm long. Petiole strongly kneed at the base.19
Cupule subsessile. Petiole slightly kneed at the base..20
19. Cupule subglobose, 2.53 22.5 cm; spines slender, irregularly arranged. Leaves
thick-coriaceous; lateral veins disappearing near the leaf margin....10. C. fulva
Cupule obovoid to ellipsoid, 44.5 33.5 cm; spines sturdy, flat to triangular,
regularly arranged. Leaves thin-coriaceous; lateral veins joining near the leaf margin
............19. C. paucispina
20. Leaves sparsely to densely covered with appressed stellate and hirsute hairs below;
intercostal venation dense, scalariform, prominent on both surfaces. Cupule ovoid
to ellipsoid, 710 4.57 cm; nut wall thick and woody, most part adnate to
the cupule.....11. C. hypophoenicea
Leaves sparsely tomentose below; intercostal venation lax, scalariform, weakly raised
or almost obscure on both surfaces. Cupule ovoid to conical, c. 1.5 1 cm; nut
wall thin, most part free from the cupule..............3. C. buruana
1. Castanopsis acuminatissima (Blume) A. DC.

(Latin, acuminatissimus = extremely long-pointed; the leaf tip)
J. Bot. 1 (1863) 182; Soepadmo l.c. (1968) 385, l.c. (1972) 307; Cockburn l.c. (1972) 201, l.c. (1976)
89; Whitmore, Tantra & Sutisna l.c. 152; PROSEA l.c. 108; Turner l.c. 234. Basionym: Castanea
acuminatissima Blume., Mus. Bot.-Lugd. Bat. 1 (1850) 283. Type: Blume s.n. (= RHL Sheet No.
901310451), Java (holotype L). Synonyms: Castanea sessilifolia Blume l.c. 284; Pasania acuminatissima
(Blume) Oerst., Vid. Medd. Naturh. For. Kjb. 8 (1867) 84; Quercus varingaefolia Miq., Pl. Jungh.
(1851) 12; Quercus junghuhnii Miq., Fl. Ind. Bat. 1, 1 (1856) 853; Castanopsis junghuhnii (Miq.)
Markgr., Bot. Jahrb. 59 (1924) 62; Quercus fagiformis Jungh., Bonplandia 6 (1858) 83; Synaedrys
fagiformis (Jungh.) Koidzumi, Bot. Mag. Tokyo 30 (1916) 187; Castanopsis schlenkerae Bailey, Queensl.
Agr. J. 22 (1909) 149.
Tree up to 30 m tall, 50 cm diameter, often coppicing. Bark flaky or sometimes smooth,

yellowish grey; inner bark granular, brownish yellow. Sapwood yellowish. Twigs densely
pubescent when young, becoming glabrous, smooth or sometimes minute-lenticellate.
Stipules triangular, c. 1.5 1 mm. Leaves coriaceous, glabrous above, sparsely brown
stellate-tomentose below; narrowly ovate to lanceolate, 6.511 23 cm, base acute,
rounded or attenuate, margin plane or sometimes revolute, toothed above the middle,
apex long-acuminate, acumen 1020(25) mm long; midrib flat or impressed above,
raised below; lateral veins 610 pairs, thin and weakly raised on both sides, disappearing
near the leaf margin; intercostal venation scalariform, obscure on both surfaces; petiole
57 mm long, glabrous. Inflorescences male, female or mixed. Male inflorescences
712 cm long; bracts ovate, 12 11.5 mm; bracteoles reduced to a cluster of hirsute
hairs. Male flowers solitary but densely arranged along the rachis; perianth lobes obtuse,
c. 2 1 mm; stamens 12, filaments c. 3 mm long; pistillode 11.5 mm diameter. Female
or mixed inflorescences 816 cm long; bracts and bracteoles ovate, 12 0.71.2
mm. Female flowers solitary along the rachis; perianth lobes acute, c. 2.5 1 mm;
staminodes 12; styles cylindrical, recurved, 11.5 mm long. Cupules (sub)sessile,
asymmetrical, ovoid, 1.21.5 0.50.7(1.2) cm, densely pubescent, densely spiny;
spines up to c. 2 mm long, unbranched, flat to triangular, slightly recurved, densely
pubescent, sparsely and irregularly or regularly arranged in lines. Nuts 1 in each cupule,
ovoid to conical, 11.5 0.7 cm, free part densely tomentose; wall thin, most part
free from the cupule.
Distribution. India, Burma, China, Taiwan, Thailand, Indo-China, and Malesia (N Sumatra,
Peninsular Malaysia, Java, Borneo, Sulawesi, New Guinea, and New Britain). In Sabah,
recorded from Kota Marudu, Keningau and Ranau districts. In Sarawak, uncommon, known
only from Sg. Menalio in Marudi district. Not known in Brunei and Kalimantan.
Ecology. In primary lowland mixed dipterocarp forest to montane forest, at 3002500m
altitude.
Uses. The bark yields tannin and the nuts are edible.
2. Castanopsis borneensis King
Fig. 1.
(of Borneo)
Ann. Bot. Gard. Calc. 2 (1889) 99; Merrill l.c. 210; Masamune l.c. 152; Soepadmo l.c. (1968) 387, l.c.
(1972) 299; Cockburn l.c. (1976) 92; PROSEA l.c. 113; Coode et al. (eds.) l.c. 123; Argent et al. (eds.)
l.c. 238. Type: Beccari PB 1212, Borneo, Sarawak (holotype FI; isotype K).
Tree up to 20 m tall, 30 cm diameter, non-coppicing. Bark smooth or slightly fissured,

pale brown; inner bark fibrous, yellowish brown. Sapwood yellowish. Twigs glabrous,
slightly fissured or smooth or sparsely lenticellate, grey when young. Stipules linear,
23.5 mm long. Leaves coriaceous, glabrous on both surfaces; ovate to elliptic, 7
11 35 cm, length:width ratio 23:1, base rounded or acute, margin entire and plane,
apex acuminate, acumen c. 10 mm long; midrib raised on both surfaces but stronger
below; lateral veins 67 pairs, thin and flat on both surfaces or sometimes weakly
raised below, joining near the leaf margin; intercostal venation dense, subscalariform,
distinct and prominent on both surfaces; petiole 1015 mm long, glabrous. Inflorescences
male and female. Male inflorescences 1015 cm long; bracts ovate, c. 1 mm across;
bracteoles lanceolate, c. 1 0.4 mm. Male flowers in clusters of 3 along the rachis;
perianth lobes acute, c. 1.5 0.5 mm; stamens 68, filaments c. 3 mm long; pistillode
11.5 mm diameter. Female inflorescences c. 10 cm long; bracts and bracteoles ovate,
11.5 0.7 mm. Female flowers solitary along the rachis; perianth lobes acute, c.
1 0.5 mm; staminodes 12; styles conical, straight, 11.5 mm long. Cupules sessile,
subglobose, 1.53 12.5 cm, sparsely pubescent, densely spiny; spines 57 mm long,
unbranched, slender, straight, sparsely pubescent except the tips, densely and irregularly
arranged. Nuts 1 in each cupule, ovoid, 1.52 11.5 cm, free apical part sparsely
pubescent; wall thin, most part adnate to the cupule.
Vernacular names. Sarawakberangan (Malay), berangan padi (Iban).
Distribution. Endemic to Borneo. In Sabah, common in Ranau, Sipitang and Beaufort
districts. In Sarawak, known from Bintulu, Kuching, Miri, and Sibu districts. Also in Brunei.
Ecology. In primary lowland mixed dipterocarp and kerangas forests, up to 200 m altitude.
3. Castanopsis buruana Miq.

(of Buru Island in Maluku)
Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 120; Soepadmo l.c. (1968) 387, l.c. (1972) 307; Cockburn l.c.
vi
(1976) 92; Whitmore, Tantra & Sutisna l.c. 152; PROSEA l.c. 113. Type: Teijsmann HB 1868, Maluku,
Buru, Kajeli (holotype U; isotypes BO, L). Synonyms: Castanea buruana (Miq.) Oerst., Knig.
Danske Vid. Selsk. Skr. 5, 9 (1867) 377; Castanopsis buruana Miq. forma grandifolia Miq. l.c.
(1863) 120.
Tree up to 30 m tall, 60 cm diameter, non-coppicing. Bark smooth or sometimes scaly,

brown; inner bark reddish or whitish. Sapwood white. Twigs sparsely stellate-hairy
when young, late-glabrescent, minutely lenticellate. Stipules linear, triangular, 34
1 mm. Leaves coriaceous, glabrous above, sparsely tomentose below; ovate to elliptic,
(7)1113 34.5 cm, base cuneate to sometimes acute, margin entire and revolute,
apex long-acuminate, acumen c. 20 mm long; midrib impressed above, raised and sparsely
pilose below; lateral veins 810 pairs, flat above, thin and weakly raised below,
disappearing or sometimes joining near the leaf margin; intercostal venation lax,
scalariform, weakly raised or almost obscure on both surfaces; petiole 1118 mm long,
slightly kneed at the base, glabrous. Inflorescences male and female. Male inflorescences
(4)710 cm long; bracts broadly ovate, 11.5 mm across; bracteoles broadly ovate,
c. 0.5 mm across. Male flowers in clusters of 34 along the rachis, densely arranged;
perianth lobes acute, c. 2 1 mm; stamens 12, filaments c. 3 mm long; pistillode
11.5 mm diameter. Female inflorescences 1012 cm long; bracts and bracteoles broadly
ovate, c. 0.5 mm across. Female flowers solitary along the rachis; perianth lobes acute,
c. 1.5 mm across; staminodes 12, well-developed and exceeding the perianth; styles
cylindrical, recurved, 22.5 mm long. Cupules subsessile, ovoid to conical, asymmetrical,
c. 1.5 1 cm, densely pubescent, spiny; spines 35 mm long, unbranched, flat to
triangular, straight, sparsely pubescent except the tips, arranged regularly in lines with
space in between. Nuts 1 in each cupule, rarely 2, ovoid to globose, 11.5 cm across,
free apical part densely pilose; wall thin, most part free from the cupule.
Vernacular name. Sabahkogopon (Dusun, Bundu Tuhan).
Distribution. Borneo, Sulawesi and Maluku. In Sabah, known only from Padang
Pentululungan in Ranau district (two collections) and Kampung Kitau in Keningau
district. So far not known in Sarawak, Brunei and Kalimantan.
Ecology. In primary lowland and hill mixed dipterocarp forests, up to 450 m altitude.
4. Castanopsis clemensii Soepadmo
Fig. 2.
(J. Clemens, 18621936, former chaplain of the American Army, plant collector)
Reinwardtia 7 (1968) 388, l.c. (1972) 302; Cockburn l.c. (1976) 92; Whitmore, Tantra & Sutisna l.c.
152. Type: Brand & Anak SAN 25315, Borneo, Sabah, Ranau district, Sg. Mentaki (holotype L; isotypes
K, SAN, SAR).
Tree up to 30 m tall, 50 cm diameter, non-coppicing. Bark cracked or lenticellate or

sometimes smooth, yellowish white; inner bark fibrous, red-brown turning to dirty brown
on exposure. Sapwood pale yellow. Twigs sparsely stellate-hairy, smooth or minutelenticellate. Stipules ovate, 79 45 mm, usually persistent. Leaves coriaceous,
glabrous on both surfaces; elliptic, 812 44.5 cm, base acute, rounded to subcordate,
sometimes asymmetrical, margin entire and plane, apex acuminate, acumen 810 mm
long; midrib faintly raised above, strongly raised below; lateral veins 810 pairs, flat
10
vi
above, slightly raised below, disappearing near the leaf margin; intercostal venation
dense, subscalariform, distinctly prominent on both surfaces; petiole 1012 mm long,
sparsely stellate-pubescent. Inflorescences male and female. Male inflorescences 4.5
7 cm long; bracts and bracteoles ovate, c. 0.6 mm across. Male flowers in clusters
of 34 along the rachis; perianth lobes obtuse, 1.52 mm across; stamens 79, filaments
c. 2 mm long; pistillode c. 1.5 mm diameter. Female inflorescences 47.5 cm long;
bracts broadly ovate, c. 1.5 1 mm; bracteoles ovate, c. 1 0.5 mm. Female flowers
solitary along the rachis; perianth lobes acute, 0.51 mm across; staminodes 12; styles
cylindrical, recurved, 11.5 mm long. Cupules obovoid or ellipsoid, 1.32.5 11.5
cm, sparsely puberulous, spiny except for the adaxial and abaxial sutures; stalk c. 0.5
cm long; spines 58 mm long, branched at the base or sometimes unbranched, flat,
straight, sparsely pubescent, sparsely and irregularly arranged. Nuts 1 in each cupule,
ellipsoid, 12 1.5 cm, free apical part pubescent; wall thin, most part adnate to the
cupule.
Vernacular name. Sabahkogopon (Dusun).
Distribution. Endemic to Borneo. In Sabah, common in Ranau district and recorded
by a few collections from Tenom, Tambunan and Keningau districts. In Sarawak, reported
from Bario, Limbang (G. Murut) and Sri Aman districts. Not known in Brunei and
Kalimantan.
Ecology. In primary lower montane forest, up to 1300 m altitude.
5. Castanopsis costata (Blume) A. DC.

(Latin, costatus = ribbed; the prominent lateral veins)
J. Bot. 1 (1863) 182; Merrill l.c. 210; Masamune l.c. 232; Burkill l.c. 493; Soepadmo l.c. (1968)
388, l.c. (1972) 312; Cockburn l.c. (1972) 203, l.c. (1976) 89 & 90; Whitmore, Tantra & Sutisna
l.c. 152; PROSEA l.c. 113; Turner l.c. 234; Coode et al. (eds.) l.c. 124; Argent et al. (eds.) l.c. 238.
Basionym: Castanea costata Blume, Mus. Bot. Lugd.-Bat. 1 (1850) 284. Type: Korthlas s.n. (=
RHL Sheet No. 901309120), Sumatra, Bt. Melintang (holotype L). Synonyms: Castanea brevicuspis
Miq., Fl. Ind. Bat. 1, 1 (1858) 866; Castanopsis brevicuspis (Miq.) A. DC. l.c. 182; Castanea
spectabilis Miq. l.c. 866; Castanopsis spectabilis (Miq.) A. DC. l.c. 182; Castanopsis trisperma
Scheff, Nat. Tijd. Ned. Ind., 31 (1870) 362.
Tree up to 36 m tall, 80 cm diameter, non-coppicing. Bark smooth or scaly or sometimes

flaky, whitish; inner bark fibrous, yellowish. Sapwood pale yellow. Twigs pubescent
when young, glabrescent, sparsely minute-lenticellate. Stipules linear, triangular, 58
1 mm. Leaves coriaceous to sometimes thick-coriaceous, glabrous above, sparsely
to densely rusty brown stellate-tomentose or very rarely glabrous below; elliptic-oblong
or ovate, (8)1115(19) 36.5 cm, base abruptly acute to rounded, margin entire,
slightly to strongly revolute, apex acute; midrib flat above, strongly raised below; lateral
veins 815 pairs, distantly spaced, impressed above, raised below, disappearing near
the leaf margin; intercostal venation scalariform, obscure or sometimes prominent above,
slightly prominent below; petiole (6)1015(25) mm long, sparsely stellate-hairy.
Inflorescences male and female. Male inflorescences 913.5 cm long; bracts and
bracteoles ovate, 11.5 1 mm. Male flowers in clusters of 23 or sometimes solitary
12
along the rachis; perianth lobes acute, c. 1.5 0.5 mm; stamens 12, filaments 22.5
mm long; pistillode 11.5 mm diameter. Female inflorescences 6.512 cm long; bracts
and bracteoles broadly ovate, c. 12 1 mm. Female flowers in clusters of 3 or sometimes
solitary along the rachis; perianth lobes acute, c. 1 1 mm; staminodes 12; styles
cylindrical, slightly recurved, 11.5 mm long. Cupules subglobose, 12.5 1.52.5
cm, densely pubescent, dehiscing into 4 segments, spiny; stalk 0.50.7 cm long; spines
712 mm long, branched, rarely unbranched, slender, strongly recurved, sparsely hairy
at the base, densely and irregularly arranged. Nuts 23 in each cupule, conical, 1
2 11.5 cm, free part densely pubescent; wall thin, most part free from the cupule.
Distribution. Sumatra, Peninsular Malaysia, Java, and Borneo. Common and widely
distributed in Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. In primary lowland mixed dipterocarp forest to montane forest, up to 1900m
altitude.
Uses. The timber is locally used for house-building. The nuts are edible.
6. Castanopsis densinervia Soepadmo

(Latin, densus = dense, nervia = nerves; densely veined, the leaves)
Reinwardtia 7 (1968) 389, l.c. (1972) 301; Cockburn l.c. (1976) 89; Whitmore, Tantra & Sutisna
l.c. 152; PROSEA l.c. 114; Argent et al. (eds.) l.c. 238. Type: Meijer SAN 48109, Borneo, Sabah,
Mt. Kinabalu, west of Mesilau Cave (holotype L; isotypes K, SAN).
Tree up to 30 m tall, 50 cm diameter, non-coppicing. Bark smooth, greyish; inner

bark greyish turning to dark grey on exposure. Sapwood dark grey on exposure. Twigs
sparsely covered with large lenticels, sparsely stellate-hairy or glabrous. Leaves thickcoriaceous, glabrous and shining above, densely brown tomentose or pilose below; ovate
or elliptic-oblong, 9.512.5 34.5 cm, base acute, rounded to subcordate, sometimes
asymmetrical, margin entire and revolute, apex acute or shortly acuminate, acumen
c. 5 mm long; midrib flat or impressed above, raised below; lateral veins 1215 pairs,
closely spaced and parallel, impressed above, strongly raised below, disappearing near
the leaf margin; intercostal venation dense, scalariform, obscure above, faintly prominent
below; petiole 1015 mm long, sparsely stellate-hairy. Inflorescences male and female.
Male inflorescences 37 cm long; bracts ovate, 11.5 mm across; bracteoles lanceolate,
c. 0.7 0.3 mm. Male flowers in clusters of 3 along the rachis; perianth lobes acute,
1.52 mm across; stamens 12, filaments c. 2.5 mm long; pistillode 23 mm diameter.
Female inflorescences 34 cm long; bracts and bracteoles ovate, 11.5 mm across.
Female flowers solitary along the rachis; perianth lobes obtuse, c. 0.5 0.5 mm;
staminodes 12; styles conical, straight, 22.5 mm long. Cupules subglobose, 2.54.5
24 cm, glabrous, spiny except for a small part of the adaxial suture; stalk c. 0.5
cm long; spines 710 mm long, branched at the base, slender, straight, sparsely pubescent
at the base, densely but irregularly arranged with space in between. Nuts 1 in each
cupule, ovoid, c. 3.5 3 cm, free apical part glabrous; wall thin, most part adnate
to the cupule.
Distribution. Endemic to Borneo. In Sabah, recorded mainly from Mt. Kinabalu in
13
Ranau and Kimanis road in Keningau districts; so far no record from Sarawak. Also
in NE Kalimantan.
Ecology. In primary hill mixed dipterocarp forest to lower montane forest, at 1000
1800 m altitude.
7. Castanopsis endertii Hatus. ex Soepadmo

(F.H. Endert, 18911953, sometime officer of the Indonesia Forestry Service, Java)
Reinwardtia 7 (1968) 390, l.c. (1972) 301; Whitmore, Tantra & Sutisna l.c. 152; Argent et al. (eds.) l.c.
238. Type: Endert 5196, Borneo, Kalimantan, W Kutai, Kombeng (holotype L; isotype BO).
Tree up to 35 m tall, 60 cm diameter, non-coppicing. Bark smooth with ring-like markings,

slightly fissured, brownish; inner bark granular, greyish. Sapwood yellowish. Twigs
glabrous, smooth. Stipules linear, triangular, 46 2 mm. Leaves thin-coriaceous,
glabrous on both surfaces; elliptic to oblong, (10.5)1318(23) (5)68(10) cm,
base rounded or acute, margin entire and plane, apex acute to shortly acuminate, acumen
c. 5 mm long; midrib slightly raised above, stronger below; lateral veins (7)1012
pairs, flat above, raised below, disappearing near the leaf margin; intercostal venation
lax, scalariform, faintly prominent on both surfaces; petiole (10)1015(25) mm long,
glabrous. Inflorescences male and female. Male inflorescences 1012 cm long; bracts
ovate. Male flowers in clusters of 3 along the rachis; perianth lobes acute, 11.5 mm
across; stamens 12, filaments 22.5 mm long; pistillode c. 2 mm diameter. Female
inflorescences 812 cm long; bracts and bracteoles ovate, 12 0.51 mm. Female
flowers solitary along the rachis; perianth lobes acute, c. 1.5 mm across; staminodes
12; styles conical, straight, c. 1.5 mm long. Cupules subsessile, globose, c. 2.54 cm
across, sparsely pubescent, spiny; spines 713 mm long, unbranched, sturdy, straight
or slightly recurved, sparsely pubescent, sparsely and irregularly arranged. Nuts 1 in
each cupule, globose, 23.5 cm across; wall thin, most part adnate to the cupule.
Vernacular name. Sabahkogopon (Dusun).
Distribution. Endemic to Borneo. Widely distributed throughout Sabah; so far not known
in Brunei and Sarawak. Also in Kalimantan.
Ecology. In primary and disturbed montane forests, up to 2500 m altitude.
Use. The nuts are edible.
8. Castanopsis evansii Elmer

(J.H. Evans, former Governor of Puerto Princesa, Palawan, the Philippines)
Leafl. Philip. Bot. 5 (1913) 1778; Soepadmo l.c. (1968) 391, l.c. (1972) 305; Cockburn l.c. (1976) 92;
Whitmore, Tantra & Sutisna l.c. 153; Kessler & Sidiyasa l.c 138; PROSEA l.c. 114. Type: Elmer
12988, the Philippines, Palawan, Mt. Pulgar (holotype A; isotypes K, L, NY, U, US, WRSL).
Synonyms: Castanopsis woodii Merr., Philip. J. Sc. 29 (1926) 362; Castanopsis elmeri Merr., PEB
(1929) 42.
14
Tree up to 20 m tall, 30 cm diameter, non-coppicing. Bark smooth or sometimes flaky,

greyish; inner bark fibrous, yellow. Sapwood pale yellow. Twigs glabrous, sparsely minutelenticellate. Stipules linear, triangular, 56 2 mm. Leaves coriaceous rarely thickcoriaceous, glabrous on both surfaces; elliptic to lanceolate, 1117 46 cm, base attenuate,
acute to rounded, margin entire and revolute, apex acute; midrib slightly raised above,
stronger below; lateral veins 1114 pairs, flat above, raised below, joining near the leaf
margin; intercostal venation lax, scalariform, thin and faintly prominent on both surfaces;
petiole (6)1220 mm long, glabrous. Inflorescences male, female or mixed. Male
inflorescences (3.5)812 cm long; bracts and bracteoles ovate, 11.5 0.71 mm. Male
flowers solitary or sometimes in clusters of 3 along the rachis; perianth lobes obtuse,
11.5 mm across; stamens 1012, filaments c. 2.5 mm long; pistillode c. 1.5 mm diameter.
Female and mixed inflorescences 610 cm long; bracts and bracteoles broadly ovate,
c. 1 mm across. Female flowers solitary along the rachis; perianth lobes acute, c. 1.5
mm across; staminodes 12; styles conical, recurved, 12 mm long. Cupules subglobose,
obovoid to ellipsoid, 34.5 34 cm, glabrous or sometimes pubescent, spiny; stalk
c. 0.5 mm long; spines 719 mm long, much-branched, slender, straight, glabrous, sparsely
and irregularly arranged. Nuts 1 in each cupule, ovoid, 33.5 cm across, free apical part
densely tomentose; wall thin, most part adnate to the cupule.
Vernacular names. Sabahbuling (Dusun, Banggi), kumpat (Dusun, Rungus).
Distribution. Borneo and the Philippines. Common throughout Sabah; in Sarawak rare,
only known by one collection from G. Gading FR in Lundu district (S. 9645). Also
in SE Kalimantan.
Ecology. In primary and disturbed hill mixed dipterocarp forest, up to 500 m altitude.
9. Castanopsis foxworthyi Schottky
Fig. 3.
(F.W. Foxworthy, 18771950, botanist of the Bureau of Science, Manila and later research officer at the Forest Department of Malay Peninsula)
Bot. Jahrb. 49 (1913) 358; Merrill l.c. (1921) 211; Masamune l.c. 232; Soepadmo l.c. (1968) 391,
l.c. (1972) 309; Cockburn l.c. (1976) 92; Anderson l.c. 198; Whitmore l.c. 153; PROSEA l.c. 114;
Turner l.c. 234; Coode et al. (eds.) l.c. 124. Type: Foxworthy 214 (n.v.), Borneo, Sarawak. Synonym:
Castanopsis kinabaluensis A. Camus, Bull. Soc. Bot. Fr. 75 (1928) 698.
Tree up to 30 m tall, 60 cm diameter, non-coppicing. Bark smooth, blackish brown;

inner bark pale red. Sapwood yellowish white. Twigs sparsely stellate-pubescent, largelenticellate or sometimes finely fissured. Stipules narrowly ovate to elliptic, 68 1
2 mm. Leaves thick-coriaceous, glabrous on both surfaces; elliptic or lanceolate, 7
12(16.5) 47.5 cm, base attenuate, rounded or acute, margin entire and revolute,
apex acuminate, acumen c. 10 mm long; midrib slightly raised above, stronger below;
lateral veins 712 pairs, impressed above, flat or sometimes raised below, disappearing
near the leaf margin; intercostal venation dense, scalariform, prominent on both surfaces;
petiole (8)1015(18) mm long, glabrous. Inflorescences male and female. Male
inflorescences 715 cm long; bracts and bracteoles ovate, c. 1 0.5 mm. Male flowers
in clusters of 3 along the rachis, perianth lobes acute, c. 2 1.5 mm; stamens 12,
15
vi
filaments c. 2.5 mm long; pistillode c. 5 mm diameter. Female inflorescences 810

cm long; bracts ovate, c. 1 0.5 mm; bracteoles broadly ovate, c. 1 mm across. Female
flowers in clusters of 3 along the rachis; perianth lobes acute, c. 1 mm across; staminodes
12; styles conical, straight, c. 1 mm long. Cupules subglobose, 35 23 cm, densely
pubescent, sparsely spiny; stalk c. 0.5 cm long; spines c. 6 mm long, branched at
the base, flat to triangular, straight, sparsely pubescent at the base, sparsely and regularly
arranged with space in between. Nuts 3 in each cupule, ovoid, 12 cm across, free
part densely hairy; wall thin, most part free from the cupule.
Vernacular names. Sarawakberangan lingkau (Iban), berangan paya (Malay); beluing,
maruwit (Tagar).
Distribution. NE Sumatra, Peninsular Malaysia, Singapore, and Borneo. Very common
and widely distributed in Sabah and Sarawak. Also in Brunei.
Ecology. In primary mixed dipterocarp, peatswamp and kerangas forests, to montane
forest, up to 2100 m altitude.
10. Castanopsis fulva Gamble

(Latin, fulvus = tawny; the indumentum)
Kew Bull. (1914) 179; Soepadmo l.c. (1968) 391, l.c. (1972) 312; Cockburn l.c. (1972) 204, l.c. (1976)
89; Anderson l.c. 198; Whitmore, Tantra, Sutisna l.c. 153; PROSEA l.c. 114; Turner l.c. 234; Coode et
al. (eds.) l.c. 124; Argent et al. (eds.) l.c. 240. Lectotype (Soepadmo 1968): Kings Collectors 7751,
Peninsular Malaysia, Perak, Batang Padang (holotype K; isotypes L, SING).
Tree up to 25 m tall, 50 cm diameter, non-coppicing. Bark smooth or slightly fissured;

inner bark granular, red-brown. Sapwood creamy yellow. Twigs velvety hairy or sometimes
glabrescent, finely fissured. Stipules triangular, 23 1 mm. Leaves thick-coriaceous,
glabrous above, sparsely to densely brown stellate-hairy below; elliptic, ovate, to oblong,
1117 47 cm, base acute, attenuate or rounded, margin entire and revolute, apex longacuminate, acumen 1015 mm long; midrib flat above, strongly raised below; lateral veins
813 pairs, flat above, raised below, disappearing near the leaf margin; intercostal venation
scalariform, weakly prominent above, stronger below; petiole 1225 mm long, strongly kneed
at the base, densely stellate-hairy. Inflorescences male, female or mixed. Male inflorescences
(5)1114.5 cm long; bracts ovate, 1.52 11.5 mm; bracteoles ovate, 0.51 mm across.
Male flowers in clusters of 3 along the rachis; perianth lobes acute, 11.5 1 mm; stamens
12, filaments 11.5 mm long; pistillode 1.52 mm diameter. Female and mixed inflorescences
911(18) cm long; bracts and bracteoles broadly ovate, 1.52 mm across. Female flowers
in clusters of 3 along the rachis; perianth lobes obtuse, c. 0.5 mm across; staminodes 12;
styles cylindrical, straight, c. 1 mm long. Cupules subglobose, 2.53 22.5 cm, densely
pubescent, spiny; stalk c. 0.5 cm long; spines c. 5 mm long, much-branched, slender, slightly
recurved or straight, pubescent except the tips, sparsely to densely and irregularly arranged.
Nuts 3 in each cupule but only 1 well-developed, ovoid-conical, 1.52 cm across, free
part pubescent; wall thin, most part free from the cupule.
Vernacular name. Sarawakberangan padi (Iban).
17
vi
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah, recorded from

Beaufort and Ranau districts. In Sarawak, common and widely distributed. Also in
Kalimantan and Brunei.
Ecology. In primary hill mixed dipterocarp and kerangas forests, up to 1100 m altitude.
11. Castanopsis hypophoenicea (Seemen) Soepadmo
Fig. 4.
(Latinised Greek, hypophoeniceus = purple-red or crimsom beneath; the indumentum of

lower leaf surface)
Reinwardtia 7 (1968) 392, l.c. (1972) 305; Cockburn l.c. (1976) 93; Anderson l.c. 198; Whitmore,
Tantra & Sutisna l.c. 153; PROSEA l.c. 115; Coode et al. (eds.) l.c. 153; Argent et al. (eds.) l.c.
240. Basionym: Quercus hypophoenicea Seemen, Bot. Jahrb. 23, Beibl. 57 (1897) 52. Type: Beccari
PB 2287, Borneo, Sarawak (holotype FI; isotypes K, L, P). Synonyms: Lithocarpus hypophoenicea
(Seemen) Barn., Trans. & Proc. Bot. Soc. Edinb. 44 (1944) 171; C. dipersispina Merr., PEB (1929)
41.
Tree up to 30 m tall, 60 cm diameter, non-coppicing. Bark rough or sometimes smooth,

greyish; inner bark fibrous, pale yellow to reddish. Sapwood yellowish white. Twigs
glabrous to sometimes densely yellowish brown pubescent with hirsute and stellate hairs,
sparsely minute-lenticellate. Stipules ovate, 68 45 mm. Leaves coriaceous, rarely
thick-coriaceous, glabrous above, sparsely to densely covered with appressed stellate and
hirsute hairs below; elliptic, oblong or ovate-lanceolate, 1017 47 cm, base rounded,
acute, to attenuate, asymmetrical, margin entire and slightly revolute, apex long-acuminate,
acumen c. 15 mm long; midrib impressed above, raised below; lateral veins 1112 pairs,
flat or impressed above, raised below, joining near the leaf margin; intercostal venation
dense, scalariform, prominent on both surfaces, stronger below; petiole 717 mm long,
slightly kneed at the base, densely covered with stellate and hirsute hairs. Inflorescences
male, female or mixed. Male inflorescences (5)8.511 cm long; bracts and bracteoles
ovate to lanceolate, 0.71 0.7 mm. Male flowers in clusters of 35 along the rachis;
perianth lobes obtuse, 11.5 mm across; stamens 10, filaments c. 1.5 mm long; pistillode
1.52 mm diameter. Female and mixed inflorescences c. 11.5 cm long; bracts and
bracteoles ovate, c. 1 0.5 mm. Female flowers solitary along the rachis; perianth lobes
acute, c. 1 mm across; staminodes 10; styles conical, recurved, c. 1.5 mm long. Cupules
subsessile, ovoid to ellipsoid, 710 4.57 cm, puberulous, spiny; spines 1222 mm
long, much-branched, slender, straight or recurved, pubescent or sometimes glabrous,
sparsely and irregularly arranged with space in between. Nuts 1 in each cupule, ovoid
to ellipsoid, 46 35 cm, free apical part glabrous; wall woody and thick, most part
adnate to the cupule.
Vernacular names. Sabahpedayang (Dusun). Sarawakberangan bo (Iban), wau
(Bidayuh). Bruneibedayang (Dusun), beranang (Malay).
Distribution. Endemic to Borneo. Widely distributed throughout Sabah. In Sarawak,
common in Kapit, Kuching and Sarikei districts. Also in Brunei and Kalimantan.
Ecology. In primary and secondary hill mixed dipterocarp forests, up to 800 m altitude.
19
12. Castanopsis javanica (Blume) A. DC.

(of Java)
Fig. 5.
J. Bot. 1 (1863) 182; Merrill l.c. (1921) 211; Masamune l.c. 233; Soepadmo l.c. (1968) 393, l.c.
(1972) 306; Cockburn l.c. (1972) 205, l.c. (1976) 90; Anderson l.c. 198; Whitmore, Tantra & Sutisna
l.c. 153; PROSEA l.c. 115; Turner l.c. 234. Basionym: Fagus javanica Blume, Flora 7 (1824) 295.
Type: Blume s.n. (= RHL Sheet No. 901309128), Java, Tjibeureum, Mt. Gede (holotype L). Synonyms:
Castanea javanica (Blume) Blume, Bijdr. Fl. Ned. Ind. (1825) 525; Castanea ?montana Blume l.c.
(1825) 526; Castanea javanica Blume var. montana (Blume.) Blume, Fl. Javae Cupul. (1829) 45, t.
24; Castanea javanica (Blume) Blume var. fuscescens (Blume) Blume l.c. (1829) 45; Castanopsis
javanica (Blume) A. DC. var. montana (Blume) A. DC., Prod. 16, 2 (1864) 111; Quercus discocarpa
Hance, J. Bot. 12 (1874) 242; Quercus javanica (Blume) Drake, J. Bot. 4 (1890) 153; Castanopsis
lentiginosa E.F. Warb., Kew Bull. (1936) 20; Castanopsis penangensis A. Camus, Bull. Soc. Bot. Fr.
94 (1947) 4.
Tree up to 40 m tall, 100 cm diameter, non-coppicing. Bark smooth or scaly, brown;

inner bark laminated, brown. Sapwood greyish white. Twigs glabrous, densely covered
with large lenticels. Stipules linear, triangular, c. 5 2 mm. Leaves coriaceous, glabrous
above, sparsely tomentose below; ovate to elliptic, (7)1113(15) (2.5)3.55 cm,
base attenuate, rounded, or acute, margin entire and slightly revolute, apex acuminate,
acumen 1015 mm long; midrib flat or impressed above, raised below; lateral veins 8
12 pairs, impressed above, slightly raised below, disappearing near the leaf margin;
intercostal venation dense, scalariform, obscure above, thin and weakly prominent below;
petiole (4)68(12) mm long, sparsely pilose. Inflorescences male, female or mixed.
Male Inflorescences 4.57 cm long; bracts broadly ovate, 11.5 mm across; bracteoles
ovate, c. 1 0.5 mm. Male flowers in clusters of 23 along the rachis; perianth lobes
acute, c. 1.5 1 mm; stamens 12; filaments c. 1 mm long; pistillode 11.5 mm diameter.
Female and mixed inflorescences 610 cm long; bracts ovate, 22.5 11.5 mm; bracteoles
broadly ovate, c. 0.5 mm across. Female flowers solitary along the rachis; perianth lobes
acute, c. 1 mm across; staminodes 8; styles cylindrical, straight, 23 mm long. Cupules
discoid, asymmetrical, 0.71.5 11.5 cm, densely pubescent, spiny; stalk slender and
straight, 0.51 cm long; spines 23 mm long, branched at the base, flat to triangular
or sometimes slender, straight, pubescent except the tips, sparsely arranged in regular
lines with spaces in between. Nuts 1 in each cupule, discoid or depressed ovoid, 1
1.5 22.5 cm, free part hairy; wall thin, most part free from the cupule.
Vernacular name. Sarawakkupat (Kelabit).
Distribution. Sumatra, Peninsular Malaysia, W Java, and Borneo. In Sabah, known from
Keningau, Penampang, Tenom and Ranau districts. In Sarawak, common in Bario, Kapit,
and Marudi districts. So far not known in Brunei and Kalimantan.
Ecology. In primary hill mixed dipterocarp forest, up to 1000 m altitude.
Uses. The timber is widely used by local people for house-building (e.g house post).
The bark has tanning properties, and the nuts are edible.
13. Castanopsis lucida (Nees in Wall.) Soepadmo

(Latin, lucidus = shining; the upper leaf surface)
20
vi
Reinwardtia 7 (1968) 394, l.c. (1972) 318; Cockburn l.c. (1972) 205; Whitmore, Tantra & Sutisna
l.c. 153; PROSEA l.c. 115; Turner l.c. 243; Argent et al. (eds.) l.c. 240. Basionym: Alseodaphne
lucida Nees in Wall., Pl. As. Rar. 2 (1831) 72. Type: Wallich 2590 (under the name of Laurus
lucida), Singapore (BM, K). Synonym: C. hullettii King ex Hook. f., Fl. Br. Ind. 5 (1888) 623.
Tree up to 20 m tall, 40 cm diameter, non-coppicing; buttresses up to 1.6 m tall, straight.

Bark glabrescent, large-lenticellate or fissured or rarely smooth, brown; inner bark fibrous,
yellowish brown. Sapwood dirty white. Twigs sparsely stellate-hairy, densely covered
with large lenticels. Stipules triangular, c. 2 1 mm. Leaves thick-coriaceous, glabrous
and shining above, sparsely pubescent below; elliptic, oblong to ovate-elliptic, (10)16
21(26) (3.5)810 cm, base rounded, sometimes attenuate or acute, margin entire and
revolute, apex obtuse; midrib broad and raised on both surfaces, stronger below; lateral
veins 1318 pairs, parallel, flat above, strongly raised below, disappearing near the leaf
margin; intercostal venation dense, scalariform, thin and weakly prominent on both surfaces;
petiole 2035 mm long, densely stellate-hairy. Inflorescences male and female. Male
inflorescences 1015 cm long; bracts and bracteoles ovate, 0.51 0.51 mm. Male
flowers in clusters of 35 along the rachis; perianth lobes obtuse, 11.5 0.51 mm;
stamens 12, filaments 2.53 mm long; pistillode 11.5 mm diameter. Female inflorescences
810 cm long; bracts broadly ovate, c. 1.5 mm across; bracteoles ovate, c. 1 mm across.
Female flowers in clusters of 3 along the rachis; perianth lobes acute, c. 0.5 mm across;
staminodes 12; styles conical, straight, 0.51 mm long. Cupules depressed subglobose,
asymmetrical, 2.54 cm diameter, sparsely pubescent, shortly spiny or tubercled, dehiscing
into 34 equal segments; stalk thick and twisted, c. 0.5 cm long; spines or tubercles
23 mm long, unbranched, flat to triangular, recurved, sturdy, sparsely tomentose, sparsely
and regularly arranged in lines. Nuts 24 in each cupule, ovoid, 1.52 cm across, free
part glabrous; wall thick and woody, most part free from the cupule.
Distribution. Peninsular Malaysia, Singapore and Borneo. So far, not known from Sabah.
In Sarawak uncommon, known only by one collection (Beccari PB 3242) from Marop.
Also in Kalimantan.
Ecology. In hill mixed dipterocarp forest, up to 500 m altitude.
Uses. The bark yields tannin and the nuts are edible.
14. Castanopsis megacarpa Gamble

(Greek, mega = large, karpos = fruit; with large fruit)
Kew Bull. (1914) 180; Burkill l.c. 494; Soepadmo l.c. (1968) 395, l.c. (1972) 305; Cockburn l.c.
(1972) 206, l.c. (1976) 90; Anderson l.c. 198; Whitmore, Tantra & Sutisna l.c. 153; PROSEA l.c. 115;
Turner l.c. 234; Argent et al. (eds.) l.c. 240. Type: Kings Collectors 7070, Peninsular Malaysia,
Perak, Kinta (holotype K; isotype L).
Tree up to 36 m tall, 90 cm diameter, non-coppicing. Bark smooth or shallowly fissured,

grey; inner bark fibrous, yellow. Sapwood white or pale yellow. Twigs sparsely tomentose
or sometimes glabrous, sparsely lenticellate or sometimes smooth. Stipules ovate, c. 5
2 mm. Leaves thick-coriaceous, glabrous and shining above, sparsely brownish tomentose
22
below; elliptic-oblong, (15)2024(30) 7.58.5(10) cm, base rounded, attenuate or

acute, sometimes asymmetrical, margin entire and strongly revolute, apex acute; midrib
impressed above, broad and strongly raised below; lateral veins 1315 pairs, flat above,
raised below, disappearing near the leaf margin; intercostal venation scalariform, prominent
on both surfaces; petiole 1225 mm long, sparsely stellate-hairy. Inflorescences male
and female. Male inflorescences (6)814(17) cm long; bracts broadly ovate, c. 1 mm
across; bracteoles ovate, c. 1 0.5 mm. Male flowers in clusters of 3 along the rachis;
perianth lobes obtuse, c. 1.5 1 mm; stamens 12, filaments 22.5 mm long; pistillode
c. 2 mm diameter. Female inflorescences 912 cm long; bracts and bracteoles ovate,
22.5 11.5 mm. Female flowers solitary along the rachis; perianth lobes acute, c.
1.5 1 mm; staminodes 12; styles conical, straight, c. 0.5 mm long. Cupules sessile,
ellipsoid or globose, 58 35.5 cm, pubescent, spiny; spines 1520 mm long, branched,
sturdy, recurved or straight, sparsely or densely and irregularly arranged. Nuts 1 in each
cupule, ellipsoid or globose, 37.5 34.5, free apical part glabrous; wall thick and
woody, most part adnate to the cupule.
Distribution. Peninsular Malaysia, Singapore and Borneo. In Sabah, known from Sipitang,
Beaufort and Ranau districts. In Sarawak, known from Bintulu, Kuching and Lundu (G.
Pueh) districts. Also in Kalimantan.
Ecology. In primary and secondary mixed dipterocarp, kerangas to lower montane forests,
up to 1400 m altitude.
Uses. The timber is used locally for making bowls and other domestic utensils.
15. Castanopsis microphylla Soepadmo

(Greek, mikro = small, phullon = leaf; small-leaved)
Reinwardtia 7 (1968) 395, l.c. (1972) 310; Anderson l.c. 198; Whitmore, Tantra & Sutisna l.c. 153;
PROSEA l.c. 116. Type: Sibat S. 22511, Borneo, Sarawak, Tatau district, Anap, Bt. Mersing (holotype
K; isotypes A, BO, KEP, L, SAN, SAR, SING).
Tree up to 30 m tall, 60 cm diameter, non-coppicing. Bark smooth or sometimes flaky;

inner bark fibrous, yellow. Sapwood yellowish. Twigs sparsely pubescent, smooth. Stipules
linear, triangular, c. 3 1 mm. Leaves coriaceous, glabrous above, sparsely tomentose
below; linear to lanceolate, 58 1.52.5 cm, base rounded, acute or attenuate, margin
entire and plane, apex long-acuminate, acumen c. 15 mm long; midrib raised on both
surfaces; lateral veins 89 pairs, flat above, raised below, disappearing near the leaf margin;
intercostal venation scalariform, obscure above, weakly prominent below; petiole c. 5
mm long, sparsely pilose. Inflorescences male, female or mixed. Male inflorescences
(3)612 cm long; bracts and bracteoles ovate, c. 1 0.5 mm. Male flowers in clusters
of 34 along the rachis; perianth lobes acute, c. 1.5 mm across; stamens 12, filaments
11.5 mm long; pistillode c. 0.5 mm diameter. Female and mixed inflorescences
(4.5)8.511 cm long; bracts ovate, c. 1 0.5 mm; bracteoles subrotundate, c. 0.5
mm across. Female flowers in clusters of 23 along the rachis; perianth lobes acute,
c. 2.5 1 mm; staminodes 12; styles conical, recurved, c. 1 mm long. Cupules (sub)globose,
asymmetrical, 23 1.53 cm, densely tomentose, shortly spiny or tubercled; stalk grooved,
23
0.52 cm long; spines 23 mm long, unbranched, flat to triangular, strongly recurved,

sparsely pubescent at the base, arranged regularly or sometimes irregularly in lines with
space in between. Nuts 23 in each cupule, equally developed, ovoid, 11.5 cm across,
free part sparsely pubescent; wall thin, most part free from the cupule.
Vernacular name. Sabahkogopon (Dusun, Kinabatangan).
Distribution. Endemic to Borneo. In Sabah known from Ranau, Beaufort and Kinabatangan
districts. In Sarawak known from Kapit, Kuching, Mukah, and Tatau districts. Also in
Kalimantan.
Ecology. In lowland mixed dipterocarp forest to lower montane forest, up to 1600 m altitude.
16. Castanopsis motleyana King
Fig. 6.
(James D. Motley, ?1859, engineer and plant collector, mainly in Borneo)

Ann. Bot. Gard. Calc. 2 (1889) 96; Merrill l.c. (1921) 211; Masamune l.c. 233; Soepadmo l.c. (1968)
396, l.c. (1972) 314; Cockburn l.c. (1976) 93; Anderson l.c. 198; Whitmore, Tantra & Sutisna l.c.
153; Kessler & Sidiyasa l.c. 138; PROSEA l.c. 116; Coode et al. (eds.) l.c. 124; Argent et al. (eds.)
l.c. 240. Type: Beccari PB 2613, Borneo, Sarawak (holotype FI; isotypes K, P). Synonym: C. pearsonii
Merr., Philip J. Sc. 30 (1920) 79.
Tree up to 40 m tall, 70 cm diameter, non-coppicing; buttresses spreading, up to 4 m

tall. Bark vertically cracked or scaly, reddish or chocolate-brown; inner bark fibrous,
brown. Sapwood white. Twigs densely rusty brown pubescent when young, glabrescent,
smooth. Stipules ovate, 710 2.53 mm. Leaves thick-coriaceous, glabrous above, densely
pubescent below, glabrescent; obovate, rarely elliptic-oblong, (9)1525(27) (5)7
11 cm, base acute to attenuate, margin entire and strongly revolute, apex short-acuminate,
acumen c. 5 mm long; midrib prominent above, broad and strongly raised below; lateral
veins 1013 pairs, flat above, strongly raised below, joining near the leaf margin; intercostal
venation scalariform, obscure above, slightly to strongly prominent below; petiole 820
mm long, glabrous or sparsely hairy. Inflorescences male, female or mixed. Male
inflorescences (4)7.513 cm long; bracts and bracteoles ovate, c. 1 0.5 mm. Male
flowers in dense clusters of 3 along the rachis; perianth lobes acute, 11.5 mm across;
stamens 12, filaments 23 mm long; pistillode 1.52.5 mm diameter. Female and mixed
inflorescences 1017 cm long; bracts and bracteoles ovate, 11.5 0.5 mm. Female
flowers in clusters of 23 along the rachis; perianth lobes obtuse, c. 5 mm across; staminodes
12; styles cylindrical, straight, 11.5 mm long. Cupules sessile, subglobose, 23 1.5
2.5 cm, closely arranged together along the rachis; densely pubescent, spiny; spines 7
15 mm long, branched at least at the base, slender, straight, densely pubescent except
the tips, densely and irregularly arranged. Nuts 23 in each cupule, ovoid to conical,
1.52.5 1.52 cm, sparsely pilose at the apex; wall thin, most part free from the cupule.
Vernacular names. Sabahbabayi (Rungus). Sarawakberangan daun besar (Malay).
Distribution. Borneo and the Philippines. In Sabah and Sarawak common and widely
distributed. Also in Brunei and Kalimantan.
24
vi
17. Castanopsis oligoneura Soepadmo

(Greek, oligo = few, neuron = nerves or veins; the few-veined leaves)
Tantra & Sutisna l.c. 154; Coode et al. (eds.) l.c. 124. Type: Meijer SAN 32836, Borneo, Sabah, Sandakan
district, Sibuga Road, Kebun China FR (holotype L; isotypes K, SAN, SAR, SING).
Tree up to 25 m tall, 45 cm diameter, non-coppicing. Bark smooth or sometimes lenticellate,

brownish; inner bark fibrous, light brown. Sapwood white. Twigs glabrous, smooth or
sometimes sparsely minute-lenticellate. Stipules ovate, 34 1 mm. Leaves thickcoriaceous, glabrous above, sparsely brownish tomentose below; ovate or elliptic,
(7.5)1315 35(6.5), base acute to attenuate, margin entire and slightly revolute,
apex acute or short-acuminate, acumen c. 5 mm long; midrib thin, flat above, raised
below; lateral veins 57 pairs, flat above, raised below, joining near the leaf margin;
intercostal venation scalariform, weakly prominent on both surfaces; petiole 1222 mm
long, glabrous. Inflorescences male, female or mixed. Male inflorescences (5)710 cm
long; bracts ovate, 11.5 0.7 mm; bracteoles ovate, c. 0.7 0.4 mm. Male flowers
in clusters of 3 along the rachis; perianth lobes ovate, acute, c. 1.5 1 mm; stamens
12, filaments 34 mm long; pistillode c. 1 mm diameter. Female and mixed inflorescences
1013 cm long; bracts and bracteoles ovate, c. 2 1 mm. Female flowers solitary along
the rachis; perianth lobes acute, c. 1.5 mm across; staminodes 12; styles cylindrical, recurved,
12 mm long. Cupules subsessile, subglobose, 23 22.5 cm, sparsely pubescent, spiny;
spines 69 mm long, unbranched, slender, straight or sometimes recurved, pubescent, sparsely
and irregularly arranged. Nuts 1 in each cupule, subglobose, 22.5 2 cm, free apical
part sparsely to densely hairy; wall woody and thick, most part adnate to the cupule.
Distribution. Endemic to Borneo. In Sabah, known from a few collections from Sandakan,
Beaufort and Sipitang districts. In Sarawak, known only from Ulu Baleh in Kapit district
and Bungoh Range in Bau district. Also in Brunei.
Ecology. In lowland mixed dipterocarp forest, up to 300 m altitude.
18. Castanopsis oviformis Soepadmo

(Latin, ovum = egg; formis = shaped; the egg-shaped cupule)
Tantra & Sutisna l.c. 154; Kessler & Sidiyasa l.c. 138; PROSEA l.c. 116; Argent et al. (eds.) l.c. 242.
Type: Sibat S. 23233, Borneo, Sarawak, Mukah district, Balingian, Ulu Arip, Bt. Iju (holotype L;
isotypes A, BO, K, SAN, SAR, SING).
Tree up to 30 m tall, 50 cm diameter, non-coppicing; buttresses small, sometimes branched.

Bark cracked or scaly, brownish; inner bark granular, pale brown. Sapwood yellowish.
Twigs pubescent when young, late-glabrescent, minutely lenticellate. Stipules caducous,
ovate, 35 23 mm. Leaves coriaceous, glabrous on both surfaces; elliptic, oblong
26
or ovate, 714 46.5 cm, base rounded to acute, sometimes asymmetrical, margin entire
and revolute, apex short-acuminate, acumen c. 5 mm long; midrib flat above, raised below;
lateral veins 913 pairs, flat above, slightly raised below, joining near the leaf margin;
intercostal venation dense, scalariform, prominent on both surfaces; petiole 815 mm
long, sparsely hairy. Inflorescences male, female or mixed. Male inflorescences (5)7
11 cm long; bracts and bracteoles ovate, 1.52 11.5 mm. Male flowers in clusters
of 3 along the rachis; perianth lobes obtuse, c. 1 mm across; stamens 12, filaments 1.5
2 mm long; pistillode c. 1.5 mm diameter. Female and mixed inflorescences 4.58 cm
long; bracts and bracteoles broadly ovate, c. 1 mm across. Female flowers solitary along
the rachis; perianth lobes acute, c. 1 mm across; staminodes 12; styles cylindrical, straight,
c. 2 mm long. Cupules ovoid, obovoid or ellipsoid, 25 1.53 cm, sparsely pubescent,
spiny except the abaxial and adaxial sutures; stalk 0.51 cm long; spines 25 mm long,
unbranched, flat to triangular, straight or recurved, sparsely hairy at the base, sparsely
and regularly arranged. Nuts 1 in each cupule, ovoid, 22.5 11.5 cm, free apical
part glabrous; wall woody, thick, most part adnate to the cupule.
Vernacular names. Sarawakberangan pipit (Iban), butoh terampayoh, terampayoh
(Kelabit).
Distribution. Endemic to Borneo. Common and widely distributed throughout Sabah and
Ecology. In primary hill mixed dipterocarp and kerangas forests, up to 900 m altitude.
Use. The nuts are edible though somewhat bitter.
19. Castanopsis paucispina Soepadmo

(Latin, pauci = few or sparse, spina = spines; the sparsely spined cupule)
Kessler & Sidiyasa l.c 139; PROSEA l.c. 116. Type: Wood & Wyatt-Smith SAN A 4385, Borneo,
Sabah, 1/2 mile east of Tambunan, Keningau district (holotype L; isotypes A, BRI, KEP, SAN, SING).
Tree up to 30 m tall, 60 cm diameter, non-coppicing. Bark smooth or slightly cracked,

brown; inner bark fibrous. Twigs densely brownish stellate-tomentose, sparsely minutelenticellate. Stipules ovate, c. 2 1 mm. Leaves thin-coriaceous, glabrous above, sparsely
to densely pubescent below; ovate-lanceolate or elliptic, 814 3.55 cm, base rounded
to acute, margin entire and plane, apex long-acuminate, rarely acute, acumen c. 10 mm
long; midrib impressed above, raised below; lateral veins 810 pairs, flat above, raised
below, joining near the leaf margin; intercostal venation scalariform, weakly prominent
on both surfaces; petiole 1017 mm long, strongly kneed at the base, sparsely stellatehairy. Inflorescences male and female. Male inflorescences 46.5 cm long; bracts and
bracteoles ovate, c. 1 0.5 mm. Male flowers in clusters of 23 along the rachis; perianth
lobes acute, c. 2 1 mm; stamens 1012, filaments 2.53.5 mm long; pistillode c. 1.5
mm diameter. Female inflorescences 510 cm long; bracts and bracteoles ovate, c. 1
0.51 mm. Female flowers solitary along the rachis; perianth lobes ovate, acute, c.
1 0.51 mm; staminodes 12; styles conical, straight, 11.5 mm long. Cupules obovoid
to ellipsoid, 44.5 33.5 cm, glabrous or rarely sparsely pubescent, spiny; stalk 0.5
27
1 cm long; spines 57 mm long, unbranched, sturdy, flat to triangular, glabrous, sparsely

but regularly arranged in 34 rows except on the adaxial suture. Nuts 1 in each cupule,
obovoid to subglobose, c. 4 3 cm, free apical part glabrous; wall woody, thick, most
part adnate to the cupule.
Vernacular name. Sarawaksaled (Kelabit).
Distribution. Endemic to Borneo (except Brunei and Kalimantan). In Sabah recorded
for Trus Madi FR in Tambunan district and in Penampang district. In Sarawak, known
only from Marudi and Tatau districts.
20. Castanopsis pedunculata Soepadmo

(Latin, pedunculatus = provided with a peduncle; the long-stalked cupule)
154. Type: Wood SAN 16805, Borneo, Sabah, Sipitang district, Mile 6, southeast of Malaman, Ulu
Mendalong (holotype L; isotypes K, KEP, SAN, SING).
Tree up to 25 m tall, 40 cm diameter, non-coppicing; buttresses up to 2 m tall. Bark

flaky or lenticellate, brown. Twigs glabrous, densely to sparsely lenticellate. Leaves
coriaceous, glabrous on both surfaces; elliptic to oblong, 1214 35 cm, base rounded
or acute, margin entire and plane, apex short-acuminate, acumen c. 5 mm long; midrib
impressed above, slightly raised below; lateral veins 910 pairs, flat above, raised below,
joining near the leaf margin; intercostal venation dense, subscalariform, prominent on
both surfaces, stronger below; petiole 1520 mm long, glabrous. Inflorescences and flowers
unknown. Cupules obovoid or subglobose, 23.5 1.52.5 cm, sometimes asymmetrical,
densely tomentose, almost smooth; stalk 11.5 cm long. Nuts 3 in each cupule, ovoid,
1.52 cm across, free part sparsely puberulous; wall thin, most part free from the cupule.
Vernacular name. Sarawakberangan (Iban, Malay).
Distribution. Endemic to Borneo (except Brunei and Kalimantan). In Sabah, known by
the type specimen from Ulu Mendalong in Sipitang district (SAN 16805), whereas in
Sarawak known only by a single collection from Similajau FR in Bintulu district (Burley
& Lee 235).
Ecology. In lowland mixed dipterocarp forest, up to 300 m altitude.
21. Castanopsis psilophylla Soepadmo
Fig. 7.
(Greek, psilos = smooth, phullon = leaves; the smooth leaf surface)

Tantra & Sutisna l.c. 154; PROSEA l.c. 116; Argent et al. (eds.) l.c. 242. Type: Muin SAN 25953,
Borneo, Sabah, Lahad Datu district, Kalumpang, Sg. Limau (holotype L; isotypes K, SAN, SAR).
28
vi
Tree up to 30 m tall, 50 cm diameter, non-coppicing. Bark smooth or finely fissured,

brown; inner bark brownish. Sapwood yellowish or whitish. Twigs glabrous, smooth.
Stipules linear, 34 mm long. Leaves thin-coriaceous, glabrous on both surfaces; ovate,
lanceolate or elliptic, 815 2.54.5 cm, base rounded, acute or cuneate, margin entire
and plane, apex acuminate, acumen 520 mm long; midrib flat or sometimes impressed
above, raised below, glabrous; lateral veins 610 pairs, flat above, weakly raised below,
disappearing near the leaf margin; intercostal venation dense, subscalariform, prominent
on both surfaces; petiole 813 mm long, glabrous. Inflorescences male, female or mixed.
Male inflorescences (4)68(10) cm long; bracts and bracteoles ovate, c. 1 0.5 mm.
Male flowers in clusters of 3 along the rachis; perianth lobes obtuse, c. 1 mm across;
stamens 9, filaments c. 2 mm long; pistillode c. 1 mm diameter. Female and mixed
inflorescences 68 cm long; bracts and bracteoles broadly ovate, 12 mm across. Female
flowers in clusters of 34 along the rachis; perianth lobes acute, c. 1 mm across; staminodes
12; styles conical, recurved, c. 1 mm long. Cupules pear-shaped, asymmetrical, 1.53
1.52 cm, dehiscing irregularly into 4 segments, densely golden tomentose all over,
covered with several ridges of short tubercles; subsessile or with stalk 0.51 cm long.
Nuts 35 in each cupule, but only one well-developed, asymmetrical, ovoid, 1.52 cm
across, free part pubescent; wall thin, most part free from the cupule.
Vernacular names. Sabahmengarung (Dusun), tukalod, kogopon (Dusun, Kinabatangan).
Sarawakberangan entadu (Iban).
Distribution. Borneo, the Philippines and Banguey Island. Common and widely distributed
in Sabah. In Sarawak, known only from Limbang and Marudi districts. Also in Kalimantan.
Use. The nuts are edible.
2. LITHOCARPUS Blume
(Greek, lithos = stone, karpos = fruit; the hard and woody cupule and acorn)
empenit, mempening (Malay); empili (Iban); salad and variants (Kayan, Kelabit, Kenyah,
Melanau, Murut, Punan); tekulud (Dusun)
Bijdr. Fl. Ned. Ind. (1825) 526; Rehder, J. Arn. Arb. 1 (1919) 122, ibid. 10 (1929) 132; Warburg, Kew
Bull. 20 (1936) 19; Barnett, Trans. & Proc. Bot. Soc. Edinb. 33 (1942) 332; Soepadmo, Reinwardtia
8 (1970) 197, FM 1, 7 (1972) 318; Cockburn, TFM 1 (1972) 209, TS 1 (1976) 95; Anderson, CLTS
(1980) 199; Corner, WSTM 3rd. edition 1 (1988) 331; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990)
154; Kubitzki in Kubitzki et al. (eds.), Fam. & Gen. Vasc. Pl. 2 (1993) 307; Turner, Gard. Bull. Sing.
47, 1 (1995) 235; Kessler & Sidiyasa, Trees of the Balikpapan-Samarinda Area, East Kalimantan
(1994) 139; PROSEA 5, 2 (1995) 284; Coode et al. (eds.), CLBD (1996) 124; Kochummen, TFPF
(1997) 246; Argent et al. (eds.), MNDT-CK 1 (1997) 242. Synonyms: Synaedrys Lindl., Introd. Nat.
Syst. ed. 3 (1836) 441; Cyclobalanus (Endl.) Oerst., Vid. Medd. Naturch. For. Kjbn. 8 (1867) 80;
Pasania (Miq.) Oerst., ibid. 81.
Trees, rarely shrubs; buttresses or stilt-roots occasionally present. Wood rays narrowuniseriate and broad-multiseriate. Bark smooth or fissured, often lenticellate; inner bark
30
granular or fibrous, with ridges protruding into the sapwood. Twigs glabrous or hairy,
lenticellate or smooth. Stipules extrapetiolar, small, caducous. Terminal bud solitary.
Leaves spirally arranged, margin entire; petiole stout, thickened throughout its length.
Inflorescences erect, male, female, androgynous or mixed, densely simple- and stellatepubescent. Male inflorescences in lateral or subterminal, lax to dense, paniculate clusters
on the new shoots or solitary in the axils of lower (proximal) leaves; bracts and bracteoles
small, caducous. Male flowers solitary or in dichasial clusters of 37(24) along the
rachis or in an androgynous inflorescence along the upper parts of the rachis; perianth
6lobed; stamens (10)12, filaments filiform, glabrous; anthers dorsifixed, reniform;
pistillode well developed, (sub)globose, densely woolly pubescent. Female, androgynous
or mixed inflorescences solitary in the axils of upper (distal) leaves or on the upper
parts of paniculate clusters or terminal on the new shoots; bracts and bracteoles usually
bigger and thicker than those of male inflorescences, persistent. Female flowers solitary
or in dichasial clusters of 210 along the rachis, or in an androgynous inflorescence
along the basal parts of the rachis; perianth 6lobed; staminodes (10)12, rudimentary;
styles 3, conical or cylindrical, straight or recurved, densely hairy at the base, stigmas
minute, terminal and punctiform. Cupules solitary or in dichasial clusters of 210 along
the rachis, partly or completely enclosing a solitary nut, indehiscent, scaly, muricate,
tuberculate, almost smooth or lamellate with the lamellae adpressed to the external cupule
wall. Fruits (acorns) rounded in cross-section, sometimes falsely multi-septate by intrusions
of the endocarp; wall woody or bony, most part adnate or free from the cupule. Seed
germination hypogeal.
Distribution. About 300 species, mainly in the tropics and subtropics of NE India (Nepal,
Bhutan, Assam), Burma, China, Korea, Japan, Formosa, Hainan, Indo-China, Thailand,
and Malesia (except for the eastern parts of Java and the Lesser Sunda Islands). One
species, Lithocarpus densiflorus, occurs in the SW parts of the United States. In Sabah
and Sarawak, the genus is represented by 61 species.
Ecology. In lowland to montane forests, from sea level up to 3000 m altitude, more
commonly below 1500 m, on different types of soils, including limestone substrate.
Uses. The timber is locally used for medium to heavy constructions but preservative
treatment is a prerequisite for permanent structures. The wood is suitable for manifacturing
pulp and paper, and the bark contains tannin. Round wood and branches of several species
are used as bed logs in mushroom cultivation. The nuts of several species are edible.
Key to Lithocarpus species

1. Leaves sessile.........58. L. tawaiensis
Leaves distinctly petioled.........2
2. Most part of acorn wall adnate to the cupule......3
Most part of acorn wall free from the cupule....13
3. Cupule scaly, muricate, echinate, echinulate or tuberculate...4
Cupule lamellate.....6
31
4. Leaves thin-coriaceous, base rounded to acute. Cupule obovoid or pear-shaped, stalk

520 mm long.....52. L. rotundatus
Leaves thick-coriaceoaus or coriaceous, base acute to cuneate. Cupule obconical, sessile
or with short stalk up to 5 mm long..................5
5. Intercostal venation scalariform, thin but clearly visible below; lateral veins dense.
Cupule tuberculate, with an opening 1.54 cm diameter at the apex...........47. L. pulcher
Intercostal venation subscalariform, obscure on both surfaces or slightly prominent
below; lateral veins lax. Cupule armed with recurved prickles or spine-like appendages,
with an opening c. 1 cm diameter at the apex................18. L. echinifer
6. Leaves glabrous or sparsely tomentose above, glabrous or sparsely to densely appressed
tomentose below.....7
Leaves densely appressed tomentose on both surfaces or glabrous above and densely
appressed tomentose below...........9
7. Leaves 1524 710 cm; petiole 1530 mm long. Cupule obovoid to subglobose
............25. L. hallieri
Leaves 616 36 cm; petiole 515 mm long. Cupule obconical, ovoid or ellipsoid8
8. Cupule obconical. Intercostal venation dense; midrib raised on both surfaces, stronger
above........15. L. corneri
Cupule ovoid to ellipsoid. Intercostal venation lax; midrib slightly raised above, strongly
raised below...............................53. L. ruminatus
9. Cupule much longer than broad, turbinate, ellipsoid or obovoid.....10
Cupule almost as long as broad, subglobose or depressed globose.....12
10. Intercostal venation dense. Cupule obovoid, strongly lamellate, lamellae raised and
folded inwards, longitidunally grooved.........31. L. keningauensis
Intercostal venation usually lax. Cupule turbinate to ellipsoid, smooth, lamellae not
strongly raised, edge minutely denticulate or entire.....11
11. Apex of cupule with an opening 0.51 cm diameter. Leaves thick-coriaceous, rigid;
lateral veins clearly joining near the leaf margin....59. L. turbinatus
Cupule without an opening at the apex. Leaves coriaceous, lateral veins faintly joining
or disappearing near the leaf margin.......3. L. beccarianus
12. Leaves thick-coriaceous, sometimes rigid, 1115.5 cm long, apex acuminate, acumen
5 12 mm long; lateral veins dense. Cupule smooth........30. L. kalkmanii
Leaves coriaceous, not rigid, 1625 cm long, apex obtuse to short-acuminate, acumen
up to 5 mm long; lateral veins lax. Cupule ridged..45. L. porcatus
13. Most part of acorn enclosed by the cupule............................14
Most part of acorn exserted.........20
14. Lateral veins clearly joining near the leaf margin...15
Lateral veins disappearing or faintly joining near the leaf margin.16
32
15. Leaves broadly ovate or elliptic-oblong; lateral veins lax, forming an angle of 60
80 with the midrib. Cupule with a stalk c. 5 mm long; appendages hook-like, recurved.
Acorn ovoid to subglobose.......6. L. brochidodromus
Leaves narrowly elliptic, lanceolate or oblanceolate; lateral veins dense, forming an
angle of 4060 with the midrib. Cupule sessile, appendages straight or slightly recurved.
Acorn conical..................................32. L. kochummenii
16. Petiole less than 5 mm long. Cupule covered with flexible spine-like appendages.
Leaves sparsely to densely tomentose with stellate hairs on both surfaces; base usually
asymmetrical.......14. L. coopertus
Petiole 525 mm long. Cupule smooth and often with longitudinal streaks, lamellate
or irregularly tuberculate and ridged. Leaves sparsely to densely appressed tomentose
below; base symmetrical......................17
17. Leaves densely appressed tomentose below; intercoastal venation lax. Cupule irregularly
tuberculate and ridged..........................................12. L. confragosus
Leaves sparsely appressed tomentose below; intercoastal venation dense. Cupule
lamellate or smooth and often with longitudinal streaks.....................18
18. Leaf margin undulate; lateral veins 12 pairs or more, dense. Acorn glabrous except
small part at the apex..............5. L. blumeanus
Leaf margin not undulate, lateral veins 12 pairs or less, lax. Acorn densely simplehairy or velvety.......19
19. Cupule smooth and often with longitudinal streaks. Acorn ovoid, scar deeply concave,
c. 2 cm diameter. Petiole 1525 mm long. Leaf base rounded......37. L. mariae
Cupule lamellate. Acorn depressesed ovoid to subglobose, scar shallowly concave,
11.5 cm diameter. Petiole 515 mm long. Leaf base mostly acute.....21. L. encleisacarpus
20. Leaves densely hairy on both surfaces or at least below.21
Leaves glabrous on both surfaces, or glabrous or subglabrous above and sparsely
tomentose below, or sparsely tomentose on both surfaces, or covered with reddish
brown papillae...........29
21. Intercostal venation scalariform..........22
Intercostal venation subscalariform, irregular or reticulate..24
22. Acorn densely tomentose. Cupule cup-shaped, 2.53.5 cm diameter, enclosing up to
half of the acorn..........60. L. urceolaris
Acorn glabrous except for small part at the apex. Cupule saucer-shaped, 1.42.5 cm
diameter, enclosing less than half of the acorn.....23
23. Leaves broadly lanceolate or obovate; petiole usually less than 5 mm long. Cupule
muricate.....28. L. hystrix
Leaves elliptic to oblong; petiole 512 mm long. Cupule scaly.....57. L. sundaicus
24. Lateral veins clearly joining near the leaf margin.....25
Lateral veins faintly joining or disappearing near the leaf margin........27
33
25. Cupule in clusters of 23 and arranged closely together along the rachis...
.........61. L. wallichianus
Cupule solitary and distantly spaced along the rachis26
26. Acorn densely tomentose. Leaves elliptic or obovate; intercostal venation dense, obscure
above.............................13. L. conocarpus
Acorn glabrous. Leaves oblanceolate, broadly elliptic to oblong; intercostal venation
lax, prominent below and impressed above..............41. L. nieuwenhuisii
27. Cupule sessile. Acorn glabrous except for the apex. Leaves less than 10 cm long and
5 cm wide; apex acute or acuminate with an acumen 510 mm long....27. L. havilandii
Cupule stalked. Acorn densely tomentose or sparsely simple-hairy. Leaves more than
10 cm long and 5 cm wide; apex caudate or acuminate with an acumen 1025 mm
long.................................28
28. Leaves thick-coriaceous, rigid, margin strongly revolute.....17. L. dasystachyus
Leaves thin-coriaceous, not rigid, margin slightly revolute.....9. L. caudatifolius
29. Intercostal venation obscure on both surfaces or thinly prominent below........30
Intercostal venation obscure above and prominent below, or prominent on both surfaces
........43
30. Leaf apex rounded, obtuse or emarginate........31
Leaf apex acute, acuminate, or caudate............33
31. Leaves covered with reddish brown papillae on both surfaces, denser below.
......44. L. papillifer
Leaves glabrous on both surfaces or sparsely to densely tomentose with a few simple
and/or stellate hairs but without reddish brown papillae...32
32. Leaves bullate, ovate, (3)4.57 24 cm. Cupule 0.51 1.51.8 cm, lamellate.
....7. L. bullatus
Leaves not bullate, obovate to elliptic, 1014(21.5) 35(7) cm. Cupule 1.52.5
3.75 cm, scaly..50. L. revolutus
33. Leaves narrower, with a length:width ratio of 44.5:1.....39. L. melataiensis
Leaves broader, with a length:width ratio of 13:134
34. Cupule scaly or armed with prickles or spine-like appendages..35
Cupule lamellate.........36
35. Leaves thin-coriaceous. Cupule armed with short, slender prickles or spine-like
appendages. Acorn sparsely tomentose, scar 2.32.6 cm diameter..19. L. echinulatus
Leaves thick-coriaceous, rigid. Cupule covered with distinct and rather broad scale-like
appendages. Acorn glabrous and shining, scar 1.51.7 cm diameter..40. L. muluensis
36. Leaves thick-coriaceous and rigid...37
Leaves usually thin-coriaceous or rarely coriaceous but not rigid..38
34
37. Leaf apex acute; lateral veins lax........51. L. rigidus

Leaf apex acuminate, acumen 1020 mm long; lateral veins dense......26. L. hatusimae
38. Mature cupule 1.31.7 cm deep........49. L. rassa
Mature cupule 0.31 cm deep..........39
39. Cupule cup-shaped, 0.61 cm deep. Lateral veins disappearing near the leaf
margin.....42. L. nodosus
Cupule saucer-shaped, 0.30.5(0.7) cm deep. Lateral veins faintly joining or rarely
disappearing near the leaf margin..........40
40. Cupule sessile.............41
Cupule stalked........42
41. Leaves narrowly elliptic or ovate, (5.5)68.5(10.5) 24 cm, base obtuse; midrib
flat above, strongly raised below; petiole 59 mm long. Cupule 0.30.5 0.81.2
cm. Acorn ovoid to conical; scar concave, 0.30.7 cm diameter.........48. L. pusillus
Leaves broadly elliptic, (9)1214(17) (4)56(7) cm, base cuneate to acute,
midrib raised on both surfaces; petiole (8)1215 mm long. Cupule 0.50.7 1.5
2 cm. Acorn depressed ovoid to subglobose; scar deeply concave, c. 1 cm diameter.
...2. L. bancanus
42. Leaf base rounded; lateral veins forming an angle of 3040 with the midrib. Petiole
57 mm long.....1. L. andersonii
Leaf base attenuate to acute; lateral veins forming an angle of 4560 with the midrib.
Petiole 1015 mm long.....4. L. bennettii
43. Cupule variously scaly.........44
Cupule lamellate........49
44. Lateral veins disappearing near the leaf margin......33. L. lampadarius
Lateral veins clearly to faintly joining near the leaf margin....45
45. Leaves sparsely tomentose with stellate and simple hairs on both surfaces. Acorn
conical.....34. L. leptogyne
Leaves glabrous on both surfaces. Acorn ovoid, cylindrical, depressed ovoid or
subglobose.......46
46. Leaf base cordate to auriculate.......29. L. jacobsii
Leaf base acute, attenuate, cuneate, obtuse or rounded....47
47. Acorn glabrous. Leaf base acute or cuneate20. L. elegans
Acorn densely or sparsely brownish simple-hairy or velvety. Leaf base rounded, attenuate
or obtuse...........................48
48. Acorn ovoid, 1.52 cm long, densely covered with reddish brown simple, long and stellate
hairs...................23. L. ferrugineus
Acorn cylindrical, 2.53.7 cm long, sparsely greyish simple-hairy especially at the
base and apex....46. L. pseudokunstleri
35
49. Acorn glabrous or hairy on the upper part only...50

Acorn variously hairy all over....56
50. Leaves thick-coriaceous; midrib strongly raised on both surfaces; intercostal venation
lax..........51
Leaves coriaceous to thin-coriaceous; midrib faintly raised on both surfaces; intercostal
venation dense.........52
51. Leaf base rounded to acute; lateral veins disappearing near the leaf margin, lax;
intercostal venation reticulate, obscure above. Cupule 23 45 cm, lamellae strongly
raised. Acorn 34 35 cm, scar deeply concave, 22.5 cm diameter56. L. stonei
Leaf base cuneate; lateral veins faintly to clearly joining near the leaf margin, dense;
intercostal venation scalariform, prominent on both surfaces. Cupule 0.51 1.5
2.5 cm, lamellae slightly distinct. Acorn 12 1.52 cm, scar shallowly concave
or flat, 11.4 cm diameter.........8. L. cantleyanus
52. Leaves oblanceolate; base cuneate, decurrent..53
Leaves elliptic, ovate or oblong; base rounded, obtuse to cordate.54
53. Leaves with a length:width ratio of 2:1, coriaceous, sometimes rigid, glabrous on
both surfaces; intercostal venation distinct on both surfaces. Cupule cup- to saucershaped......35. L. lucidus
Leaves with a length:width ratio of 2.53:1, thin-coriaceous, not rigid, glabrous above,
sparsely appressed tomentose below; intercostal venation obscure above, prominent
below. Cupule saucer-shaped............43. L. oblancifolius
54. Leaf margin often undulate; lateral veins forming an angle of 3045(60) with the
midrib. Cupule lamellae with dentate edge........24. L. gracilis
Leaf margin not undulate; lateral veins forming an angle of 4560 with the midrib.
Cupule lamellae with minutely denticulate or entire edge.......55
55. Lateral veins 711 pairs. Cupule subsessile, saucer-shaped, 1.21.7 cm diameter, edge
of lamellae minutely denticulate.........11. L. confertus
Lateral veins 1216 pairs. Cupule with stalk 510 mm long, cup-shaped, 22.5 cm
diameter, lamellae entire.........22. L. ewyckii
56. Leaves 2646 cm long........54. L. sandakanensis
Leaves 727 cm long......57
57. Intercostal venation subscalariform or reticulate...58
Intercostal venation scalariform.........59
58. Leaves thick-coriaceous; lateral veins disappearing near the leaf margin. Acorn densely
greyish hairy......10. L. clementianus
Leaves thin-coriaceous; lateral veins faintly joining near the leaf margin. Acorn sparsely
hairy.......................16. L. daphnoideus
59. Scar of acorn flat or shallowly concave.......36. L. luteus
Scar of acorn deeply concave..................60
36
60. Cupule sessile. Acorn densely silvery grey tomentose with simple hairs; scar 1.5
2 cm diameter. Leaves (11)1316(20) (4)57(8.5) cm; base cuneate, decurrent;
midrib flat above, raised below...........38. L. meijeri
Cupule with stalk 510 mm long. Acorn densely sericeous; scar 23 cm diameter.
Leaves (9)1213.5 (3)46 cm; base acute; midrib raised on both surfaces...
...............................................55. L. sericobalanus
1. Lithocarpus andersonii Soepadmo

(J.A.R. Anderson, former Forest Officer in Sarawak and Brunei)
Reinwardtia 8 (1970) 215, l.c. (1972) 365; Anderson l.c. 199; Whitmore, Tantra & Sutisna, l.c. 154;
PROSEA 5, 2 (1995) 288; Coode et al. (eds.) l.c. 124; Argent et al. (eds.) l.c. 243. Type: Anderson
5102, Borneo, Sarawak, Mukah (holotype L; isotypes K, SAR).
Tree up to 30 m tall, 60 cm diameter; stilt-roots occasionally present, up to 1.8 m tall.

Bark smooth, brownish; inner bark fibrous, brownish. Sapwood yellowish. Twigs glabrescent,
sparsely to densely lenticellate. Stipules ovate, acute, 34 12 mm. Leaves thincoriaceous to coriaceous, not rigid, glabrous above, sparsely appressed greyish tomentose
below; broadly elliptic, (6)710(12) 24(5) cm; base rounded, rarely acute, margin
slightly revolute, apex acuminate, acumen 1020 mm long; midrib raised on both surfaces,
stronger and glabrescent below; lateral veins thin, (6)911 pairs, lax, flat but clearly
visible on both surfaces, faintly joining near the leaf margin, forming an angle of 30
40 with the midrib; intercostal venation reticulate, lax, obscure on both surfaces; petiole
57 mm long. Inflorescences male or androgynous. Male inflorescences 510 cm long;
bracts and bracteoles ovate, 0.30.6 0.20.5 mm. Male flowers in clusters of 23 along
the rachis; perianth lobes acute, c. 1.2 0.7 mm; stamen filaments 23 mm long; pistillode
subglobose, 11.2 mm diameter. Androgynous inflorescences 710 cm long; bracts and
bracteoles ovate, c. 0.4 0.2 mm. Female flowers solitary along the rachis; perianth
lobes obtuse, 0.50.6 mm across; staminodes 12; styles conical, straight, c. 1 mm long.
Cupules solitary along the rachis, saucer-shaped, 0.50.7 1.51.7(2) cm, densely
appressed tomentose, lamellate; stalk 34 mm long; wall woody, thin, enclosing less than
half of the acorn; lamellae distinct, edge denticulate, set in 56 regular lines. Acorns
ovoid to conical, 1.22.3 1.61.8 cm, most part exserted, glabrous, dark purplish brown;
base flat, apex acute; scar deeply concave, 0.81.3 cm diameter; wall bony, thin, most
part free from the cupule.
Vernacular names. Sarawakempenit batu, empili pipit (Iban), empenit padang, empenit
jangkar (Malay), kayu kikai, penyibong (Melanau Oya), keraki boras (Bidayuh), tekaled
(Melanau).
Distribution. Endemic to Borneo. So far not known in Sabah. In Sarawak, uncommon
but distributed throughout Sibu, Kapit, Sarikei, Sri Aman, and Kuching districts. Also
in Brunei.
Ecology. In primary and secondary peatswamp and kerangas forests, at low altitude.
37
2. Lithocarpus bancanus (Scheff.) Rehder

(of Bangka, Sumatra)
J. Arn. Arb. 10 (1929) 132; Soepadmo l.c. (1970) 218, l.c. (1972) 360; Cockburn l.c. (1972) 213;
Anderson l.c. 199; Whitmore, Tantra & Sutisna l.c. 154; Turner l.c. 235; PROSEA l.c. 288; Coode et
al. (eds.) l.c. 124; Argent et al. (eds.) l.c. 243. Basionym: Quercus bancana Scheff., Nat. Tijd. Ned.
Ind. 31 (1870) 361. Type: Teijsmann 224, Sumatra, Bangka, Batu Balai (holotype BO; isotype U).
Synonyms: Synaedrys bancana (Scheff.) Koidzumi, Bot. Mag. Tokyo 30 (1916) 190; Pasania bancana
(Scheff. ) Markgr., Bot. Jahrb. 59 (1924) 79; Q. rajah Hance, J. Bot. 16 (1878) 198; S. rajah (Hance)
Koidzumi l.c. (1916) 192; P. rajah (Hance) S. Moore, J. Bot. 63 (1925) 115; L. rajah (Hance) A.
Camus, Riviera Scientific 18 (1932) 41; Q. scyphigera Hance var. riedelii King, Ann. Roy. Bot. Gard.
Calc. 2 (1889) 39; L. scyphigera (Hance) A. Camus var. riedelii (King) A. Camus, Chnes 3 (1954)
962.
Tree up to 25 m tall, 50 cm diameter. Bark smooth or scaly, yellowish to greyish brown;

inner bark fibrous, pinkish. Sapwood yellowish. Twigs sparsely appressed tomentose,
late-glabrescent, smooth or sparsely lenticellate. Stipules deltoid, 0.51 0.5 mm. Leaves
thin-coriaceous, rarely coriaceous but not rigid, glabrous above, sparsely yellowish to
brownish appressed tomentose with simple hairs and sometimes mixed with a few stellate
hairs below; broadly elliptic, (9)1214(17) (4)56(7) cm, base cuneate to acute,
margin slightly revolute, apex acuminate, acumen 515 mm long; midrib thin, raised
on both surfaces, stronger and sparsely to densely appressed tomentose below; lateral
veins thin, (9)1012(14) pairs, lax, flat on both surfaces or silghtly raised below, faintly
joining near the leaf margin, forming an angle of 4060 with the midrib; intercostal
venation subscalariform to subreticulate, dense, obscure on both surfaces; petiole
(8)1215 mm long. Inflorescences male, androgynous or mixed. Male inflorescences
520 cm long; bracts and bracteoles ovate, 0.51 0.5 mm. Male flowers solitary
along the rachis; perianth lobes acute, 46 23 mm; stamen filaments 2.53.5 mm
long; pistillode globose, c. 1 mm diameter. Androgynous or mixed inflorescences 10
16 cm long; bracts and bracteoles ovate, 0.51 0.5 mm. Female flowers solitary along
the rachis; perianth lobes acute, 45 mm across; staminodes 12; styles conical, recurved,
11.5 mm long. Cupules solitary along the rachis, sessile, saucer-shaped, 0.50.7 1.5
2 cm, densely appressed tomentose, lamellate; wall woody, thin, enclosing less than half
of the acorn; lamellae distinct, edge denticulate, set in 56 regular lines. Acorns depressed
ovoid to subglobose, 1.52 cm across, most part exserted, glabrous, brownish; base rounded,
apex rounded and abruptly acute, scar deeply concave, c. 1 cm diameter; wall woody,
thin, most part free from the cupule.
Distribution. Sumatra, Peninsular Malaysia and Borneo. Uncommon. In Sabah, only
recorded from Mt. Silam in Lahad Datu district. In Sarawak, only known from Kuching
district. Also in Brunei.
Ecology. In hill mixed dipterocarp forest on clay and sandstone-derived soils, up to 900
m altitude. In Sarawak also recorded in peatswamp forest.
3. Lithocarpus beccarianus (Benth.) A. Camus

Riviera Scientific 18 (1932) 39; Soepadmo l.c. (1970) 220, l.c. (1972) 329; Cockburn l.c. (1976) 100;
38
Whitmore, Tantra & Sutisna l.c. 154; PROSEA l.c. 289; Coode et al. (eds.) l.c. 124; Argent et al.
(eds.) l.c. 247. Basionym: Quercus beccariana Benth. in Hooker f., Icon. Pl. 14 (1880) 10. Type:
Beccari PB 3310, Borneo, Sarawak, Marop, near Batang Lupar (holotype FI; isotypes A, BM, K).
Synonyms: Pasania beccariana (Benth.) Prantl in Engler & Prantl, Nat. Pflz. Fam. 3, (1894) 55;
Synaedrys beccariana (Benth.) Koidzumi l. c. (1916) 188.
Tree up to 30 m tall, 60 cm diameter. Bark scaly, brown; inner bark fibrous, brown
to yellowish brown. Sapwood yellowish brown. Twigs densely tomentose, late-subglabrescent,
smooth or finely fissured. Stipules linear, 1.52 0.50.7 mm, caducous. Leaves
coriaceous, glabrous above, densely tomentose with stellate hairs below; narrowly elliptic
or ovate, (6)812 35(6) cm, base acute to cuneate, margin revolute, apex abruptly
acute to acuminate, acumen 1015 mm long; midrib impressed or flat above, raised below;
lateral veins thin, 68(12) pairs, lax, flat above, raised below, faintly joining or disappearing
near the leaf margin, forming an angle of 3045 with the midrib; intercostal venation
scalariform, lax, slightly prominent on both surfaces or obscure above; petiole 518 mm
long. Inflorescences male or androgynous. Male inflorescences 58 cm long; bracts and
bracteoles ovate, c. 0.5 mm across. Male flowers in clusters of 23 along the rachis;
perianth lobes acute, c. 1 0.5 mm; stamen filaments c. 3 mm long; pistillode subglobose,
c. 2 mm diameter. Androgynous inflorescences 510 cm long; bracts ovate, c. 1.5
0.5 mm; bracteoles ovate, c. 0.5 0.3 mm. Female flowers solitary along the rachis;
perianth lobes acute, c. 0.5 mm across; staminodes 12; styles conical, straight, c. 1 mm
long. Cupules ellipsoid, solitary along the rachis, sessile, 67(12.5) 2.53.5(6.5)
cm, tomentose, lamellate; wall woody, thick, smooth, densely hairy, enclosing the acorn
completely, without apical opening; lamellae distinct but not strongly raised, edge entire
or minutely denticulate, set in 68 lines. Acorns ellipsoid, 56 45 cm, tomentose,
brown; base rounded, apex rounded or somewhat flat; wall woody, thick, most part adnate
to the cupule.
Distribution. Endemic to Borneo. In Sabah recorded for Beaufort, Tambunan and Tawau
districts. In Sarawak known from Mt. Buri in Samarahan district, Mt. Hose in Kapit
district and from the type locality. Also in Brunei and Kalimantan.
Ecology. In primary mixed dipterocarp forest, up to 300 m altitude.
4. Lithocarpus bennettii (Miq.) Rehder

(J.J. Bennett, 18011876, English botanist)
J. Arn. Arb. 1 (1919) 123; Soepadmo l.c. (1970) 220, l.c. (1972) 356; Cockburn l.c. (1972) 215, l.c.
(1976) 102; Anderson l.c 199; Corner l.c. 339; Whitmore, Tantra & Sutisna l.c. 155; Turner l.c. 235;
PROSEA l.c. 289; Coode et al. (eds.) l.c. 124; Argent et al. (eds.) l.c. 247. Basionym: Quercus
bennettii Miq., Fl. Ind. Bat. 1, 1 (1856) 857. Type: Horsfield HB 3402, Sumatra, Bangka, Djebus
(holotype U; isotypes CGE, K). Synonyms: Cyclobalanus bennettii (Miq.) Oerst., Kng. Danske
Vid. Selsk. Skrift. 5, 9 (1871) 375; Pasania bennettii (Miq.) Gamble, J. Roy. As. Soc. Beng. 75, 2
(1915) 433; Synaedrys bennettii (Miq.) Koidzumi l.c. 190.
Tree up to 35 m tall, 80 cm diameter; buttresses up to 3 m high, stilt-root occasionally

present. Bark smooth or fissured, greyish brown; inner bark fibrous, yellowish brown.
Sapwood yellowish. Twigs glabrescent, sparsely lenticellate or smooth. Stipules linear,
39
2.53.5 1 mm. Leaves thin-coriaceous to coriaceous, not rigid, glabrous above, sparsely
yellowish appressed tomentose below; broadly elliptic, (7.5)9.512(15) (3)3.55.5(
7) cm, base attenuate to acute, margin slightly revolute, apex acuminate, acumen 5
15 mm long; midrib strongly raised above, slightly raised or flat and glabrescent below;
lateral veins thin, (8)912 pairs, lax, slightly raised or flat on both surfaces, faintly
joining or disappearing near the leaf margin, forming an angle of 4560 with the midrib;
intercostal venation subscalariform or reticulate, dense, obscure above and thinly prominent
below or obscure on both surfaces; petiole 1012(15) mm long. Inflorescences male
or androgynous. Male inflorescences 720 cm long; bracts and bracteoles ovate, 0.5
1 0.30.6 mm. Male flowers solitary along the rachis; perianth lobes obtuse, c. 1
0.8 mm; stamen filaments c. 2.5 mm long; pistillode subglobose, c. 1.3 mm diameter.
Androgynous inflorescences 1020 cm long; bracts ovate, c. 0.9 0.6 mm; bracteoles
subrounded, 0.20.3 mm across. Female flowers solitary along the rachis; perianth lobes
acute, c. 1 0.5 mm; staminodes 12; styles conical, straight, c. 1 mm long. Cupules
solitary along the rachis, with stalk c. 5 mm long, saucer-shaped, 0.30.5 12 cm,
densely appressed tomentose with simple hairs, lamellate; wall bony, thin, enclosing less
than half of the acorn; lamellae slightly distinct, edge minutely denticulate, set in 5
7 regular lines. Acorns ovoid to conical, 11.7 1.32 cm, most part exserted, glabrous
except for the sparsely simple-hairy apex, dark purplish brown; base flat, apex acute;
scar deeply concave, c. 1 cm diameter; wall bony, thin, most part free from the cupule.
Vernacular name. Sarawakempili pipit (Iban).
Distribution. Sumatra, Peninsular Malaysia, Singapore and Borneo. Uncommon but widely
distributed throughout Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. In primary and secondary mixed dipterocarp and kerangas forests, on a wide
range of soils, at 2001200 m altitude. In Sarawak also known in limestone hills.
5. Lithocarpus blumeanus (Korth.) Rehder

(C.L. Blume, 17961862, German medical doctor; 18221826 Assistant Director of the
Bogor Botanic Garden, 1829 Director of the Rijksherbarium, Leiden)
J. Arn. Arb. 10 (1929) 132; Soepadmo l.c. (1970) 221, l.c. (1972) 339; Cockburn l.c. (1976) 105;
Anderson l.c. 199; Whitmore, Tantra & Sutisna l.c. 155; Coode et al. (eds.) l.c. 125. Basionym:
Quercus blumeana Korth., Kruidk. (1844) 208. t. 44. Type: Korthals s.n. (= RHL Sheet No 901309475),
Borneo, SE Kalimantan, Dusun R. (Barito) (holotype L; isotype A). Synonyms: Cyclobalanus
blumeana (Korth.) Oerst., Vid. Medd. Naturh. For. Kjbn. 8 (1867); Synaedrys blumeanus (Korth.)
Koidzumi l.c. 186; Castanopsis blumeanus (Korth.) Rehder l.c. (1919) 122.
Tree up to 15 m tall, 25 cm diameter. Bark scaly, greyish brown; inner bark fibrous,
brown. Sapwood yellowish. Twigs subglabrous, sparsely lenticellate. Stipules triangular,
c. 2 1 mm. Leaves thin-coriaceous, glabrescent above, sparsely appressed brownish
tomentose below; broadly ovate or broadly elliptic or sometimes elliptic-oblong, (8)12
16(23) (3.5)58.5(11) cm, base rounded, subcordate or broadly acute, symmetrical,
margin undulate, revolute, apex acuminate, acumen 1015 mm long; midrib raised on
both surfaces, stronger and sparsely tomentose below; lateral veins thin, 1215(17) pairs,
dense, flat above, raised below, faintly joining near the leaf margin, forming an angle
40
of 3045 with the midrib; intercostal venation reticulate, dense, obscure above, prominent
below; petiole 1013 mm long. Inflorescences male, female or androgynous. Male
inflorescences 1520 cm long; bracts triangular, c. 0.9 0.6 mm; bracteoles broadly
ovate, acute, c. 0.3 0.2 mm. Male flowers in clusters of 3 along the rachis; perianth
lobes acute, c. 0.7 0.5 mm; stamen filaments c. 3 mm long; pistillode globose, c. 1
mm diameter. Female or androgynous inflorescences 1015 cm long; bracts ovate, c.
0.4 0.2 mm; bracteoles subrounded, 0.20.3 mm across. Female flowers solitary along
the rachis; perianth lobes acute, c. 0.8 0.5 mm; staminodes 12; styles cylindrical, slightly
recurved, c. 1.2 mm long. Cupules solitary along the rachis, with stalk 510 mm long,
11.5 2.53 cm, tomentose, lamellate; wall bony, thin, enclosing the acorn almost
completely or for its most part; lamellae slightly distinct, edge denticulate, set in 5
6 regular lines. Acorns depressed ovoid, 1.52 2.53 cm, glabrous except for a small
part at the apex which is densely simple and stellate hairy; base rounded, apex acute;
scar deeply concave or flat, c. 1.4 cm diameter; wall bony, thin, most part free from
the cupule.
Distribution. Peninsular Malaysia and Borneo. In Sabah, recorded in all districts. In
Sarawak, known in Kuching, Sri Aman, Kapit, Miri, and Limbang districts. Also in Brunei
and Kalimantan.
Ecology. In primary mixed dipterocarp forest to lower montane forest, up to 1600 m
altitude, mainly on clay soils of igneous origin.
6. Lithocarpus brochidodromus S. Julia & Soepadmo
Fig. 8.
(Latin, brochidodromus = loop-veined; the leaves)

Gard. Bull. Sing. 50 (1998) 125. Type: Dewol & Lideh SAN 105591, Borneo, Sabah, Pensiangan,
Sapulut, Sg. Saburan (holotype SAN).
Tree up to 30 m tall, 55 cm in diameter. Bark flaky, greyish or brownish; inner bark

reddish or greyish or greenish. Sapwood brownish or whitish. Twigs densely tomentose
with appressed stellate and simple hairs, subglabrescent, smooth or sparsely large-lenticellate,
sometimes scaly. Stipules linear or narrowly ovate, 515 25 mm. Leaves coriaceous,
rigid, sparsely appressed yellowish tomentose with simple and stellate hairs or rarely
glabrous above, densely appressed yellowish tomentose below; elliptic-oblong or broadly
ovate, 1422(32) 59(11) cm, base rounded or acute, margin strongly revolute, apex
acuminate, acumen 1520 mm long; midrib broad, slightly raised above, strongly raised
below; lateral veins thick, 712 pairs, lax, strongly impressed above, strongly raised below,
clearly and strongly joining near the leaf margin, forming an angle of 6080 with the
midrib; intercostal venation scalariform, lax, prominent and impressed above, strongly
prominent below; petiole 412 mm long. Inflorescences male and female. Male inflorescences 610 cm long; bracts linear to narrowly ovate, 1.53.5 12 mm; bracteoles
linear, c. 1.7 0.6 mm. Male flowers solitary along the rachis; perianth lobes acute,
2.53 11.5 mm; stamen filaments c. 2 mm long; pistillode subglobose, 1.52 mm
in diameter. Female inflorescences 1418 cm long; bracts ovate, 22.5 11.7 mm;
bracteoles linear to narrowly elliptic, c. 2 0.5 mm. Female flowers solitary along the
rachis; perianth lobes obtuse, 0.40.7 mm across; staminodes 12; styles conical, slightly
41
viTREE
recurved, 1.52.5 mm long. Cupules solitary along the rachis, with stalk c. 5 mm long,
1.52.5 22.5 cm, densely appressed stellate-tomentose; set with recurved hook-like
appendages; wall bony, thick, enclosing the acorn completely or more than half of the
acorn; hook-like appendages distinct, sturdy, set irregularly. Acorns ovoid to subglobose,
1.52 cm across, sparsely tomentose with simple hairs, brown; base flat, apex rounded
to acute; scar convex, c. 1 cm in diameter; wall thin, most part free from the cupule.
Vernacular names. Sabahtikalod (Dusun, Ranau). Sarawaktekalat (Murut).
Distribution. Endemic to Borneo. In Sabah, recorded from Sapulut in Keningau, Ulu Tungud
in Beluran, and Sg. Timbulanan in Labuk Sugut districts. In Sarawak, collected from Belaga
district and Lambir NP in Miri district. Not yet reported for Brunei and Kalimantan.
Ecology. In primary and secondary lowland and hill mixed dipterocarp forests, including
riparian forest, up to 500 m altitude.
7. Lithocarpus bullatus Soepadmo

(Latin, bullatus = inflated or blistered; the leaf surface)
Reinwardtia 8 (1970) 223, l.c. (1972) 356; Cockburn l.c. (1976) 102. Type: Clemens 32715, Borneo,
Sabah, Mt. Kinabalu (holotype BO; isotypes A, BM, K, L, NY, UC).
Tree up to 25 m tall, 40 cm diameter. Bark fissured or smooth or lenticellate, greyish

brown; inner bark granular, reddish. Sapwood reddish. Twigs sparsely tomentose, lateglabrescent, sparsely minute-lenticellate. Stipules lanceolate, 13 1 mm. Leaves thickcoriaceous, bullate, rigid, sparsely tomentose or glabrescent above, sparsely greyish
appressed tomentose below; ovate, (3)4.57 24 cm, base acute, margin strongly
revolute, apex obtuse, rounded or rarely emarginate; midrib slightly raised above, strongly
raised and sparsely tomentose below; lateral veins thick, 78(10) pairs, lax, impressed
above, strongly raised below, disappearing near the leaf margin, forming an angle of c.
45 with the midrib; intercostal venation scalariform, lax, obscure on both surfaces or
slightly prominent below; petiole 47(9) mm long. Inflorescences male, female and mixed.
Male inflorescences 610 cm long; bracts and bracteoles ovate, 11.2 0.70.8 mm.
Male flowers solitary along the rachis; perianth lobes obtuse, 11.2 mm across; stamen
filaments c. 2 mm long; pistillode subglobose, c. 1 mm diameter. Female or mixed
inflorescences 510 cm long; bracts and bracteoles ovate, c. 1.5 0.9 mm. Female flowers
in clusters of 3 or solitary along the rachis; perianth lobes acute, c. 1.5 1 mm; staminodes
12; styles conical, straight, 22.5 mm long. Cupules solitary along the rachis, rarely in
clusters of 23, sessile, saucer-shaped, 0.51 1.51.8 cm, densely tomentose, lamellate;
wall woody, thin, enclosing less than half of the acorn; lamellae slightly distinct, edge
minutely denticulate, set in 58 regular lines. Acorns ovoid to conical, 1.52 cm across,
most part exserted, glabrous, dark brown, base rounded, apex abruptly acute; scar concave,
c. 1 cm diameter; wall bony, thin, most part free from the cupule.
Distribution. Endemic to Borneo. In Sabah common in Tambunan district (Kinabalu Park
and Mt. Alab) and in Beluran district. Uncommon in Sarawak, so far only known from
G. Mulu NP in Miri district. Also in Kalimantan.
Ecology. In lower and upper montane forests, at 15003000 m altitude, on organic and
ultrabasic soils.
43
8. Lithocarpus cantleyanus (King ex Hook. f.) Rehder

(Nathaniel Cantley, 18801888 Superintendent of the Singapore Botanic Gardens)
J. Arn. Arb. 1 (1919) 123, l.c. (1929) 132; Soepadmo l.c. (1970) 225, l.c. (1972) 352; Cockburn l.c.
(1972) 216, l.c. (1976) 107; Anderson l.c. 199; Whitmore, Tantra & Sutisna l.c. 155; Turner l.c. 235;
PROSEA l.c. 289; Coode et al. (eds.) l.c. 125; Argent et al. (eds.) l.c. 247. Basionym: Quercus
cantleyana King ex Hook. f., Fl. Brit. Ind. 5 (1888) 613. Lectotype (Soepadmo, 1970): Kings Collectors
5623, Peninsular Malaysia, Perak, Chanderiang (hololectotype K; isolectotypes BM, L, P, SING).
Synonyms: Pasania cantleyana (King ex Hook. f.) Gamble l.c. (1915) 434; Synaedrys cantleyana
(King ex Hook. f.) Koidzumi l.c. 190.
Tree up to 36 m tall, 70 cm diameter. Bark finely fissured or scaly or lenticellate, greyish

brown; inner bark granular, yellowish or reddish. Sapwood yellowish. Twigs glabrescent,
sparsely lenticellate. Stipules narrowly ovate to linear, 35 12 mm. Leaves thickcoriaceous, glabrous and shining above, sparsely appressed greyish tomentose below;
elliptic, (10)11.516(26.5) (4)57(11) cm, base cuneate, margin strongly revolute,
apex acute to acuminate, acumen 515 mm long; midrib strongly raised on both surfaces;
lateral veins thick, (12)1416(18) pairs, dense, strongly impressed or rarely flat above,
raised below, faintly or clearly joining near the leaf margin, forming an angle of 30
60 with the midrib; intercostal venation scalariform, lax, prominent on both surfaces;
petiole 1015 mm long. Inflorescences male, female or androgynous. Male inflorescences
1020 cm long; bracts and bracteoles narrowly lanceolate, c. 1.2 0.5 mm. Male flowers
in clusters of 34 along the rachis; perianth lobes acute, c. 1.3 0.8 mm; stamen filaments
c. 3 mm long; pistillode subglobose, c. 1.2 mm diameter. Female or androgynous
inflorescences 1015 cm long; bracts ovate, c. 0.5 0.3 mm; bracteoles narrowly ovate,
c. 0.3 0.2 mm. Female flowers solitary along the rachis; perianth lobes acute, c. 0.6
0.4 mm; staminodes 12; styles conical, slightly recurved, c. 1 mm long. Cupules solitary
along the rachis, with stalk 57 mm long, cup-shaped, 0.51 1.52(2.5) cm, densely
tomentose, lamellate; wall woody, thin, densely hairy, enclosing less than half of the
acorn; lamellae slightly distinct, edge entire or denticulate, set in 58 regular lines. Acorns
ovoid, 11.8(2) 1.52 cm, most part exserted, glabrous, brown; base rounded, apex
abruptly acute; scar shallowly concave or flat, 11.4 cm diameter; wall thin, most part
Vernacular names. Sabahtikalod (Dusun, Bundu Tuhan). Sarawakkeraki (Iban).
Distribution. Peninsular Malaysia, Singapore and Borneo. In Sabah, common in all districts.
In Sarawak, recorded for Miri, Limbang, Kapit, Sri Aman, and Kuching districts. Also
Ecology. In primary hill mixed dipterocarp forest, mainly on clay soils, up to 900 m
altitude.
9. Lithocarpus caudatifolius (Merr.) Rehder

(Latin, caudatus = ending with a tail-like appendage, folium = leaf; the leaf with caudate
apex)
J. Arn. Arb. 1 (1919) 123; Soepadmo l.c. (1970) 226, l.c. (1972) 350; Cockburn l.c. (1976) 99; Anderson
44
l.c. 199; Whitmore, Tantra & Sutisna l.c. 155; PROSEA l.c. 290. Basionym: Quercus caudatifolius
Merr., Philip. J. Sc. Bot. 3 (1908) 324. Type: Borden FB 806, the Philippines, Luzon, Bataan Prov., Mt.
Mariveles, Lamao R. (holotype BM). Synonyms: Q. minahassae Koord. ex Elmer, Leafl. Philip. Bot. 3
(1910) 941; L. minahassae (Koord. ex Elmer) Rehder l.c. (1929) 133; Synaedrys caudatifolia (Merr.)
Koidzumi l.c. 190; Q. bulusanensis Elmer, Leafl. Philip. Bot. 10 (1939) 3736; L. bulusanensis (Elmer)
A. Camus, Not. Syst. 13 (1948) 265.
Tree up to 30 m tall, 60 cm diameter; buttresses c. 0.3 m tall. Bark scaly or smooth

or fissured, greyish brown; inner bark fibrous, yellowish or brownish. Sapwood yellowish.
Twigs densely tomentose with simple erect hairs, densely lenticellate. Stipules narrowly
ovate to linear, 34 1 mm, persistent. Leaves thin-coriaceous, not rigid, sparsely to
densely tomentose with simple and stellate hairs above, densely greyish tomentose with
simple-erect and stellate hairs below; elliptic to obovate, (5.5)913(16) (2.5)3
5(7) cm, base acute to cuneate, rarely rounded, margin slightly revolute, apex caudate
or acuminate, acumen 1025 mm long, rarely acute; midrib raised on both surfaces, stronger
below; lateral veins thin, 710(11) pairs, lax, flat above, raised below, disappearing
subscalariform or reticulate, lax or dense, obscure on both surfaces or thinly prominent
below; petiole 510 mm long. Inflorescences male, androgynous or mixed. Male
inflorescences 1318 cm long; bracts ovate, c. 1 0.5 mm; bracteoles narrowly ovate,
c. 0.5 0.2 mm. Male flowers solitary along the rachis; perianth lobes obtuse, c. 1
0.8 mm; stamen filaments 1.52.5 mm long; pistillode globose, c. 5 mm diameter.
Androgynous or mixed inflorescences 1126 cm long; bracts ovate, c. 1 0.6 mm;
bracteoles subrounded, c. 0.5 mm across. Female flowers solitary along the rachis; perianth
lobes acute, c. 0.7 0.5 mm; staminodes 12; styles cylindrical, straight, c. 1 mm long.
Cupules solitary along the rachis, with stalk c. 5 mm long, cup- to saucer-shaped, 0.3
0.7(0.9) 1.22 cm, densely tomentose with simple-erect hairs, lamellate; wall bony,
thin, densely tomentose, enclosing less than half of the acorn; lamellae slightly distinct,
edge minutely denticulate, set in 57 regular lines. Acorns ovoid to conical or rarely
obovoid, 12.5 12 cm, most part exserted, densely tomentose with simple-erect hairs,
brown; base flat, apex acute; scar concave, c. 1 cm diameter; wall bony, thin, most part
Distribution. Borneo and the Philippines. Very common in Sabah. In Sarawak, documented
only from Kuching and Marudi districts. Also in Kalimantan.
Ecology. In primary and secondary mixed dipterocarp forest to lower montane forest,
10. Lithocarpus clementianus (King ex Hook. f.) A. Camus

(Sir Cecil Clementi Smith, former Governor of the Straits Settlements)
Riviera Scientific 18 (1932) 40; Soepadmo l.c. (1970) 229, l.c. (1972) 365; Cockburn l.c. (1972) 216,
l.c. (1976) 103; Anderson l.c. 199; Corner l.c. 339; Whitmore, Tantra & Sutisna l.c. 155; Turner l.c.
235; PROSEA l.c. 290. Basionym: Quercus clementiana King ex Hook. f. l.c. (1888) 614. Lectotype
(Soepadmo, 1970): Maingay 1529, Peninsular Malaysia, Penang (hololectotype K; isolectotypes A,
L). Synonyms: Pasania clementiana (King ex Hook. f.) Gamble l.c. (1915) 439; Synaedrys clementiana
(King ex Hook. f.) Koidzumi l.c. 191; Q. teysmannii (non Blume) Heine in Feddes Rep. 54 (1951)
225.
45
Tree up to 36 m tall, 70 cm diameter. Bark deeply fissured or cracked, brownish; inner

bark reddish or brownish. Sapwood yellowish. Twigs densely or sparsely appressed
tomentose, finely fissured or sparsely large-lenticellate. Stipules ovate, c. 2 1 mm.
Leaves thick-coriaceous, subglabrous above, sparsely yellowish appressed tomentose with
simple hairs below; elliptic to elliptic-lanceolate, (10)1419(22) (3)4.57.5 cm, base
acute, margin strongly revolute, apex acute to acuminate, acumen 1015 mm long; midrib
broad, strongly raised on both surfaces; lateral veins thick, (8)1012(14) pairs, lax,
flat or impressed above, strongly raised below, disappearing near the leaf margin, forming
an angle of 4050 with the midrib; intercostal venation subscalariform, dense, obscure
above, prominent below; petiole 510(12) mm long. Inflorescences male, androgynous
or mixed. Male inflorescences 712 cm long; bracts subrounded, c. 1 0.8 mm; bracteoles
rounded, c. 0.5 mm across. Male flowers in clusters of 3 along the rachis; perianth lobes
obtuse, c. 1.5 1 mm; stamen filaments c. 1.5 mm long; pistillode subglobose, c. 1
mm diameter. Androgynous or mixed inflorescences 810 cm long; bracts and bracteoles
ovate, c. 0.5 0.30.4 mm. Female flowers in clusters of 3 or solitary along the rachis;
perianth lobes acute, 0.50.8 0.50.6 mm; staminodes 10; styles conical, straight, c.
1 mm long. Cupules in clusters of 23 or solitary along the rachis, sessile, cup-shaped,
11.5 2.33.5 cm, densely appressed tomentose with stellate hairs, lamellate; wall woody,
thick, enclosing up to half of the acorn; lamellae distinct, edge minutely denticulate, set
in 610 regular lines. Acorns depressed subglobose, 12 23.5 cm, most part exserted,
densely covered with greyish, simple, long hairs, purplish; base flat, apex acute; scar
deeply concave, c. 1.5 cm diameter; wall woody, thick, most part free from the cupule.
Distribution. Peninsular Malaysia and Borneo. In Sabah, very common in Ranau district
and also recorded for Keningau, Sipitang and Lahad Datu districts. In Sarawak, widely
distributed throughout. Also in Kalimantan.
Ecology. In hill mixed dipterocarp forest to lower montane forest on sandy soils, up
to 1800m altitude.
11. Lithocarpus confertus Soepadmo

(Latin, confertus = crowded; the inflorescences and infructescences)
Tantra & Sutisna l.c. 155. Type: Clemens 29265, Borneo, Sabah, Mt. Kinabalu, Tenompok (holotype
L).
Tree up to 30 m tall, 60 cm diameter. Bark densely lenticellate or scaly, greyish brown;

inner bark fibrous, reddish. Sapwood whitish. Twigs densely tomentose with stellate hairs,
sparsely to densely lenticellate. Stipules lanceolate, 23 1 mm. Leaves thin-coriaceous,
subglabrous above, sparsely appressed yellowish brown tomentose below; ovate to elliptic,
(6)912(13) (3)46 cm, base rounded or obtuse, margin revolute, not undulate,
apex acute to acuminate, acumen 1015 mm long; midrib faintly raised on both surfaces,
stronger below; lateral veins thick, (7)910(11) pairs, dense, flat or impressed above,
strongly raised below, disappearing or faintly joining near the leaf margin, forming an
angle of 4560 with the midrib; intercostal venation scalariform, dense, obscure above,
prominent below; petiole 510 mm long. Inflorescences male, androgynous or mixed.
Male inflorescences 1015 cm long; bracts and bracteoles ovate, c. 0.5 0.3 mm. Male
46
flowers in clusters of 3 along the rachis; perianth lobes obtuse, c. 1.2 1 mm; stamen
filaments c. 3 mm long; pistillode subglobose, c. 1 mm diameter. Androgynous or mixed
inflorescences 510 cm long; bracts and bracteoles narrowly ovate, 0.51.2 0.30.6
mm. Female flowers solitary along the rachis; perianth lobes acute, 0.40.5 0.3 mm;
staminodes 12; styles cylindrical, straight, c. 1.5 mm long. Cupules solitary along the
rachis, subsessile, saucer-shaped, 0.20.5 1.21.7 cm, densely tomentose with stellate
and simple hairs, lamellate; wall bony, thin, enclosing less than half of the acorn; lamellae
slightly distinct, edge minutely denticulate, set in 68 concentric lines. Acorns depressed
ovoid, 11.7 1.42 cm, most part exserted, glabrous except for the apex which is densely
tomentose with stellate hairs, dark brown or purplish; base rounded, apex acute; scar
concave, c. 1 cm diameter; wall bony, thin, most part free from the cupule.
Vernacular name. Sarawaksaled birar (Kelabit).
Distribution. Endemic to Borneo. In Sabah, very common in Ranau, Sandakan, Tambunan,
and Keningau districts. In Sarawak, uncommon, recorded for Bario, Lawas, Kapit and
Sri Aman districts. Also in Kalimantan.
Ecology. In primary and secondary mixed dipterocarp, peatswamp and lower montane
forests, at 9001200 m altitude; on sandstone-derived and clay soils.
Uses. In Sabah, trunks and branches are used as bed logs in mushroom cultivation.
12. Lithocarpus confragosus (King ex Hook. f.) A. Camus

(Latin, con- = together, fragosus = crumpled; the external surface of cupule wall)
l.c. (1976) 107; Anderson l.c. 199; Whitmore, Tantra & Sutisna l.c. 155; Turner l.c. 235; PROSEA l.c.
290. Basionym: Quercus confragosa King ex Hook. f. l.c. (1888) 616. Lectotype (Soepadmo, 1970):
Kings Collectors 8123, Peninsular Malaysia, Perak, Gopeng district (holotype K; isotypes A, L, SING).
Synonym: Pasania confragosa (King ex Hook. f.) Schottky, Bot. Jahrb. 47 (1912) 662.
Tree up to 30 m tall, 65 cm diameter. Bark smooth or scaly, lenticellate, greyish brown;

inner bark fibrous, yellowish or reddish. Sapwood whitish. Twigs tomentose, lateglabrescent, densely warty lenticellate. Stipules linear, 812 3 mm. Leaves thickcoriaceous, glabrous above, densely appressed yellowish tomentose below; elliptic-oblong
or broadly elliptic, (10)1218(25) 57(10) cm, base acute to cuneate, symmetrical,
margin revolute, apex acute to acuminate, acumen 510(15) mm long; midrib raised
above, strongly raised below; lateral veins thick, 79(11) pairs, lax, impressed or flat
above, strongly raised below, disappearing near the leaf margin, forming an angle of
4060 with the midrib; intercostal venation subscalariform, lax, obscure on both surfaces
or thinly prominent below; petiole 1020 mm long. Inflorescences male, female or
androgynous. Male inflorescences 1016 cm long; bracts and bracteoles linear to narrowly
lanceolate, c. 1 0.3 mm. Male flowers in clusters of 23 along the rachis; perianth
lobes obtuse, c. 1 0.5 mm; stamens filaments 1.53 mm long; pistillode subglobose,
c. 0.5 mm diameter. Female or androgynous inflorescences 915 cm long; bracts and
bracteoles elliptic, c. 1 0.5 mm. Female flowers solitary along the rachis; perianth
47
lobes rounded, 0.51 mm across; staminodes 12; styles cylindrical, straight, c. 1 mm

long. Cupules solitary along the rachis, with stalk 510 mm long, depressed ovoid to
depressed globose, 23 35.5 cm, densely tomentose, irregularly tuberculate and ridged;
wall woody thick, completely enclosing the acorn except for an apical opening c. 0.5
cm diameter; tubercles crumpled together, distinct, set irregularly. Acorns depressed ovoid
to depressed globose, 1.52.5 24 cm, free part densely tomentose with simple hairs,
brown; base flat, apex flat or depressed, umbonate at the centre; scar flat, c. 2 cm diameter;
wall woody, thick, most part free from the cupule.
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah known by a few
collections from Tambunan, Lahad Datu, Ranau, Sandakan, and Tawau districts. In Sarawak,
recorded for Kuching, Miri and Sri Aman districts. Also in Kalimantan.
13. Lithocarpus conocarpus (Oudem.) Rehder

(Greek, konos = cone, karpos = fruit; the conical acorn)
(1976) 108; Anderson l.c. 200; Corner l.c. 340; Whitmore, Tantra & Sutisna l.c. 155; Turner l.c. 235;
Kessler & Sidiyasa, Trees of the Balikpapan-Samarinda Area, East Kalimantan (1994) 140; PROSEA
l.c. 291; Coode et al. (eds.) l.c. 125; Argent et al. (eds.) l.c. 247. Basionym: Quercus conocarpa
Oudem., Versl. Med. Kon. Ak. Wet. Natuurk. 12 (1861) 206. Type: Junghuhn s.n. (1857), Java, Mt.
Malabar (holotype BO; isotype L). Synonyms: Cyclobalanus conocarpa (Oudem.) Oerst. l.c. (1867)
81; Pasania conocarpa (Oudem.) Schottky l.c. (1913) 357; Synaedrys conocarpa (Oudem.) Koidzumi
l.c. 191.
Tree up to 45 m tall, 90 cm diameter. Bark scaly, greyish brown; inner bark granular,
reddish or brownish. Sapwood yellowish. Twigs densely tomentose with simple-erect and
stellate hairs, late-subglabrescent, sparsely to densely lenticellate. Stipules ovate, 23
1.5 mm. Leaves coriaceous, densely tomentose with stellate and simple hairs above, densely
appressed yellowish tomentose with simple-erect and stellate hairs below; elliptic or
obovate, (6)811(14) (2)34.5(5.5) cm, base acute, sometimes asymmetrical, margin
revolute, apex acute to acuminate, acumen 1015(20) mm long; midrib raised on both
surfaces; lateral veins thin, 1012(16) pairs, dense or lax, flat or impressed above, raised
below, clearly joining near the leaf margin, forming an angle of 4060 with the midrib;
intercostal venation subscalariform, dense, obscure above, prominent below; petiole 5
8 mm long. Inflorescences male, androgynous or mixed. Male inflorescences 816 cm
long; bracts and bracteoles linear, 0.31.2 0.10.4 mm. Male flowers solitary along
the rachis; perianth lobes rounded, c. 1 mm across; stamen filaments c. 2 mm long; pistillode
globose, 0.51 mm diameter. Androgynous or mixed inflorescences 712 cm long; bracts
ovate, c. 1 0.5 mm; bracteoles elliptic, c. 0.6 0.4 mm. Female flowers solitary along
the rachis; perianth lobes acute c. 1 0.5 mm; staminodes 12; styles cylindrical, straight,
c. 1 mm long. Cupules solitary and distantly spaced along the rachis, with stalk 3
8 mm long, cup-shaped, 0.40.8(1) 1.32 cm, densely tomentose with simple, erect
hairs, lamellate; wall thin, densely tomentose, enclosing up to half of the acorn; lamellae
slightly distinct, edge denticulate or rarely entire, set in 56 regular lines. Acorns conical
or ovoid, 1.62 1.31.5 cm, most part exserted, densely tomentose with simple, erect
48
hairs, rarely glabrescent, dark brown; base rounded, apex acute; scar concave or flat,
0.81.2 cm diameter; wall woody, thin, most part free from the cupule.
Vernacular names. Sabahtikalod (Dusun, Bundu Tuhan). Sarawakpadeh siah, saled
(Kelabit), palan (Kenyah), piraus (Bidayuh).
Distribution. Sumatra, Peninsular Malaysia, Singapore, Java, and Borneo. Very common
and widely distributed throughout Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. In lowland to lower montane forests, including limestone forest, mainly on clayrich soils, up to 1400 m altitude.
Use. The timber is locally used for house-building.
14. Lithocarpus coopertus (Blanco) Rehder

(Latin, coopertus = completely concealed; the acorn)
(1976) 105; Anderson l.c. 200; Whitmore, Tantra & Sutisna l.c. 155; Kessler & Sidiyasa l.c. 140;
PROSEA l.c. 291; Turner l.c. 235; Coode et al. (eds.) l.c. 125; Argent et al. (eds.) l.c. 247. Basionym:
Quercus cooperta Blanco, Fl. Filip. (1845) 503. Neotype: Elmer 14012, the Philippines, Mindanao,
Agusan Prov., Cabadbaran (holoneotype UC; isoneotypes A, BM, L, NY). Synonyms: Q. fernandezii
Vidal, Sinops. (1883) 41, t. 92; Q. reflexa King l.c. (1889) 78, t. 72; Castanea cooperta (Blanco)
Oerst. l.c. (1871) 379; Synaedrys cooperta (Blanco) Koidzumi l.c. 186; Castanopsis costata (non A.
DC.) F.Vill., Nov. App. Fl. Filip. 3 (1880) 209; S. reflexa (King) Koidzumi l.c. 187; Castanopsis
reflexa (King) Rehder l.c. (1919) 122; L. reflexa (King) A. Camus l.c. (1932) 41; Q. pruinosa (non
Blume) Elmer, Leafl. Philip. Bot. 3 (1910) 940; Q. boholensis Merr., Philip. J. Sc. 29 (1926) 476; L.
boholensis (Merr.) Rehder l.c. (1929) 132.
Tree up to 35 m tall, 70 cm diameter. Bark smooth or flaky, sometimes lenticellate,

greyish brown; inner bark fibrous, reddish. Sapwood whitish. Twigs densely tomentose
with stellate hairs, late-subglabrescent, sparsely minute-lenticellate. Stipules linear, 2
3 1 mm. Leaves thin-coriaceous, sparsely to densely appressed tomentose with stellate
hairs on both surfaces; elliptic or ovate, (5)914(17) (2)47 cm, base rounded to
acute, usually asymmetrical, margin revolute, apex acute to acuminate, acumen 1020 mm
long; midrib raised on both surfaces, stronger below; lateral veins thin, (8)1013(16)
pairs, lax, flat or impressed above, raised below, disappearing near the leaf margin, forming
an angle of 4060 with the midrib; intercostal venation scalariform, dense, obscure on
both surfaces or thinly prominent below; petiole up to 5 mm long. Inflorescences male,
androgynous or mixed. Male inflorescences 715 cm long; bracts and bracteoles
narrowly ovate, triangular, c. 1 0.5 mm. Male flowers solitary along the rachis; perianth
lobes acute, c. 1 0.5 mm; stamen filaments c. 2 mm long; pistillode globose, c. 1.5
mm diameter. Androgynous or mixed inflorescences 715 cm long; bracts narrowly ovate,
triangular, c. 1 0.5 mm; bracteoles subrounded, c. 0.5 mm across. Female flowers
solitary along the rachis; perianth lobes obtuse, 0.40.5 mm across; staminodes 12; styles
cylindrical, straight, c. 1 mm long. Cupules solitary along the rachis, subsessile or rarely
with a stalk c. 5 mm long, ovoid to conical, 1.52 1.52.5 cm, densely tomentose,
covered with irregularly set, long, recurved, spine-like appendages; wall bony, thin, densely
hairy, completely enclosing the acorn. Acorns ovoid to conical, 1.52 cm diameter, free
49
part densely tomentose, brown; base flat, apex acute; scar deeply concave to flat, 1
1.5 cm diameter; wall bony, thin, most part free from the cupule.
Distribution. Peninsular Malaysia, Borneo and the Philippines. Common and widespread
throughout Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. In lowland to hill mixed dipterocarp forest, including kerangas and swamp forests,
Uses. In Borneo, the timber is used by local people for firewood and in house construction.
15. Lithocarpus corneri S. Julia & Soepadmo

(E.J.H. Corner, 19061996, former prominent Professor of Tropical Botany at the University of Cambridge, U.K.)
Gard. Bull. Sing. 50 (1998) 128. Type: Berhaman SAN 132620, Borneo, Sabah, Tenom district, Lumaku
FR (holotype SAN; isotypes KEP, L, SAR).
Tree up to 15 m tall, 30 cm in diameter. Bark rough, brown. Twigs sparsely tomentose,

glabrescent, greyish brown, sparsely lenticellate, sometimes smooth. Stipules linear, 3
4 1 mm, persistent. Leaves thin-coriaceous, sparsely appressed yellowish tomentose
above, densely appressed yellowish tomentose below; elliptic, (6)812 34.5 cm, base
acute, margin revolute, apex acuminate, acumen 510 mm long; midrib raised on both
surfaces, stronger above; lateral veins thin, 58 pairs, lax, flat above, slightly raised below,
disappearing near the leaf margin, forming an angle of 4560 with the midrib; intercostal
venation subscalariform, dense, obscure on both surfaces or slightly prominent below;
petiole 510 mm long. Inflorescences and flowers unknown. Cupules solitary along the
rachis, sessile, obconical, 2.54.5 3.55.5 cm, base attenuate, apex flat, densely appressed
tomentose, lamellate; wall woody, 23 cm thick, completely enclosing the acorn except
for an apical opening 12 cm diameter; lamellae distinct, edge entire, set in 1215 regular
lines, denser towards the apex. Acorns obconical, 24 35 cm, free apical part densely
tomentose with simple and stellate hairs, brown; wall woody, thick, most part adnate
to the cupule.
Distribution. Endemic to Sabah, Borneo. So far only known from Lumaku FR in Tenom
district and Mark Pang area in Ranau district.
Ecology. In primary and secondary forests on hillsides.
16. Lithocarpus daphnoideus (Blume) A. Camus

(Greek, daphnoideus = resembling those of Daphne, a genus in the Thymelaeaceae; the
leaves)
Riviera Scientific 18 (1932) 40; Warburg, Kew Bull. 20 (1936) 19; Soepadmo l.c. (1970) 234, l.c.
(1972) 352; Cockburn l.c. (1972) 218; Anderson l.c. 200; Whitmore, Tantra & Sutisna l.c. 156; Turner
l.c. 236; Coode et al. (eds.) l.c. 125. Basionym: Quercus daphnoidea Blume, Fl. Jav. Cupul. (1829)
28. Type: van Hasselt s.n. (= RHL Sheet No. 901310119), W Java, Mt. Pajung (holotype L; isotypes
50
CGE, K). Synonyms: Cyclobalanus daphnoidea (Blume) Oerst. l.c. (1867) 81; Synaedrys daphnoidea
(Blume) Koidzumi l.c. 191; Pasania daphnoidea (Blume) S. Moore, J. Bot. 63 (1925) 114; Quercus
nitida Blume, Mus. Bot. Lugd.-Bat. 1 (1850) 294; Cyclobalanus nitida (Blume) Oerst. l.c. (1867) 81; L.
nitida (Blume) A. Camus, Bull. Soc. Bot. Fr. 92 (1945) 255; Q. poculiformis Seemen, Bot. Jahrb. 27,
Beibl. 64 (1900) 13; S. poculiformis (Seemen) Koidzumi l.c. 192; L. poculiformis (Seemen) A. Camus
l.c. (1932) 41; L. sarawakensis E.F. Warb. l.c. 19.
Tree up to 24 m tall, 50 cm diameter; buttresses c. 1 m tall. Bark smooth or fissured,

greyish brown; inner bark granular, reddish. Sapwood greyish. Twigs sparsely tomentose,
late-glabrescent, smooth or sparsely to densely minute-lenticellate. Stipules linear, 34
1 mm. Leaves thin-coriaceous, glabrous above, sparsely appressed yellowish tomentose
below; elliptic or broadly oblanceolate, (10)1316(18) 46 cm, base acute to cuneate
or rarely rounded, margin revolute, apex acuminate, acumen 1015 mm long; midrib raised
on both surfaces; lateral veins thin, (9)1013 pairs, lax, slightly raised or flat above,
raised below, faintly joining near the leaf margin, forming an angle of 3045 with the
midrib; intercostal venation subscalariform, dense, obscure above and prominent below
or prominent on both surfaces; petiole (7)1015 mm long. Inflorescences male or
androgynous. Male inflorescences 1315 cm long; bracts and bracteoles ovate, 0.60.8
0.30.4 mm. Male flowers solitary along the rachis; perianth lobes acute, c. 1.2
0.8 mm; stamen filaments c. 2.5 mm long; pistillode globose, c. 1 mm diameter.
Androgynous inflorescences 1015 cm long; bracts broadly ovate, c. 1.4 1 mm; bracteoles
broadly ovate, c. 0.8 0.6 mm. Female flowers in clusters of 3 or solitary along the
rachis; perianth lobes acute, 0.60.8 0.5 mm; staminodes 10; styles conical, straight,
c. 1 mm long. Cupules in clusters of 2 or solitary along the rachis, sessile, cup-shaped,
12 1.52.5 cm, densely tomentose, lamellate; wall woody, thick, enclosing less than
half of the acorn; lamellae distinct, edge denticulate, set in 810 regular lines. Acorns
ovoid to conical, 22.5 1.82.5 cm cm, most part exserted, sparsely simple-hairy, dark
brownish or red-brown; base flat, apex abruptly acute; scar flat to concave, 11.5 cm
diameter; wall woody, thick, most part free from the cupule.
Distribution. Sumatra, Peninsular Malaysia, Java, and Borneo. In Sabah, recorded for
Ranau, Keningau, and Tawau districts. In Sarawak, only known from Bt. Raya in Kapit
district. Also in Brunei and Kalimantan.
Ecology. In primary hill mixed dipterocarp forest to lower montane forest, up to 1400 m
altitude.
17. Lithocarpus dasystachyus (Miq.) Rehder

(Greek, dasus = thickly hairy, stachyus = spike; the densely pubescent inflorescences)
J. Arn. Arb. 1 (1919) 124; Soepadmo l.c. (1970) 234, l.c. (1972) 350; Cockburn l.c. (1976) 102; Anderson
l.c. 200; Whitmore, Tantra & Sutisna l.c. 156; Coode et al. (eds.) l.c. 125; Argent et al. (eds.) l.c. 248.
Basionym: Quercus dasystachya Miq., Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 221. Type: de Vries s.n.,
(1860) (= RHL Sheet No. 901310124), Borneo, Kalimantan, Pontianak (holotype L; isotypes BO, U).
Synonyms: Pasania dasystachya (Miq.) Schottky l.c. (1913) 356; Synaedrys dasystachya (Miq.)
Koidzumi l.c. (1916) 194; P. winkleriana Schottky l.c. 357; Q. winkleriana (Schottky) Merr., EB (1921)
216; L. winkleriana (Schottky) A. Camus l.c. (1931) 42.
51
Tree up to 20 m tall, 30 cm diameter; stilt-roots occasionally present. Bark smooth or

flaky or fissured, greyish brown; inner bark granular, brownish. Sapwood yellowish. Twigs
sparsely to densely stellate-hairy, smooth or sparsely minute-lenticellate. Stipules ovate,
deltoid to linear, 34 12 mm. Leaves thick-coriaceous, rigid, densely stellate-hairy
or sparsely tomentose, or rarely glabrescent above, densely brownish tomentose with simpleerect and stellate hairs below; ovate, broadly ovate, elliptic or obovate, (9)1319(23)
(4.5)68(10) cm, base acute or rounded, margin strongly revolute, apex acuminate,
acumen 1025 mm long; midrib strongly raised on both surfaces; lateral veins thin,
(8)1015 pairs, lax, strongly impressed or flat above, raised below, faintly joining
or disappearing near the leaf margin, forming an angle of 4060 with the midrib;
intercostal venation subscalariform, lax, rarely dense, obscure above, thinly prominent
below; petiole 515 mm long. Inflorescences male, androgynous or mixed. Male
inflorescences 13.525 cm long; bracts and bracteoles broadly ovate, 0.20.3 mm across.
Male flowers solitary along the rachis; perianth lobes acute, c. 0.5 0.3 mm; stamen
filaments 23 mm long; pistillode subglobose, 0.71 mm diameter. Androgynous or mixed
inflorescences 1115 cm long; bracts and bracteoles ovate, c. 1 0.5 mm. Female flowers
solitary along the rachis; perianth lobes broadly ovate, obtuse, c. 0.8 0.5 mm; staminodes
12; styles cylindrical, straight, c. 1.5 mm long. Cupules solitary along the rachis, with
stalk 25 mm long, saucer- to cup-shaped, 0.50.9 11.5 cm, densely tomentose, lamellate;
wall bony, thin, enclosing less than half of the acorn; lamellae slightly distinct, edge
minutely denticulate, set in 78 regular lines. Acorns ovoid to conical, 1.21.5 1
1.7(2) cm, most part exserted, sparsely simple-erect hairy or glabrescent, dark brown;
base flat, apex abruptly acute; scar deeply concave, c. 1 cm diameter; wall bony, thin,
most part free from the cupule.
Vernacular names. Sarawakbangas (Matu Malay), empenit padang (Malay).
Distribution. Peninsular Malaysia and Borneo. In Sabah, known mainly from Sandakan
and Lahad Datu districts. In Sarawak, very common in Kuching, Miri, Sri Aman, and
Sibu districts. Also in Brunei and Kalimantan.
Ecology. Frequent in kerangas and peatswamp forests, but also found in primary hill
mixed dipterocarp forest on ultrabasic soils, up to 1000 m altitude.
18. Lithocarpus echinifer (Merr.) A. Camus
Fig. 9.
(Latin, echinus = straight prickles or spines, fer = bearing; the cupule)

Bull. Soc. Bot. France 81 (1934) 818; Soepadmo l.c. (1970) 235, l.c. (1972) 333; Cockburn l.c. (1976)
103; Anderson l.c. 200; Whitmore, Tantra & Sutisna l.c. 156; PROSEA l.c. 292; Coode et al. (eds.)
l.c. 125. Basionym: Quercus echinifera Merr., PEB (1929) 43. Lectotype: Elmer 21627, Borneo,
Sabah, near Tawau (hololectotype A; isolectotypes BM, BR, K, L, P, U).
Tree up to 35 m tall, 80 cm diameter. Bark fissured or smooth, greyish brown; inner

bark granular, yellowish to brownish. Sapwood yellowish. Twigs sparsely tomentose, warty
lenticellate. Stipules linear, 510 25 mm. Leaves thick-coriaceous, glabrous above,
densely greyish tomentose with appressed hairs below; elliptic, oblong or broadly ovate,
1018 (4.5)710(11) cm, base acute to cuneate, margin revolute, apex acute or
acuminate, acumen 510 mm long; midrib raised on both surfaces, stronger below; lateral
52
vi
veins thin, 1013 pairs, lax, flat above, slightly raised below, disappearing near the leaf
margin, forming an angle of 4560 with the midrib; intercostal venation subscalariform,
dense, obscure on both surfaces or slightly prominent below; petiole 1525 mm long.
Inflorescences male or female. Male inflorescences 621 cm long; bracts narrowly ovate
to linear, 1.52 0.5 mm; bracteoles ovate, c. 1 0.5 mm. Male flowers solitary along
the rachis; perianth lobes obtuse, c. 1.5 1 mm; stamen filaments 35 mm long; pistillode
globose, c. 1.5 mm diameter. Female inflorescences 918 cm long; bracts broadly ovate,
c. 1 0.8 mm; bracteoles rounded, c. 1 1 mm. Female flowers solitary along the
rachis; perianth lobes acute, c. 1.2 0.8 mm; staminodes 12; styles cylindrical, straight,
c. 1 mm long. Cupules solitary along the rachis, sessile or with stalk c. 5 mm long,
obconical, 23 2.54 cm, densely tomentose, sparsely but regularly armed with prickles
or spine-like, mostly recurved appendages; wall bony, thin, hairy, enclosing the acorn
completely except for an apical opening c. 1 cm diameter. Acorns subhemispherical, 2
3 34 cm, free part sparsely tomentose, brown; base rounded, apex flat, umbonate;
wall woody, thick, most part adnate to the cupule.
Distribution. Endemic to Borneo. In Sabah, widely distributed in all districts. In Sarawak,
recorded mainly for Miri and Kapit districts. Also in Brunei.
Ecology. In primary and secondary hill mixed dipterocarp forests, including riparian forest,
up to 1000 m altitude, mainly on clay soils.
19. Lithocarpus echinulatus Soepadmo

(Latin, echinulatus = armed with short, slender prickles or spine-like appendages; the cupule)
PROSEA l.c. 293. Type: Kostermans 13438, Borneo, Kalimantan, Sangkulirang district, Mt. Medadam
(holotype BO; isotypes BM, K, KEP, L, SING).
Tree up to 40 m tall, 80 cm diameter; buttresses c. 2 m tall. Bark smooth, brownish.

Sapwood brownish. Twigs sparsely appressed tomentose, smooth or sparsely lenticellate.
Leaves thin-coriaceous, glabrescent above, sparsely brownish or greyish appressed tomentose
below; elliptic to oblong, (11)1622(23) (4.5)57.5 cm, base attenuate, acute or
cuneate, margin slightly revolute, apex acuminate, acumen 510 mm long; midrib slightly
raised above, strongly raised below; lateral veins thin, (6)810(18) pairs, lax, slightly
raised or impressed or flat above, strongly raised below, disappearing near the leaf margin,
forming an angle of 3045 with the midrib; intercostal venation subscalariform, lax,
obscure on both surfaces or thinly prominent below; petiole 618 mm long. Inflorescences
male or androgynous. Male inflorescences 2025 cm long; bracts ovate, 0.81 0.3
0.6 mm; bracteoles broadly ovate, c. 0.5 mm across. Male flowers solitary along the
rachis; perianth lobes obtuse, 11.5 1 mm; stamen filaments c. 2 mm long; pistillode
subglobose, 0.50.8 mm diameter. Androgynous inflorescences 1126 cm long; bracts
and bracteoles ovate, c. 1 0.30.6 mm. Female flowers solitary along the rachis; perianth
lobes acute, 0.81 0.30.7 mm; staminodes 10; styles conical, straight, c. 2 mm long.
Cupules solitary along the rachis, sessile, saucer-shaped, 0.61 2.23.2 cm, densely
tomentose, echinulate; wall woody, thick, enclosing less than half of the acorn; short
and slender prickles or spine-like appendages distinct, straight or recurved, sharp-tipped,
54
densely but irregularly set. Acorns depressed globose, 1.32.5 2.23.3 cm, most part
exserted, sparsely tomentose with simple hairs, brown; base rounded, apex abruptly
rounded-apiculate; scar flat to concave, 2.32.6 cm diameter; wall woody, thick, most
Distribution. Endemic to Borneo. In Sarawak, uncommon, known from Dulit Range in
Belaga district and Sg. Joh and Ulu Katibas in Kapit district. So far there is no record
for Sabah. Also in Kalimantan.
Ecology. In lowland to hill mixed dipterocarp forests on igneous and clay soils, usually
along riverbanks, up to 400 m altitude.
20. Lithocarpus elegans (Blume) Hatus. ex Soepadmo

(Latin, elegans = elegant; the cupules and acorns)
Reinwardtia 8 (1970) 236, l.c. (1972) 366; Cockburn l.c. (1972) 219, l.c. (1976) 100; Anderson l.c.
200; Corner l.c. 340; Whitmore, Tantra & Sutisna l.c. 156; Turner l.c. (1995) 236; PROSEA l.c. 293;
Coode et al. (eds.) l.c. 125. Basionym: Quercus elegans Blume, Verh. Batav. Gen. K. & W. 9 (1823)
208. Type: Blume s.n., Java, Bantam Residency, Sadjira (holotype L; isotypes CGE, K, P). Synonyms:
Q. spicata (non Humb. & Bonpl. 1809) Smith in Rees, Cyclop. (1814) 12; Q. racemosa (non Lam.
1783) Jack, Mal. Misc. 2 (1822) 86; Q. arcaula Hamilt. ex G. Don var. racemosa (Jack) Blume l.c.
(1850) 290; Q. spicata (non Humb. & Bonpl.) Smith var. racemosa (Jack) Miq., Ann. Mus. Lugd.-Bat
1 (1863) 106; Q. depressa Blume l.c. (1823) 209, t. 1; Q. spicata (non Humb. & Bonpl.) Smith var.
depressa (Blume) King, Ann. Roy. Bot. Gard. Calc. 2 (1889) 48, t. 43; Q. glaberrima Blume l.c.
(1823) 210, t. 2; Q. spicata (non Humb. & Bonpl.) Smith var. glaberrima (Blume) Miq. l.c. (1856)
848; L. spicata (Smith) Rehder var. glaberrima (Blume) A. Camus l.c. (1949) 103, t. 482; L. spicata
(Smith) Rehder var. elegans (Blume) A. Camus l.c. (1949) 102, t. 481; Q. placentaria Blume, l.c.
(1825) 518; Q. spicata (non Humb. & Bonpl.) Smith var. placentaria (Blume) Schottky l.c. (1912)
664; Q. grandifolius D. Don in Lamb., Gen. Pin. 2 (1824) 27, t. 8; L. grandifolius (D. Don) S.N.
Biswas, Bull. Bot. Surv. India 10 (1968) 258; P. spicata (Smith) Oerst. var. placentaria (Blume)
Oerst. l.c. (1867) 83; Pasania spicata (Smith) Oerst. var. placentaria (Blume) Schottky l.c. (1912)
664; L. spicata (Smith) Rehder var. placentaria (Blume) Rehder l.c. (1929) 133; Q. glomerata Roxb.,
Fl. Ind. 3 (1832) 640; P. glomerata (Roxb.) Oerst. l.c. (1891) 379; Q. anceps Korth. l.c. (1844) 204;
Q. microcalyx Korth. l.c. (1844) 206; Q. arcaula Hamilt. ex G. Don var. microcalyx (Korth.) Blume
l.c. (1850) 290; Q. spicata (non Humb. & Bonpl.) Smith var. microcalyx (Korth.) Miq. l.c. (1856)
848; P. spicata (Smith) Oerst. var. microcalyx (Korth.) Gamble l.c. (1915) 421; L. microcalyx (Korth.)
A. Camus l.c. (1945) 83; Q. pyrifolia Blume l.c. (1850) 304; Q. sphacelata Blume l.c. (1850) 304; Q.
gracilipes Miq., Fl. Ned. Ind. 4, Suppl. (1861) 347; Q. spicata (non Humb. & Bonpl.) Smith var.
gracilipes (Miq.) Miq. l.c. (1863) 106; P. spicata (Smith) Oerst. var. gracilipes (Miq.) Schottky l.c.
(1912) 664; L. spicata (Smith) Rehder var. gracilipes (Korth.) Rehder l.c. (1919) 131; Q. spicata (non
Humb. & Bonpl.) Smith var. latifolia Scheff. l.c. (1870) 359; Q. rhioensis Hance l.c. 198; L. rhioensis
(Hance ) A. Camus l.c. (1932) 40.
Tree up to 30 m tall, 70 cm diameter. Bark deeply fissured or lenticellate, greyish brown;

inner bark granular, reddish or brownish. Sapwood yellowish or brownish. Twigs sparsely
tomentose, late-glabrescent, sparsely to densely large-lenticellate. Stipules ovate to linear,
36 12 mm. Leaves coriaceous, rarely thick-coriaceous, glabrous on both surfaces;
narrowly to broadly obovate or elliptic, (9)1217(21) 36(8) cm, base acute or
cuneate, margin revolute, apex bluntly acute or acuminate, acumen 1015(20) mm long;
55
midrib raised on both surfaces; lateral veins thin, (11)1316(18) pairs, lax, slightly
raised or flat or sometimes impressed above, raised below, faintly joining near the leaf
margin, forming an angle of 4060 with the midrib; intercostal venation subscalariform
or rarely reticulate, lax, obscure above, prominent below; petiole 715 mm long.
Inflorescences male, androgynous or mixed. Male inflorescences 1020 cm long; bracts
ovate, c. 1.5 1 mm; bracteoles elliptic, c. 0.5 0.2 mm. Male flowers solitary but
closely arranged along the rachis; perianth lobes rounded, 11.2 mm across; stamens
filaments c. 2 mm long; pistillode globose, c. 1 mm diameter. Androgynous or mixed
inflorescences 1220 cm long; bracts and bracteoles broadly ovate, 11.3 0.91 mm.
Female flowers in cluster of (2)37(10) along the rachis; perianth lobes acute, c. 0.7
0.5 mm; staminodes 12; styles cylindrical, straight, c. 1 mm long. Cupules in clusters
37 or rarely solitary along the rachis, sessile or with stalk c. 5 mm long, cup-shaped,
11.5 (1)22.5(3) cm, densely appressed tomentose, scaly; wall woody, thin, enclosing
up to half of the acorn; scale-like appendages distinct, appressed, woody, imbricate,
set in regular lines. Acorns ovoid or depressed ovoid to subglobose, (1)1.52(2.3)
(1)22.5(3) cm, most part exserted, glabrous, brownish; base flat, apex rounded;
scar flat or concave, 11.5 cm diameter; wall woody, thin, most part free from the cupule.
Vernacular names. Sabahtihaig (Dusun, Banggi). Sarawakberungulad (Kelabit).
Distribution. India, Burma, Thailand, Laos, Cambodia, Vietnam, China, Sumatra, Peninsular
Malaysia, Singapore, Java, Borneo, and Sulawesi. Common and widespread throughout
Sabah. In Sarawak, uncommon, only known from Sg. Menalio in Marudi district and
G. Gading in Lundu district. Also in Kalimantan.
Ecology. In hill mixed dipterocarp forest to lower montane forest, up to 1500 m altitude,
on a variety of soils.
Uses. The wood is used locally as firewood and for making charcoal. The acorns are
edible but rarely eaten.
21. Lithocarpus encleisacarpus (Korth.) A. Camus

(Greek, enkleiein = enclosed, karpos = fruit; with acorn completely enclosed by the cupule)
236; PROSEA l.c. 294. Basionym: Quercus encleisacarpus Korth. Kruidk. (1844) 209. t. 45. Type:
Korthals s.n. (= RHL Sheet No. 901310172), C Sumatra, Mt. Melintang (holotype L; isotypes A, K,
P). Synonyms: Cyclobalanus encleisocarpa (Korth.) Oerst. l.c. (1867) 81; Pasania encleisocarpa
(Korth.) Gamble l.c. (1915) 449; Synaedrys encleisocarpa (Korth.) Koidzumi l.c. 186; Castanopsis
encleisocarpa (Korth.) Rehder l.c. (1919) 122.
Tree up to 40 m tall, 75 cm diameter. Bark smooth or scaly and sparsely lenticellate,

greyish brown; inner bark yellowish. Sapwood whitish. Twigs densely tomentose with
appressed stellate hairs, late-subglabrescent, sparsely lenticellate. Leaves thin-coriaceous,
glabrous above, sparsely appressed yellowish or brownish tomentose below; elliptic or
ovate, (7)1215 (2)46 cm, base symmetrical, acute or obtuse, rarely rounded, margin
not undulate, weakly revolute, apex acute to acuminate, acumen (5)1015 mm long;
56
midrib slightly raised or impressed above, raised below; lateral veins thin, (7)810
(12) pairs, lax, flat or impressed above, thinly raised below, faintly joining near the
leaf margin, forming an angle of 4070 with the midrib; intercostal venation subscalariform
or irregular, dense, obscure above, prominent below; petiole 515 mm long. Inflorescences
male, female or androgynous. Male inflorescences 613 cm long; bracts rounded or broadly
ovate, 0.40.5 0.30.5 mm; bracteoles broadly obovate, c. 0.3 0.5 mm. Male flowers
in clusters of 3 along the rachis; perianth lobes obtuse, c. 1 0.7 mm; stamen filaments
c. 2 mm long; pistillode globose, 11.2 mm diameter. Female or androgynous inflorescences
1020 cm long; bracts rounded, 0.40.5 mm across; bracteoles ovate, c. 0.4 0.2 mm.
Female flowers solitary along the rachis; perianth lobes acute, c. 0.3 0.2 mm; staminodes
1012; styles cylindrical, straight, c. 1.5 mm long. Cupules solitary but arranged closely
together along the rachis, with stalk 1015 mm long, deeply cup-shaped to almost completely
covering the acorn, 1.52.5 23 cm, tomentose, lamellate; wall bony, thin, densely
hairy; lamellae slightly distinct, edge minutely denticulate or entire, set in 57 regular
lines. Acorn depressed ovoid to subglobose, 1.52 1.53 cm, free part densely yellowish
simple hairy or velvety, dark brown; base rounded, apex acute; scar shallowly concave,
11.5 cm diameter; wall bony, thin, most part free from the cupule.
Distribution. Thailand, Sumatra, Peninsular Malaysia, Singapore, and Borneo. In Sabah,
known from Ranau, Tawau, Sandakan, and Beaufort districts. In Sarawak, known mainly
from Kapit, Kuching, Miri, and Limbang districts. Also in Brunei and Kalimantan.
Ecology. In primary hill mixed dipterocarp forest to lower montane forest, at 10001500m altitude.
Uses. The timber is locally used for house construction and as firewood. The bark contains
tannin which is used for dyeing cotton and rattans.
22. Lithocarpus ewyckii (Korth.) Rehder

(D.J. van Ewijck van Oostbroek en De Bilt, 17861858, officer of the Dutch Colonial Service)
J. Arn. Arb. (1929) 132; Soepadmo l.c. (1970) 240, l.c. (1972) 351; Cockburn l.c. (1972) 220, l.c.
(1976) 108; Anderson l.c. 200; Whitmore, Tantra & Sutisna l.c. 156; Turner l.c. 236; PROSEA l.c.
(1995) 294; Coode et al. (eds.) l.c. 125; Kochummen l.c. 248; Argent et al. (eds.) l.c. 248. Basionym:
Quercus ewyckii Korth., Kruidk. (1844) 212. Type: Korthals s.n. (= RHL Sheet No. 901310184),
Central Sumatra, Mt. Melintang (holotype L; isotypes A, BM, NY, P). Synonyms: Cyclobalanus
ewyckii (Korth.) Oerst. l.c. (1867) 80; Pasania ewyckii (Korth.) Gamble l.c. (1915) 431; Synaedrys
ewyckii (Korth.) Koidzumi l.c. (1916) 191; P. lamponga (Miq.) Gamble var. ewyckioides Gamble l.c.
425; L. pseudolamponga A. Camus l.c. (1954) 110, t. 504.
Tree up to 50 m tall, 90 cm diameter. Bark smooth or scaly, greyish brown to reddish

brown; inner bark fibrous, brownish. Sapwood brownish. Twigs subglabrous, sparsely
lenticellate or smooth. Stipules linear, 45 1 mm. Leaves thin-coriaceous, subglabrous
above, sparsely appressed yellowish tomentose with simple hairs below; elliptic or ovate,
(6)9.513(15) (2.5)36(7) cm, base rounded or obtuse, margin not undulate, revolute,
apex bluntly to sharply acuminate, acumen (5)1020 mm long; midrib faintly raised
on both surfaces, stronger below; lateral veins thin, (12)1315(16) pairs, dense, flat
or sometimes obscure above, raised below, disappearing or faintly joining near the leaf
margin, forming an angle of 4560 with the midrib; intercostal venation scalariform,
57
dense, obscure above, prominent below; petiole 515(24) mm long. Inflorescences male,
androgynous or mixed. Male inflorescences 1018 cm long; bracts and bracteoles ovate,
c. 0.5 0.3 mm. Male flowers solitary or in clusters of 23 along the rachis; perianth
lobes acute, c. 1.2 1 mm; stamen filaments 23 mm long; pistillode globose, c. 0.7
mm diameter. Androgynous or mixed inflorescences 510 cm long; bracts and bracteoles
ovate, 11.2 0.50.6 mm. Female flowers solitary along the rachis; perianth lobes
acute, c. 0.7 0.5 mm; staminodes 12; styles conical, slightly recurved, c. 1 mm long.
Cupules solitary along the rachis, with stalk 510 mm long, cup-shaped, 0.41 2
2.5 cm, sparsely tomentose, lamellate; wall woody, thick, sparsely hairy, enclosing less
than half of the acorn; lamellae distinct, edge entire, set in 67 regular lines. Acorns
conical to ovoid, 1.72.3 1.52.5 cm, most part exserted, glabrous, brownish; base
rounded, apex abruptly acute; scar deeply concave, 11.5 cm diameter; wall bony, thick,
Vernacular names. Sarawaksalad repak (Kelabit), taalan (Punan).
Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo. Very common and
widespread throughout Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. In primary hill mixed dipterocarp forest, including kerangas forest, to montane
forest, at 2002000 m altitude, on sandy loam and clay soils.
23. Lithocarpus ferrugineus Soepadmo

(Latin, ferrugineus = red-brown; the indumentum of the acorn)
Tantra & Sutisna l.c. 156; Coode et al. (eds.) l.c. 126; Argent et al. (eds.) l.c. 248. Type: Ilias S.
17030, Borneo, Sarawak, Serian district, Sabal FR (holotype L; isotypes A, BO, K, KEP, M, P, S,
SAN, SAR, SING).
Tree up to 30 m tall, 50 cm diameter; buttresses c. 0.7 m tall. Bark scaly or smooth

or lenticellate, greyish brown; inner bark fibrous, brownish or reddish. Sapwood yellowish.
Twigs sparsely tomentose, finely fissured or sparsely lenticellate. Leaves thin-coriaceous
to coriaceous, glabrous on both surfaces; lanceolate, oblanceolate, elliptic or oblong,
(12)1622.5(30) (3)4.56.5(9) cm, base obtuse or rounded, margin revolute, apex
abruptly acute to acuminate, acumen 1015 mm long; midrib slightly raised above,
strongly raised below; lateral veins thick, (8)913(14) pairs, lax, impressed or flat above,
raised below, faintly joining near the leaf margin, forming an angle of 3050 with the
midrib; intercostal venation subscalariform or irregular, lax, obscure above, prominent
below; petiole (6)715(18) mm long. Inflorescences male, androgynous or mixed. Male
inflorescences 1020 cm long; bracts and bracteoles ovate, 0.51.2 0.30.8 mm. Male
flowers solitary along the rachis; perianth lobes obtuse, c. 1.2 1.5 mm; stamen filaments
c. 2 mm long; pistillode subglobose, 0.81 mm diameter. Androgynous or mixed
inflorescences 1015 cm long; bracts ovate, c. 1.2 1 mm; bracteoles obtuse, 0.20.3
mm across. Female flowers solitary along the rachis; perianth lobes acute, c. 1 0.8
mm; staminodes 12; styles conical, straight, c. 1.5 mm long. Cupules solitary along the
rachis, subsessile, cup- or saucer-shaped, 0.51 1.22 cm, densely appressed tomentose
with stellate hairs, scaly; wall woody, thin, enclosing up to half of the acorn; scale-like
58
appendages distinct, appressed, imbricate, irregularly set. Acorns ovoid, 1.52 1.5
2.3 cm, most part exserted, densely reddish brown tomentose with simple, long and stellate
hairs; base rounded, apex obtuse; scar deeply concave, c. 1 cm diameter; wall woody,
Distribution. Peninsular Malaysia and Borneo. Very common and widespread throughout
Ecology. In primary hill mixed dipterocarp forest to lower montane forest, up to 1200
m altitude, on sandy clay and yellow podsolic soils.
24. Lithocarpus gracilis (Korth.) Soepadmo

(Latin, gracilis = slender; the twig)
Reinwardtia 8 (1970) 243, l.c. (1972) 362; Cockburn l.c. (1972) 220, l.c. (1976) 110; Anderson l.c. 201;
Whitmore, Tantra & Sutisna l.c. 157; Turner l.c. 236; Kessler & Sidiyasa l.c. 141; PROSEA l.c. (1995)
294; Coode et al. (eds.) l.c. 126; Argent et al. (eds.) l.c. 248. Basionym: Quercus gracilis Korth. l.c.
207. Type: Korthals s.n. (= RHL Sheet No 901310314), Borneo, Kalimantan, Balaran (holotype L;
isotypes A, P). Synonyms: Cyclobalanus gracilis (Korth.) Oerst. l.c. (1871) 376; Q. cyrtorhyncha Miq.
l.c. (1861) 350; Pasania cyrtorhyncha (Miq.) Gamble l.c. (1915) 432; Synaedrys cyrtorhyncha (Miq.)
Koidzumi l.c. 191; L. cyrtorhyncha (Miq.) Rehder l.c. (1919) 124; Q. diepenhorstii Miq. l.c. (1861)
349; Cyclobalanus diepenhorstii (Miq.) Oerst. l.c. (1867) 80; S. diepenhorstii (Miq.) Koidzumi l.c. 191;
L. diepenhorstii (Miq.) Barnett l.c. (1944) 177; L. cyathiformis A. Camus l.c. (1947) 4.
Tree up to 40 m tall, 90 cm diameter; buttresses up to 1.5 m tall. Bark smooth or fissured

or scaly, greyish brown; inner bark fibrous, brownish or yellowish. Sapwood yellowish
brown. Twigs sparsely tomentose, late-glabrescent, sparsely lenticellate or smooth. Stipules
linear, 24 1 mm. Leaves thin-coriaceous, glabrous above, sparsely brownish appressed
tomentose below; elliptic, oblong or ovate, (8)12.515.5(18) (4)4.56.5(7) cm, base
obtuse to cordate, margin revolute, often undulate, apex bluntly acute to acuminate, acumen
1015 mm long; midrib faintly raised on both surfaces, stronger below; lateral veins
thin, (12)1317(19) pairs, dense, raised on both surfaces or flat above, faintly joining
near the leaf margin, forming an angle of 3045(60) with the midrib; intercostal venation
subscalariform to irregular, dense, obscure above, prominent below; petiole (3)57
(8) mm long. Inflorescences male, androgynous or mixed. Male inflorescences 10
20 cm long; bracts narrowly ovate to linear, c. 1.2 0.3 mm; bracteoles narrowly ovate,
c. 0.7 0.3 mm. Male flowers in clusters of 35 along the rachis; perianth lobes obtuse,
c. 0.9 0.7 mm; stamen filaments 23 mm long; pistillode subglobose, 0.81 mm diameter.
Androgynous or mixed inflorescences 1220 cm long; bracts linear, c. 1.2 0.3 mm;
bracteoles narrowly ovate, c. 0.4 0.2 mm. Female flowers solitary or in clusters of
3 along the rachis; perianth lobes obtuse, c. 0.6 0.3 mm; staminodes 12; styles cylindrical,
straight, 0.71 mm long. Cupules solitary, rarely in clusters of 3 along the rachis, sessile
or with stalk c. 5 mm long, saucer- to cup-shaped, 0.51 1.82.5 cm, densely tomentose
with simple hairs, lamellate; wall bony, thin, enclosing less than half of the acorn; lamellae
slightly distinct, edge dentate, set in 68 regular lines. Acorns ovoid to conical, 1.5
1.8(2) 1.52.5 cm, most part exserted, glabrous, brown; base rounded, apex abruptly
acute; scar shallowly concave or flat, 0.81.5 cm diameter; wall bony, thin, most part
59
Vernacular names. Sabahdiparak, tikalod (Dusun, Banggi). Sarawaksaled (Kelabit),

tekalat (Kayan), tekalat uak (Kenyah).
Distribution. Sumatra, Peninsular Malaysia and Borneo. Common and widespread throughout
Ecology. In primary or secondary hill mixed dipterocarp forest, including riparian and
kerangas forests, up to 1200 m altitude, on sandy-clay and alluvium soils.
Use. The wood is used locally for general construction.
25. Lithocarpus hallieri (Seemen) A. Camus

(J.G. Hallier, 18681932, sometime research assistant at the Herbarium Bogoriense, Bogor,
and Rijksherbarium, Leiden)
Riviera Scientific 18 (1932) 40; Soepadmo l.c. (1970) 244, l.c. (1972) 327; Cockburn l.c. (1976) 103;
Anderson l.c. 201; Whitmore, Tantra & Sutisna l.c. 157; PROSEA l.c. 295; Argent et al. (eds.) l.c.
248. Basionym: Quercus hallieri Seemen, Bull. Dpt. Agr. Ind. Nerl. 1 (1906) 189. Type: Hallier
2655, Borneo, Kalimantan, Mt. Lianggagang (holotype BO; isotype L). Synonym: Synaedrys hallieri
(Seemen) Koidzumi l.c. 189.

greyish brown; inner bark granular, yellowish or brownish. Sapwood brownish. Twigs
glabrous, smooth or sparsely minute-lenticellate. Leaves thick-coriaceous, glabrous on
both surfaces or glabrous above and sparsely tomentose beneath; elliptic to oblong, 15
20(24) 79(10) cm, base rounded, acute to cuneate, margin revolute, apex acute or
acuminate, acumen c. 10 mm long; midrib raised on both surfaces, stronger below; lateral
veins thin, 611 pairs, lax, flat above, raised below, faintly joining near the leaf margin,
forming an angle of 4560 with the midrib; intercostal venation subscalariform, dense,
obscure above, thinly prominent below, or obscure on both surfaces; petiole 1530 mm
long. Inflorescences male, female and androgynous. Male inflorescences 2026 cm long;
bracts linear, 22.5 0.5 mm; bracteoles ovate, c. 0.8 0.6 mm. Male flowers in clusters
of 3 along the rachis; perianth lobes obtuse, 1.52 1 mm; stamen filaments 1.53.5
mm long; pistillode subglobose, c. 1 mm diameter. Female or androgynous inflorescences
1015 cm long; bracts broadly ovate, c. 1 mm across; bracteoles subrounded, c. 0.5 mm
across. Female flowers solitary along the rachis; perianth lobes acute, c. 1.5 1 mm;
staminodes 12; styles conical, straight, c. 1 mm long. Cupules solitary along the rachis,
sessile or with stalk c. 10 mm long; obovoid to subglobose, 47 cm diameter, sparsely
tomentose, lamellate; wall woody, thick, sparsely hairy, completely enclosing the acorn
except for an apical opening c. 0.5 cm diameter; lamellae obscure or slightly distinct,
edge entire, set in 45 irregular or rarely regular lines. Acorns subglobose, 34.5 4
4.5 cm, free apical part sparsely tomentose, brown; base rounded, apex flat; wall woody,
thick, most part adnate to the cupule.
Vernacular names. Sarawakkeraki (Iban), salad (Kelabit), saled kup (Murut).
Distribution. Endemic to Borneo. In Sabah, very common in Ranau (especially in Kinabalu
60
Park), Kota Belud, Penampang, Tenom, Keningau, and Beaufort districts. In Sarawak,
known from Kapit, Miri and Kuching districts. Also in Kalimantan.
Ecology. In primary lowland mixed dipterocarp forest to montane forest on clay soils,
at 4002600 m altitude.
26. Lithocarpus hatusimae Soepadmo

(S. Hatusima, 19431945 Head of the Herbarium Bogoriense, Bogor, later Professor at
Kagoshima College, Faculty of Agriculture, Kyushu, Japan)
Tantra & Sutisna l.c. 157. Type: Clemens 29206, Borneo, Sabah, Mt. Kinabalu (holotype L; isotypes
A, BM, K, NY, UC).
Tree up to 20 m tall, 35 cm diameter. Bark smooth or fissured, greyish brown; inner

bark fibrous, brownish. Sapwood yellowish or brownish. Twigs densely appressed tomentose,
sparsely or rarely densely lenticellate. Stipules linear, c. 3 1 mm. Leaves thick-coriaceous,
rigid, subglabrous or sparsely appressed tomentose above, sparsely brownish or yellowish
appressed tomentose below; elliptic or ovate, (5.5)8.511.5(18) (2.5)4.56(7.5) cm,
base acute or cuneate, rarely rounded, margin strongly revolute, apex sharply acuminate,
acumen 1020 mm long; midrib broad, strongly raised on both surfaces; lateral veins
thick, (7)1012(13) pairs, dense, impressed or flat above, strongly raised below, faintly
venation subscalariform, dense, obscure on both surfaces or thinly prominent below; petiole
(8)913(20) mm long. Inflorescences male, female, androgynous or mixed. Male
inflorescences 1012 cm long; bracts and bracteoles ovate, c. 0.8 0.4 mm. Male flowers
solitary along the rachis; perianth lobes obtuse, c. 1 0.4 mm; stamen filaments c. 2
mm long; pistillode subglobose, 0.50.8 mm diameter. Female, androgynous or mixed
inflorescences 510 cm long; bracts broadly ovate, 0.20.3 mm across. Female flowers
solitary or in clusters of 23 along the rachis; perianth lobes acute, 0.30.4 mm across;
staminodes 10; styles conical, straight, c. 1.5 mm long. Cupules solitary or rarely in
clusters of 2 along the rachis, with stalk 35 mm long, cup-shaped, 0.50.9 1.21.8
cm, densely appressed tomentose, lamellate; wall woody, thin, enclosing less than half
of the acorn; lamellae distinct, edge minutely denticulate, set in 56 regular lines. Acorns
ovoid to conical, 12 1.22.2 cm, most part exserted, glabrous except for the densely
tomentose small part at the apex, dark brown; base rounded to cordate, apex abruptly
acute; scar concave, c. 1 cm diameter; wall bony, thin, most part free from the cupule.
Vernacular name. Sarawaksalad (Murut).
Distribution. Endemic to Borneo. In Sabah, common in Ranau and Sipitang districts.
In Sarawak, documented from Miri (common in G. Mulu area), Limbang, Sri Aman,
Kapit, and Kuching districts. Also in Brunei and Kalimantan.
Ecology. In primary and secondary hill mixed dipterocarp forest, including kerangas forest,
to montane forest, at 8002200 m altitude, mainly on clay soils.
61
27. Lithocarpus havilandii (Stapf) Barnett

(J.D. Haviland, 18571901, First Sarawak Medical Officer, plant and insect collector)
Trans. & Proc. Bot. Soc. Edinb. 33 (1942) 176; Soepadmo l.c. (1970) 245, l.c. (1972) 373; Cockburn
l.c. (1976) 97; Anderson l.c. 201; Whitmore, Tantra & Sutisna l.c. 157. Basionym: Quercus havilandii
Stapf, FMK (1894) 231, t. 18. Type: Haviland 1069, Borneo, Sabah, Mt. Kinabalu (holotype K;
isotypes BM, SING). Synonyms: Q. abendanonii Valeton, Icon. Bog. 4 (1914) 179, t. 354; L.
abendanonii (Valeton) A. Camus l.c. (1945) 84.
Tree up to 25 m tall, 40 cm diameter. Bark smooth or scaly or lenticellate, greyish brown;

inner bark fibrous, reddish or brownish. Sapwood reddish or whitish. Twigs densely or
sparsely tomentose with stellate and simple hairs, smooth. Stipules ovate, c. 3 1 mm.
Leaves coriaceous to thick-coriaceous, rigid, sparsely stellate-hairy or subglabrous above,
densely brownish stellate-hairy below; elliptic or ovate, (4.5)5.57.5(9.5) (2.5)34
(5) cm, base broadly acute, rounded, or rarely subcordate, margin strongly revolute, apex
acute or acuminate, acumen 510 mm long; midrib raised on both surfaces, stronger below;
lateral veins thick, (6)712 pairs, lax, impressed above, raised below, faintly joining or
disappearing near the leaf margin, forming an angle of 4560 with the midrib; intercostal
venation subscalariform, lax, obscure or prominent above, prominent below; petiole
(3) 58 mm long. Inflorescences male or androgynous. Male inflorescences 36 cm long;
bracts and bracteoles narrowly ovate, 11.2 0.30.5 mm. Male flowers solitary along
the rachis; perianth lobes obtuse, c. 1 0.5 mm; stamen filaments c. 2.5 mm long; pistillode
globose, c. 1 mm diameter. Androgynous inflorescences 36 cm long; bracts narrowly
ovate to linear, 1.23 0.5 mm; bracteoles ovate, c. 0.8 0.5 mm. Female flowers solitary
along the rachis; perianth lobes acute, c. 0.8 0.6 mm; staminodes 12; styles conical,
recurved, c. 2 mm long. Cupules solitary along the rachis, sessile, cup-shaped, 0.30.5
11.5(2) cm, densely appressed tomentose with stellate and simple hairs, scaly; wall
bony, thin, enclosing less than half of the acorn; scale-like appendages distinct, irregularly
set. Acorns ovoid to conical, 1.21.7(2) 1.52 cm, most part exserted, glabrous except
for the sparsely simple-hairy apex, brownish; base flat, apex acute; scar flat or slightly
concave, 0.51 cm diameter; wall bony, thin, most part free from the cupule.
Distribution. Borneo and Sulawesi. In Sabah, common in Ranau and Tambunan districts.
In Sarawak, only known from Sg. Badas Sirau and Ba Kelalan in Lawas district. Also
in Kalimantan.
Ecology. Common on ridge tops in primary lower and upper montane forests, at 1300
2500 m altitude.
28. Lithocarpus hystrix (Korth.) Rehder

(Latinised Greek, hystrix = set with slender prickles; the cupule)
J. Arn. Arb. 1 (1919) 127; Soepadmo l.c. (1970) 246, l.c. (1972) 384; Corner l.c. 340; Whitmore,
Tantra & Sutisna l.c. 157; Turner l.c. 236; PROSEA l.c. 295; Coode et al. (eds.) l.c. 126. Basionym:
Quercus hystrix Korth. l.c. 201, t. 43. Type: Korthals s.n., Sumatra (holotype L; isotypes A, K, P).
Synonyms: Q. korthalsii Blume var. hystrix (Korth.) Blume, Mus. Bot. Lugd.-Bat. 1 (1850) 293; Q.
mappacea Korth. var. hystrix (Korth.) Miq. l.c. (1856) 869; Cyclobalanus hystrix (Korth.) Oerst. l.c.
(1867) 81; Pasania hystrix (Korth.) Gamble l.c. (1915) 428; Synaedrys hystrix (Korth.) Koidzumi l.c.
62
(1916) 195; Q. cyrtopoda Miq. l.c. (1858) 869; Cyclobalanus cyrtopoda (Miq.) Oerst. l.c. (1867) 80;
L. cyrtopoda (Miq.) A. Camus l.c. (1949) 74; Castanea furfurella Miq. l.c. (1861) 352; Q. brevipetiolata
Scheff. l.c. (1870) 359.
Tree up to 36 m tall, 75 cm diameter. Bark scaly or lenticellate, brownish or reddish.

Twigs densely stellate-hairy, smooth or sparsely minute-lenticellate. Stipules linear, c.
3 1 mm. Leaves thin-coriaceous, densely tomentose with stellate and simple hairs above,
densely or rarely sparsely appressed tomentose below; broadly lanceolate or obovate,
(8)10.513.5(16) (2.5)35.5(6) cm, base rounded, margin revolute, apex acute to
acuminate, acumen 515 mm long; midrib raised on both surfaces, stronger below; lateral
veins thin, 1012(15) pairs, dense, slightly raised or flat above, raised below, disappearing
or faintly joining near the leaf margin, forming an angle of 4060 with the midrib;
intercostal venation scalariform, dense, obscure above, slightly prominent below; petiole
35 mm long. Inflorescences male, androgynous or mixed. Male inflorescences 511
cm long; bracts and bracteoles narrowly ovate to linear, 0.51.2 0.20.5 mm. Male
flowers in clusters of 23 along the rachis; perianth lobes obtuse, c. 1 0.8 mm; stamen
filaments c. 2 mm long; pistillode globose, 11.2 mm diameter. Androgynous or mixed
inflorescences 717 cm long; bracts narrowly ovate, c. 1 0.30.5 mm; bracteoles broadly
ovate, c. 0.7 0.5 mm. Female flowers solitary along the rachis; perianth lobes acute,
c. 0.7 0.5 mm; staminodes 1012; styles conical, recurved, c. 1 mm long. Cupules
solitary along the rachis, sessile, saucer-shaped, 0.71 1.82.5 cm, densely tomentose;
wall woody, thick, enclosing less than half of the acorn, irregularly set with pricklelike, slender appendages. Acorns ovoid to conical, 1.82 1.72 cm, most part exserted,
glabrous, purplish; base rounded, apex acute; scar deeply concave, c. 1.3 cm diameter;
Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo. So far not known
in Sabah. In Sarawak, recorded for Miri, Kapit, Kuching, and Sri Aman districts. Also
Ecology. In lowland mixed dipterocarp forest to lower montane forest, at 3001300 m
altitude, on yellow sandy-clay soils.
Use. The bark contains tannin.
29. Lithocarpus jacobsii Soepadmo

(M. Jacobs, 19291983, former botanist at the Rijksherbarium, Leiden)
Coode et al. (eds.) l.c. 126; Argent et al. (eds.) l.c. 249. Type: Jacobs 5398, Borneo, Sarawak, Belaga
district, Batang Rajang (holotype L; isotypes G, K, SAR, US).
Tree up to 36 m tall, 60 cm diameter. Bark smooth or lenticellate, brown; inner bark

brownish. Sapwood whitish. Twigs sparsely tomentose with stellate hairs, smooth or
sparsely large-lenticellate. Stipules linear, 510 1 mm. Leaves thick-coriaceous, glabrous
on both surfaces; oblong or obovate, (17)2440(56) (4.5)610.5(16) cm, base cordate
to auriculate, margin very strongly revolute, apex bluntly to sharply acute to acuminate,
acumen c. 10 mm long; midrib broad, strongly raised on both surfaces; lateral veins
63
thick, (10)1317(19) pairs, lax, flat or impressed above, strongly raised below, clearly
venation subscalariform to irregular, lax, obscure above, prominent below; petiole (4)5
12(15) mm long. Inflorescences male and female. Male inflorescences 610 cm long;
bracts broadly ovate, c. 1 0.8 mm; bracteoles narrowly ovate, c. 0.7 0.4 mm. Male
flowers in clusters of 3 along the rachis; perianth lobes acute, 46 23 mm; stamen
filaments c. 5 mm long; pistillode c. 5 mm diameter. Female inflorescences c. 25 cm
long; bracts broadly ovate, c. 1 0.7 mm; bracteoles elliptic, c. 0.8 0.6 mm. Female
flowers in clusters of 3 along the rachis; perianth lobes acute, 45 mm across; staminodes
12; styles conical, straight, 23 mm long. Cupules in clusters of 23 along the rachis
and arranged closely together, with stalk c. 5 mm long, saucer-shaped, 0.20.7 22.5
cm, densely appressed tomentose with stellate hairs, scaly; wall woody, thick, enclosing
less than half of the acorn; scale-like appendages slightly distinct, set in regular lines.
Acorns depressed ovoid to subglobose, 1.52.5 22.8 cm, most part exserted, glabrous
except for the sparsely simple-hairy apex, dark brown; base flat, apex rounded; scar deeply
Vernacular names. Bruneitakalot (Dusun), tusuk pending memaluh (Iban). Sarawak
kalit (Iban, Engkari), saled arong (Kelabit).
Distribution. Peninsular Malaysia and Borneo. In Sabah, only known from Sandakan
district (Sepilok FR, Sg. Kun-Kun and Telupid). In Sarawak, known mainly from Kapit,
Miri and Sri Aman districts. Also in Brunei and Kalimantan.
Ecology. In primary and secondary hill mixed dipterocarp forest to lower montane forest,
up to 1300 m altitude, on sandy clay soils.
30. Lithocarpus kalkmanii S. Julia & Soepadmo

(C. Kalkman, 19281998; former Director of the Rijksherbarium, Leiden)
Gard. Bull. Sing. 50 (1998) 130. Type: Meijer & Hendry SAN 42460, Borneo, Sabah, Ranau district,
Kinabalu Park, western border north of Sosopodon (holotype SAN; isotypes AA, K, L).
Tree up to 30 m tall, 60 cm in diameter. Bark large-lenticellate or cracked, brownish

or greyish; inner bark reddish brown. Sapwood yellowish. Twigs sparsely brownish
tomentose, late-glabrescent, smooth or sparsely large-lenticellate. Leaves thick-coriaceous,
sometimes rigid, densely appressed yellowish tomentose above, densely appressed greyish
brown tomentose below; broadly elliptic, 1115.5 58 cm, base acute, margin revolute,
apex acuminate, acumen 512 mm long; midrib slightly raised above, strongly raised
below; lateral veins thick, 1214 pairs, dense, flat above, strongly raised below, disappearing
scalariform, dense, obscure above, slightly prominent below; petiole 1520 mm long.
Inflorescences and flowers unknown. Cupules solitary along the rachis, sessile, subglobose,
57 45.5 cm, base rounded, apex flat, glabrous, smooth, lamellate; wall woody, thick,
completely enclosing the acorn or with an apical opening 0.51 cm diameter; lamellae
distinct, edge entire or minutely denticulate, especially the upper-most ones, set in 6
7 regular lines. Acorns subglobose, 4.56 35 cm, free apical part depressed at the
centre, densely tomentose, dark brown; wall woody, thick, most part adnate to the cupule.
64
TREE FLORA OF SABOL. 3
vi
Distribution. Endemic to Sabah. So far only known in Kinabalu Park and Sosopodon
FR in Ranau district, and one collection from Nabawan in Keningau district.
Ecology. In primary hill mixed dipterocarp forest to lower montane forest, at 10001500
m altitude.
31. Lithocarpus keningauensis S. Julia & Soepadmo
Fig. 10.
(of Keningau, Sabah)

Gard. Bull. Sing. 50 (1998) 132. Type: Amin SAN 95311, Borneo, Sabah, Keningau district, Bt. Kitau
(holotype SAN; isotypes K, KEP, L, SAR).
Tree up to 30 m tall, 60 cm in diameter. Bark fissured to scaly, brown or reddish; inner

bark brown or reddish. Sapwood yellowish to brownish. Twigs densely appressed tomentose,
late-glabrescent, slightly fissured or sparsely large-lenticellate or smooth. Leaves thickcoriaceous, rigid, glabrous or densely appressed yellowish tomentose above, densely
appressed yellowish or brownish tomentose below; elliptic to broadly elliptic, 12.516.5
(29) 4.57(12.5) cm, base acute to cuneate, margin revolute, apex shortly acuminate,
acumen c. 10 mm long; midrib broad, slightly raised above, strongly raised below; lateral
veins thick, 1214 pairs, lax, slightly raised or flat above, raised below, disappearing
scalariform, dense, obscure above, slightly prominent below; petiole 1015 mm long.
Inflorescences and flowers unknown. Cupules solitary along the rachis, sessile, obovoid,
711 56.5 cm, base attenuate, apex flat and umbonate, glabrescent, strongly lamellate;
wall woody, thick, completely enclosing the acorn; lamellae distinct and prominent even
in the young cupule, folded inwardly, longitudinally grooved, set in 610 regular lines,
raised from the cupule surface. Acorns obovoid, 4.56 35 cm, free apical part depressed
and umbonate, densely tomentose, dark brown; wall woody, thick, most part adnate to
the cupule.
Distribution. Endemic to Sabah. Known only from Ulu Biah, Bt. Kitau and Keningau
trail in Keningau district.
Ecology. In primary and secondary forests on hill slopes, up to 300 m, on dark brown
ultrabasic soils.
32. Lithocarpus kochummenii S. Julia & Soepadmo
Fig. 11.
(K.M. Kochummen, 19311999, former senior forest botanist at the Forest Research Institute Malaysia, Kepong)
Gard. Bull. Sing 50 (1998) 134. Type: Lee S. 38884, Borneo, Sarawak, Miri district, G. Mulu NP
(holotype SAR; isotypes K, KEP, SAN).
Tree up to 30 m tall, 60 cm in diameter; stilt-root up to 2 m high. Bark fissured or

lenticellate, reddish brown. Twigs densely tomentose, late-glabrescent, sparsely to densely
large-lenticellate. Stipules linear, 610 1 mm. Leaves thick-coriaceous, rigid, sparsely
66
vi
appressed tomentose or glabrescent above, densely appressed brownish or yellowish

tomentose below; narrowly elliptic, lanceolate or oblanceolate, (10)1418(22) (4)
57(8.5) cm, base acute, obtuse to rounded, margin strongly revolute, apex acute to
acuminate, acumen 1520 mm long; midrib broad, raised on both surfaces, stronger below;
lateral veins thick, (11)1418(20) pairs, dense, strongly impressed or rarely flat above,
strongly raised below, clearly joining near the leaf margin, forming an angle of 40
60 with the midrib; intercostal venation thin, scalariform, rarely subscalariform, lax,
obscure above, thinly prominent below; petiole 510 mm long. Inflorescences male, female
or androgynous. Male inflorescences 513 cm long; bracts elliptic, 1.51.7 0.8 mm;
bracteoles oblong, c. 0.5 0.2 mm. Male flowers solitary along the rachis; perianth
lobes acute, c. 1.2 1 mm; stamen filaments 22.5 mm long; pistillode c. 1 mm in
diameter. Female or androgynous inflorescences 715 cm long; bracts and bracteoles
broadly ovate, 11.2 0.61 mm. Female flowers solitary along the rachis; perianth
lobes acute, c. 1 0.8 mm; staminodes 1012; styles conical, recurved, c. 2 mm long.
Cupules solitary along the rachis, sessile, conical to ovoid, 22.5 cm across, densely
tomentose with simple and stellate hairs; wall woody, thin, completely enclosing the acorn;
spine-like appendages distinct, broad and sturdy, straight or slightly recurved, set irregularly. Acorns conical, 1.52 cm across, free part densely tomentose with simple hairs,
rarely glabrous, brown; base flat, apex acute; scar deeply concave, 1.31.5 cm in diameter;
wall woody, thin, most part free from the cupule.
Vernacular name. Sarawaksalad (Murut).
Distribution. Endemic to Borneo. So far not known in Sabah. In Sarawak, recorded for
G. Mulu in Miri district, Ulu Sg. Kayan in Lundu district, Batu Lawi and Kota FR in
Limbang district. Also in Brunei.
Ecology. In kerangas, riparian, to lower montane forests, at 9001300 m altitude.
33. Lithocarpus lampadarius (Gamble) A. Camus

(Greek, lampas = torch; twigs are apparently used for torches in fishing by the Sakai tribe in
Peninsular Malaysia)
237; PROSEA l.c. 297. Basionym: Pasania lampadaria Gamble, Kew Bull. (1914) 177. Type: Wray
1174, Peninsular Malaysia, Perak, Mt. Batu Putih (holotype K; isotype SING). Synonyms: Syanedrys
lampadaria (Gamble) Koidzumi l.c. 192; Quercus lampadaria (Gamble) Burkill, Kew Bull. (1935)
318.
Tree up to 36 m tall, 100 cm diameter. Bark scaly or fissured, brown; inner bark fibrous,
brownish. Sapwood brownish or yellowish. Twigs sparsely appressed tomentose, sparsely
to densely large-lenticellate. Stipules linear, 615 13 mm. Leaves thick-coriaceous,
rigid, glabrous above, sparsely brownish appressed tomentose below; broadly ovate, elliptic,
or obovate, (13)1527(38) (5)814(23) cm, base acute to cuneate, margin revolute,
apex acute to acuminate, acumen 1525 mm long; midrib broad, strongly raised on both
surfaces; lateral veins thick, (6)1015(20) pairs, dense or lax, impressed or flat above,
68
raised below, disappearing near the leaf margin, forming an angle of 4560 with the
midrib; intercostal venation usually scalariform, dense, obscure above, prominent below;
petiole 1027 mm long. Inflorescences male or androgynous. Male inflorescences 6
20 cm long; bracts ovate, c. 1 0.7 mm; bracteoles linear, 0.80.9 0.2 mm. Male
flowers solitary along the rachis; perianth lobes obtuse, c. 1.6 1.3 mm; stamen filaments
c. 2 mm long; pistillode globose, c. 1 mm diameter. Androgynous inflorescences 10
18 cm long; bracts narrowly ovate, c. 3 1.5 mm; bracteoles broadly ovate, c. 2.8
2.5 mm. Female flowers in clusters of 35 along the rachis; perianth lobes acute, c.
1 0.5 mm; staminodes 10; styles conical, straight, c. 1 mm long. Cupules in clusters
of 25 or rarely solitary along the rachis, sessile, cup-shaped, 12 24 cm, densely
appressed tomentose, scaly; wall woody, very thick, enclosing up to half of the acorn;
scale-like appendages slightly distinct, set in regular lines. Acorns depressed ovoid to
subglobose, 22.5 33.5 cm, most part exserted, glabrous, dark brown; base flat, apex
rounded; scar deeply concave, 12 cm diameter; wall woody, thick, most part free from
the cupule.
Vernacular name. Sarawaksalad kup (Murut).
Distribution. Peninsular Malaysia and Borneo. In Sabah known from Ranau, Tambunan
and Tenom districts. In Sarawak only known from Sg. Buong in Belaga district and Kota
FR in Limbang district. Also in Kalimantan.
Ecology. In primary and secondary hill mixed dipterocarp forest to lower montane forest,
at 3001800 m altitude, on sandstone-derived soils.
34. Lithocarpus leptogyne (Korth.) Soepadmo

(Greek, leptogyne = with slender female organ; the slender young female flower and acorn)
Reinwardtia 8 (1970) 254, l.c. (1972) 379; Cockburn l.c. (1972) 223, l.c. (1976) 110; Anderson l.c. 201;
Whitmore, Tantra & Sutisna l.c. 157; Kessler & Sidiyasa l.c. 141; PROSEA l.c. 298; Coode et al. (eds.) l.c.
126; Argent et al. (eds.) l.c. 249. Basionym: Quercus leptogyne Korth., Kruidk. (1844) 206. Type: Korthals
s.n. (= RHL Sheet No 90131137), Borneo, Kalimantan, Mt. Sakumbang (holotype L; isotypes A, P). Synonyms:
Cyclobalanus leptogyne (Korth.) Oerst. l.c. (1871) 376; Q. lamponga (non Miq.) Hook. f., Fl. Brit. Ind. 5
(1888) 611; Q. conocarpa (non Oudem.) Merr. l.c. (1929) 43.
Tree up to 40 m tall, 90 cm diameter; buttresses up to 1.5 m tall. Bark smooth or lenticellate,

greyish; inner bark granular, brownish. Sapwood yellowish. Twigs densely tomentose
with stellate hairs, sparsely minute-lenticellate. Stipules linear, 36 1 mm. Leaves
coriaceous, sparsely yellowish tomentose with stellate and simple hairs on both surfaces;
elliptic, lanceolate, or elliptic-oblong, (7.5)1216(19) (2.5)3.54.5(6.5) cm, base
acute, rarely rounded, margin revolute, apex acuminate, acumen 1015(20) mm long;
midrib thin, raised on both surfaces; lateral veins thick, 1114(18) pairs, dense, impressed
or flat above, raised below, clearly joining near the leaf margin, forming an angle 30
60 with the midrib; intercostal venation subscalariform, lax, obscure above, prominent
below; petiole 68(10) mm long. Inflorescences male, androgynous or mixed. Male
inflorescences 1020 cm long; bracts ovate, c. 0.7 0.5 mm; bracteoles broadly ovate,
c. 0.5 mm across. Male flowers solitary along the rachis; perianth lobes obtuse, c. 0.7
69
0.5 mm; stamen filaments 23 mm long; pistillode subglobose, c. 1 mm diameter.

Androgynous or mixed inflorescences 6.510 cm long; bracts and bracteoles ovate, c.
1 0.5 mm. Female flowers solitary along the rachis; perianth lobes acute, 0.50.7
mm across; staminodes 12; styles conical, slightly recurved, c. 1 mm long. Cupules solitary
along the rachis, with stalk c. 5 mm long, saucer-shaped, 0.30.8 1.32 cm, densely
tomentose, scaly; wall woody, thick, densely tomentose, enclosing less than half of the
acorn; scale-like appendages distinct, appressed, dense, regularly set. Acorns conical, 1.2
2 1.52 cm, most part exserted, densely simple-hairy, brownish or purplish; base rounded,
apex acute; scar deeply concave or flat, c. 1.2 cm diameter; wall woody, thin, most part
Vernacular names. Sabahpening-peningan (Kedayan), tikalod (Dusun, Bundu Tuhan),
turug (Sungai). Sarawakkalan (Berawan).
Distribution. Peninsular Malaysia and Borneo. Very common and widespread throughout
on various types of soils.
35. Lithocarpus lucidus (Roxb.) Rehder
Fig. 12.
(Latin, lucidus = shining; the acorn and upper leaf surface)

PROSEA l.c. 298; Coode et al. (eds.) l.c. 126; Argent et al. (eds.) l.c. 249; Kochummen l.c. 248.
Basionym: Quercus lucida Roxb., Fl. Ind. 3 (1832) 635. Type: Roxburgh s.n., Peninsular Malaysia,
Penang (holotype BR; isotype BM). Synonyms: Q. cuneata A. DC., Prod. 16, 2 (1864) 108; Pasania
lucida (Roxb.) Gamble l.c. (1915) 440; Synaedrys lucida (Roxb.) Koidzumi l.c. 192; Q. omalokos
Korth. l.c. 214; Cyclobalanus omalokos (Korth.) Oerst. l.c. (1867) 80; P. omalokos (Korth.) Schottky
l.c. (1912) 676; S. omalokos (Korth.) Koidzumi l.c. 192; L. omalokos (Korth.) Rehder l.c. (1919) 129.
Tree up to 50 m tall, 120 cm diameter; buttresses up to 1 m tall. Bark smooth, brown;

inner bark granular, yellowish brown. Sapwood yellowish to brownish. Twigs glabrescent,
sparsely lenticellate. Stipules linear, 23 1 mm. Leaves coriaceous, sometimes rigid,
glabrous on both surfaces, shining above; oblanceolate, (4)812(16) (2)46(8)
cm, base cuneate, decurrent, margin revolute sometimes undulate, apex rounded, obtuse,
acute to abruptly cuspidate; midrib faintly raised on both surfaces; lateral veins thin,
(10)1416(20) pairs, dense, flat on both surfaces or slightly raised below, faintly joining
reticulate, dense, thin but distinct on both surfaces; petiole 27 mm long. Inflorescences
male, female or androgynous. Male inflorescences 920 cm long; bracts and bracteoles
ovate, 0.61 0.30.6 mm. Male flowers in clusters of 34 along the rachis; perianth
lobes obtuse, c. 1 0.8 mm; stamen filaments c. 2 mm long; pistillode globose, c. 0.8
mm diameter. Female or androgynous inflorescences 1020 cm long; bracts narrowly
ovate, c. 1.4 0.30.6 mm; bracteoles broadly ovate, c. 1 0.9 mm. Female flowers
solitary or in clusters of 3 along the rachis; perianth lobes acute, c. 1 0.7 mm; staminodes
70
vi
10; styles conical, straight, c. 1.2 mm long. Cupules solitary but arranged closely together
or sometimes in clusters of 3 along the rachis, sessile or with stalk c. 5 mm long, cupor saucer-shaped, 0.51(1.5) 23.5 cm, glabrescent, lamellate; wall woody, thick,
glabrescent, enclosing half or less of the acorn; lamellae dense and distinct, edge minutely
denticulate or undulate, set in 811 regular lines. Acorns depressed ovoid, 1.52.5
23.5 cm, most part exserted, glabrous and shining, brown purplish; base flat, apex rounded
or acute; scar deeply concave 1.52 cm diameter; wall woody, thin, most part free from
the cupule.
Vernacular names. Sabahsarusup tembaung (Dusun, Kimanis). Sarawaksaled arong
(Kelabit).
Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo. Common and
widespread throughout Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. In primary hill mixed dipterocarp forest to lower montane forest, up to 1400 m
altitude, on clay, sandy-clay and sandstone-derived soils.
Use. The wood is locally used for house construction.
36. Lithocarpus luteus Soepadmo

(Latin, luteus = golden yellow; the indumentum of the acorn)
Tantra & Sutisna l.c. 158; PROSEA l.c. 298; Argent et al. (eds.) l.c. 249. Type: Clemens 32817, Borneo,
Sabah, Mt. Kinabalu, Mt. Nuntok ridges (holotype L; isotypes A, BM, K, NY, UC).
Tree up to 40 m tall, 100 cm diameter; buttresses c. 2 m tall. Bark deeply fissured to scaly,
lenticellate, reddish brown; inner bark fibrous, reddish or brownish. Sapwood greyish.
Twigs glabrescent, densely warty lenticellate. Stipules linear, 36 1 mm. Leaves thickcoriaceous, glabrous above, sparsely appressed tomentose below; elliptic, (7)913(27)
(2.5)46.5(12) cm, base acute to cuneate, margin revolute, apex acute to acuminate,
acumen 1015 mm long; midrib flat above, broad and strongly raised below; lateral veins
thin, 912(14) pairs, dense, flat above, raised below, disappearing near the leaf margin,
forming an angle of c. 45 with the midrib; intercostal venation scalariform, very dense,
obscure above, thin but distinct below; petiole 815(20) mm long. Inflorescences male
or androgynous. Male inflorescences 915 cm long; bracts ovate, 3 1.82 mm; bracteoles
elliptic, c. 1 0.3 mm. Male flowers solitary or in clusters of 3 along the rachis; perianth
lobes obtuse, 11.2 0.6 mm; stamen filaments 23 mm long; pistillode subglobose, 1
1.2 mm diameter. Androgynous inflorescences 513 cm long; bracts and bracteoles narrowly
ovate, 12 0.31 mm. Female flowers solitary or in clusters of 23 along the rachis;
perianth lobes obtuse, 0.81 0.61 mm; staminodes 1012; styles conical, straight, c. 1
mm long. Cupules in clusters of 23 or solitary along the rachis, sessile or with stalk c.
5 mm long, cup-shaped, 11.5 23 cm, tomentose, lamellate; wall woody, thick, hairy,
enclosing up to half of the acorn; lamellae obscure or slightly distinct, edge denticulate,
set in 810 regular lines. Acorns ovoid to subglobose, 12 22.5 cm, most part exserted,
densely covered with golden yellow hairs, brown; base flat, apex acute; scar flat or shallowly
concave, c. 2 cm diameter; wall woody, thick, most part free from the cupule.
72
Distribution. Endemic to Borneo. In Sabah, very common in Ranau district, and also
recorded for Tambunan, Tenom and Sandakan districts. In Sarawak, known mainly in
Kapit, Miri (G. Mulu and Lambir NP) and Lawas districts. Also in Kalimantan.
Ecology. In primary hill mixed dipterocarp forest to lower montane forest, at 5001800 m
altitude.
37. Lithocarpus mariae Soepadmo

(Mrs. M.J. van Steenis-Kruseman, 19041999, the highly respected bibliographer and biographer of Malesian botany)
Tantra & Sutisna l.c. 158; Argent et al. (eds.) l.c. 249. Type: Anderson et al. S. 15404, Borneo, Sarawak,
Lundu district, Mt. Lundu (holotype L; isotypes A, BK, K, M, S, SAR, SING).
Tree up to 35 m tall, 60 cm diameter; buttresses c. 0.7 m tall. Bark smooth, lenticellate,

greyish; inner bark granular, red-brown. Sapwood yellowish. Twigs glabrescent, smooth
or sparsely lenticellate. Leaves thin-coriaceous, glabrous above, sparsely appressed yellowish
tomentose below; ovate or elliptic, (8)1012(13) 45(9) cm, base rounded, symmetrical,
margin revolute, not undulate, apex acute to acuminate, acumen 1015 mm long; midrib
flat above, raised below; lateral veins thin, (9)1012 pairs, lax, flat or impressed above,
raised below, faintly joining or disappearing near the leaf margin, forming an angle of
4560 with the midrib; intercostal venation scalariform, dense, obscure on both surfaces
or thinly prominent below; petiole 1525 mm long. Inflorescences male or androgynous.
Male inflorescences 1015 cm long; bracts ovate, c. 0.5 0.3 mm; bracteoles linear,
c. 0.5 0.1 mm. Male flowers solitary along the rachis; perianth lobes obtuse, c. 0.5
mm across; stamen filaments 2.53 mm long; pistillode globose, c. 0.5 mm diameter.
Androgynous inflorescences 810 cm long; bracts triangular, c. 0.7 0.5 mm; bracteoles
linear, c. 0.7 0.2 mm. Female flowers solitary along the rachis; perianth lobes rounded,
c. 0.5 mm across; staminodes 12; styles triangular, straight, c. 1 mm long. Cupules solitary
along the rachis, with stalk 1020 mm long, ovoid, 24 2.53 cm, densely tomentose,
smooth, often with longitidunal streaks; wall bony, thin, completely enclosing the acorn.
Acorns ovoid, 23.5 22.5 cm, free part densely tomentose with simple hairs, dark
brown; base rounded, apex acute; scar deeply concave, c. 2 cm diameter; wall woody,
Distribution. Endemic to Borneo; uncommon. In Sabah, documented from Tawau, Beaufort,
Kota Belud, and Kudat districts. In Sarawak, known from Kuching, Kapit and Limbang
districts. Also in Kalimantan.
Ecology. In primary hill mixed dipterocarp forest, on flat lands and hilly sides, up to 800 m
altitude.
38. Lithocarpus meijeri Soepadmo

(W. Meijer, 1923; 19591968 forest botanist at Sandakan, Sabah, and later Professor of
Biology, University of Kentucky, Lexington, U.S.A.)
73
Tantra & Sutisna l.c. 158; PROSEA l.c. 299; Coode et al. (eds.) l.c. 126; Argent et al. (eds.) l.c. 251.
Type: Meijer SAN 39608, Borneo, Sandakan district, Sepilok FR (holotype L; isotypes K, SAN,
SING).
Tree up to 40 m tall, 100 cm diameter. Bark smooth or fissured or scaly, greyish to

reddish brown; inner bark brownish. Sapwood yellowish brown. Twigs sparsely tomentose
with simple hairs, sparsely to densely lenticellate. Stipules linear, 34 mm long. Leaves
coriaceous, glabrous above, sparsely yellowish appressed tomentose below; elliptic, obovate,
or broadly elliptic, (11)1316(20) (4)57(8.5) cm, base cuneate, decurrent, margin
revolute, apex acute to acuminate, acumen 510 mm long, rarely rounded; midrib flat
above, raised below; lateral veins thin, (9)1113 pairs, dense, flat above, raised below,
disappearing near the leaf margin, forming an angle of c. 45 with the midrib; intercostal
venation scalariform, dense, prominent on both surfaces; petiole 1015 mm long.
Inflorescences male, female, androgynous or mixed. Male inflorescences 714 cm long;
bracts linear, c. 1 0.2 mm; bracteoles broadly ovate, c. 0.5 0.4 mm. Male flowers
solitary or in clusters of 23 along the rachis; perianth lobes obtuse, c. 1.3 0.6 mm;
stamen filaments c. 2.5 mm long; pistillode subglobose, 0.81 mm diameter. Female,
androgynous or mixed inflorescences 815 cm long; bracts and bracteoles narrowly ovate,
0.81.2 0.5 mm. Female flowers solitary or in clusters of 23 along the rachis; perianth
lobes acute, c. 0.8 0.5 mm; staminodes 12; styles conical, straight, c. 1.5 mm long.
Cupules solitary or in clusters of 23 along the rachis, sessile, cup-shaped, 11.8 2
2.5 cm, densely appressed tomentose, lamellate; wall woody, thick, enclosing less than
half of the acorn; lamellae distinct, edge entire or undulate, sometimes obscure, set in
67 regular lines. Acorns depressed ovoid, 0.52 1.73 cm, most part exserted, densely
silvery grey tomentose with simple hairs, brown; base flat, apex acute; scar deeply concave,
1.52 cm diameter; wall woody, thick, most part free from the cupule.
Distribution. Peninsular Malaysia and Borneo. Common throughout Sabah. In Sarawak,
known from Kapit, Miri and Kuching districts. Also in Brunei and Kalimantan.
Ecology. In primary hill mixed dipterocarp forest, including kerangas forest, usually on
steep hilly sides, up to 1100 m altitude.
39. Lithocarpus melataiensis S. Julia & Soepadmo

(of Bt. Melatai, Sarawak)
Gard. Bull. Sing. 50 (1998) 137. Type: Yii S. 48455, Borneo, Sarawak, Kapit district, Bt. Melatai
(holotype SAR; isotypes BKF, K, KEP, KLU, L).
Tree up to 30 m tall, 70 cm in diameter. Twigs sparsely tomentose, smooth or fissured.

Stipules linear, c. 10 1 mm. Leaves thin-coriaceous, sparsely appressed brownish
tomentose with simple hairs on both surfaces; narrowly elliptic, 913.5 23 cm, base
acute or cuneate, margin revolute, apex caudate or long-acuminate, acumen 1522 mm
long; midrib slightly raised on both surfaces; lateral veins thin, (8) 912 pairs, dense,
flat above, raised below, faintly joining near the leaf margin, forming an angle of 40
50 with the midrib; intercostal venation subscalariform, lax, obscure on both surfaces
74
or thinly prominent below; petiole 612 mm long. Inflorescences male and female. Male
inflorescences 511 cm long; bracts linear, triangular, 11.3 0.3 mm; bracteoles ovate,
0.40.6 0.30.5 mm. Male flowers solitary along the rachis; perianth lobes rounded,
11.2 0.51 mm; stamen filaments 22.5 mm long; pistillode globose, c. 1 mm in
diameter. Female inflorescences 9.512.5 cm long; bracts linear, triangular, c. 1.2 0.3
0.5 mm; bracteoles obtuse, 0.20.3 mm across. Female flowers solitary along the rachis;
perianth lobes acute, 0.40.6 0.20.3 mm; staminodes 10; styles conical, straight, c.
1 mm long. Cupules solitary along the rachis, sessile, deeply saucer-shaped, 0.40.7
1.21.7 cm, densely tomentose with stellate and simple hairs, lamellate; wall bony, thin,
enclosing less than half of the acorn; lamellae distinct, edge minutely denticulate, set
in 68 regular lines. Acorns conical, 1.72 11.5 cm, most part exserted, densely simplehairy, brown; base flat, apex acute; scar concave, 0.71 cm in diameter; wall bony, thin,
Distribution. Endemic to Sarawak. So far only known from Bt. Melatai and Ulu Batang
Baleh in Kapit district.
Ecology. In hill mixed dipterocarp forest, at 300900 m altitude.
40. Lithocarpus muluensis S. Julia & Soepadmo

(of G. Mulu, Sarawak)
Gard. Bull. Sing. 50 (1998) 139. Type: Ilias S. 15082, Borneo, Sarawak, G. Mulu NP (holotype SAR).
Tree up to 30 m tall, 75 cm in diameter. Bark flaky. Twigs glabrescent, fissured or sparsely

lenticellate. Leaves thick-coriaceous, rigid, glabrous above, sparsely appressed yellowish
tomentose below; elliptic to broadly elliptic, (8)1117 (4.5)5.58.5(9) cm, base acute,
margin strongly revolute, apex acute to acuminate, acumen (5)1323 mm long; midrib
raised on both surfaces, stronger above; lateral veins thin, 915 pairs, dense or lax, slightly
raised on both surfaces, faintly joining near the leaf margin, forming an angle of 4050
with the midrib; intercostal venation subscalariform, lax, obscure on both surfaces; petiole
(8)1013(15) mm long. Inflorescences and flowers unknown. Cupules in clusters
of 2 along the rachis when young, later solitary along the rachis, sessile, saucer-shaped,
1.82.4 22.5 cm, sparsely stellate-tomentose, scaly; wall woody, thick, enclosing less
than half of the acorn; scale-like appendages distinct, broad and rigid, set irregularly. Acorns
depressed subglobose, 1.82.4 22.5 cm, most part exserted, glabrous and shining, dark
brown; base flat, apex rounded; scar deeply concave, 1.51.7 cm in diameter; wall woody,
thick, most part free from the cupule.
Distribution. Endemic to Sarawak. So far only known from G. Mulu NP in Miri district.
m altitude.
41. Lithocarpus nieuwenhuisii (Seemen) A. Camus

(A.W. Nieuwenhuis, 18641900, surgeon and amateur botanist)
75
114; Anderson l.c. 202; Whitmore, Tantra & Sutisna l.c. 158; PROSEA l.c. 299; Coode et al. (eds.) l.c.
126; Argent et al. (eds.) l.c. 251. Basionym: Quercus nieuwenhuisii Seemen, Bull. Dpt. Agr. Ind.
Nerl. 1 (1906) 6. Type: Jaheri 1546, Borneo, Kalimantan, Sg. Salut (holotype BO). Synonyms:
Synaedrys nieuwenhuisii (Seemen) Koidzumi l.c. 197; Q. clementis Merr. l.c. (1908) 321; Pasania
clementis (Merr.) Schottky l.c. (1913) 358; S. clementi (Merr.) Koidzumi l.c. 194; L. clementis (Merr.)
A. Camus l.c. (1932) 39; P. ochracea Schottky l.c. (1913) 357; Q. ochracea (Schottky) Merr. l.c. (1921)
214; L. ochracea (Schottky) A. Camus l.c. (1932) 41; Q. borneensis Merr. l.c. (1922) 516; L. borneensis
(Merr.) Rehder l.c. (1929) 132; L. crateriformis Merr. ex A. Camus l.c. (1949) 97.
Tree up to 30 m tall, 70 cm diameter; buttresses and stilt-roots sometimes present. Bark

scaly or lenticellate, greyish brown; inner bark fibrous, yellowish or brownish. Sapwood
greyish or yellowish. Twigs densely tomentose with stellate and simple hairs, sparsely
lenticellate. Stipules ovate, 56 2 mm. Leaves thin-coriaceous, sparsely stellate-hairy
above especially on the midrib and veins, densely pubescent with stellate and simple
hairs below; oblanceolate, broadly elliptic to oblong, (6)1120.5(28) (2)3.56.5(
8.5) cm, base attenuate, acute, or obtuse, occasionaly asymmetrical, margin revolute, apex
acute with emarginate tip to acuminate with acumen 1020 mm long; midrib raised on
both surfaces, stronger below; lateral veins thick, (7)1012(15) pairs, lax, impressed
or flat above, strongly raised below, clearly joining near the leaf margin, forming an
angle of 4060 with the midrib; intercostal venation subscalariform, lax, impressed above,
prominent below; petiole (3)58(15) mm long. Inflorescences male, androgynous or
mixed. Male inflorescences 1020 cm long; bracts narrowly ovate, c. 1.3 0.5 mm;
bracteoles ovate, c. 0.6 0.3 mm. Male flowers solitary along the rachis; perianth lobes
acute, c. 1 0.8 mm; stamen filaments 1.52 mm long; pistillode subglobose, c. 0.5
mm diameter. Androgynous or mixed inflorescences 1025 cm long; bracts narrowly
ovate, c. 1 0.4 mm; bracteoles broadly ovate, c. 0.9 0.7 mm. Female flowers solitary
along the rachis; perianth lobes acute, 0.51 mm across; staminodes 12; styles cylindrical,
recurved, c. 2 mm long. Cupules solitary and distantly spaced along the rachis, subsessile,
cup-shaped, 1.52.5 23 cm, densely appressed tomentose with stellate hairs, scaly;
wall woody, thin, enclosing up to half of the acorn; scale-like appendages distinct, appressed,
imbricate, sparsely and irregularly arranged. Acorns ovoid, subglobose to cylindrical, (1.5
)24.5 1.52.5 cm, most part exserted, glabrous except for the sparsely simple-hairy
apex, dark purple; base flat, apex acute; scar deeply concave or flat, 0.51 cm diameter;
Vernacular name. Sarawakempili batu (Iban).
Distribution. Peninsular Malaysia, Borneo and the Philippines. Very common and widely
distributed in Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. In primary hill mixed dipterocarp, swamp, kerangas, riparian to lower montane
forests, up to 1500 m altitude, mainly on sandy clay soils.
42. Lithocarpus nodosus Soepadmo

(Latin, nodosus = knobby; the knobby twigs)
76
Tantra & Sutisna l.c. 158. Type: Meijer SAN 21980, Borneo, Sabah, Mt. Kinabalu, Kundasang (holotype
L; isotypes K, SAN).
Tree up to 20 m tall, 30 cm diameter. Bark smooth, brownish; inner bark granular, brownish.
Sapwood whitish. Twigs glabrous, with thickened nodes, sparsely large-lenticellate. Stipules
narrowly ovate to linear, 12 0.5 mm. Leaves thin-coriaceous to coriaceous, not rigid,
glabrous above, sparsely appressed yellowish tomentose below; elliptic, 610(12)
(3) 45(6) cm, base acute, margin strongly revolute, apex acute to acuminate, acumen
1015 mm long; midrib broad, flat above, raised below; lateral veins thin, (6)810 pairs,
lax, flat or slightly impressed above, raised below, disappearing near the leaf margin,
forming an angle of 3050 with the midrib; intercostal venation subscalariform, dense,
obscure on both surfaces; petiole 813 mm long. Inflorescences male or androgynous.
Male inflorescences 310 cm long; bracts and bracteoles linear, 0.91 0.20.3 mm.
Male flowers in clusters of 3 along the rachis; perianth lobes obtuse, c. 1 0.7 mm;
stamen filaments 12 mm long; pistillode subglobose, 0.50.7 mm diameter. Androgynous
inflorescences 6.59 cm long; bracts linear, c. 1.2 0.4 mm; bracteoles ovate, c. 1
0.6 mm. Female flowers solitary along the rachis; perianth lobes obtuse, c. 1 mm across;
staminodes 1012; styles conical, straight, 1.52 mm long. Cupules solitary along the
rachis, sessile, cup-shaped, 0.61 1.82.5 cm, densely appressed tomentose with simple
hairs, lamellate; wall woody, thin, enclosing less than half of the acorn; lamellae distinct,
edge minutely denticulate, set in 45 regular lines. Acorns ovoid to subglobose, 12
(2.5) 1.72.3 cm, most part exserted, glabrous, dark brown; base rounded, apex acute;
scar deeply concave, c. 1 cm diameter; wall woody, thin, most part free from the cupule.
Distribution. Endemic to Borneo. In Sabah, common in Ranau district especially in
Kinabalu Park, and also recorded for Mt. Trus Madi in Tambunan district. In Sarawak,
known only from Miri and Sri Aman districts. Also in Kalimantan.
Ecology. In primary hill mixed dipterocarp forest to montane forest, including limestone
habitat, at 3002400 m altitude, on different types of soils including ultrabasic.
43. Lithocarpus oblancifolius S. Julia & Soepadmo.

(Latin, oblancifolius = with reversed lance-shaped leaves)
Gard. Bull. Sing. 50 (1998) 141. Type: Leopold SAN 133942, Borneo, Sabah, Tawau district, Tawau
Hill Park (holotype SAN).
Tree up to 24 m tall, 45 cm in diameter. Bark lenticellate or smooth, brown or greyish;

inner bark greyish to yellowish. Sapwood whitish to yellowish. Twigs sparsely tomentose,
late-subglabrescent, smooth or sparsely large-lenticellate. Stipules narrowly ovate, c. 2
1 mm. Leaves thin-coriaceous, not rigid, glabrous above, sparsely appressed greyish
tomentose below; oblanceolate, (6)813.5 (2.5)34.5 cm, base cuneate, margin
revolute, apex rounded or acute; midrib faintly raised on both surfaces, stronger below;
lateral veins thin, 813 pairs, lax, flat or impressed above, raised below, joining near
the leaf margin, forming an angle of 3050 with the midrib; intercostal venation reticulate
or subscalariform, dense, obscure above, prominent below; petiole 35 mm long. Inflorescences male, female, androgynous or mixed. Male inflorescences 311 cm long;
bracts narrowly ovate, c. 1 0.3 mm; bracteoles linear, c. 5 0.1 mm. Male flowers
77
solitary along the rachis; perianth lobes obtuse, 0.81 0.60.8 mm; stamen filaments
c. 2 mm long; pistillode globose, 0.70.8 mm diameter. Female, androgynous or mixed
inflorescences 412 cm long; bracts linear, 11.2 0.20.4 mm; bracteoles narrowly
ovate, c. 0.5 0.2 mm. Female flowers solitary along the rachis; perianth lobes acute
or obtuse, 0.50.9 0.50.7 mm; staminodes 12; styles conical, recurved, 11.2 mm
long. Cupules solitary along the rachis, sessile, saucer-shaped, 11.2 1.62 cm, densely
appressed tomentose with stellate and simple hairs, lamellate; wall woody, thin, enclosing
less than half of the acorn; lamellae distinct, edge entire, set in 68 regular lines. Acorns
ovoid to conical, 1.61.8 1.62 cm, most part exserted, glabrous, brown; base flat, apex
acute; scar concave, c. 1.5 cm in diameter; wall bony, thick, most part free from the cupule.
Distribution. Endemic to Sabah. Known from Tawau Hill Park in Tawau district and
Sg. Piso in Labuk Sugut district.
Ecology. In lowland mixed dipterocarp forest, near riverbanks, in rather open area, up
to 100 m altitude.
44. Lithocarpus papillifer Soepadmo

(Latin, papillia = soft superficial protuberances, -fer = bearing; the papillate leaves and
twigs)
Tantra & Sutisna l.c. 158; Coode et al. (eds.) l.c. 126. Type: Clemens 32493, Borneo, Sabah, Labuk
Sugut district, Kolombuon R. (holotype L; isotypes A, BM, BO, K, NY, UC).
Tree up to 25 m tall, 40 cm diameter. Bark smooth or fissured or lenticellate, greyish

brown; inner bark fibrous, reddish. Sapwood yellowish. Twigs sparsely reddish brown
papillate, densely minute-lenticellate. Stipules ovate, 12 1 mm. Leaves thick-coriaceous,
rigid, covered with reddish brown papillae on both surfaces, denser below; elliptic or
ovate, (4)610(12) (2)34(5.5) cm, base acute, margin strongly revolute, apex obtuse,
rounded, or emarginate; midrib slightly raised above, raised below; lateral veins thin,
57(8) pairs, dense or lax, impressed or flat above, strongly raised below, faintly joining
or disappearing near the leaf margin, forming an angle of 4560 with the midrib; intercostal
venation scalariform, dense, obscure on both surfaces or slightly prominent below; petiole
510 mm long. Inflorescences male or androgynous. Male inflorescences 35 cm long;
bracts and bracteoles ovate, 0.91.2 0.60.7 mm. Male flowers in clusters of 24 along
the rachis; perianth lobes obtuse, c. 0.8 0.3 mm; stamen filaments 12 mm long; pistillode
subglobose, 0.81 mm diameter. Androgynous inflorescences 48 cm long; bracts ovate,
0.91 0.6 mm; bracteoles broadly ovate, c. 0.5 0.4 mm. Female flowers solitary
along the rachis; perianth lobes acute, c. 0.4 0.3 mm; staminodes 10; styles conical,
straight, 0.91 mm long. Cupules solitary along the rachis, sessile, saucer-shaped, 0.3
0.9 1.52.3 cm, densely appressed tomentose with stellate hairs mixed with a few reddish
brown papillae, scaly; wall bony, thin, enclosing less than half of the acorn; scale-like
appendages distinct, imbricate, set regularly. Acorns depressed ovoid, 1.51.8 1.22.3
cm, most part exserted, glabrous, chocolate brown; base flat, apex acute; scar deeply
Vernacular name. Sarawaksalad padeh (Kelabit).
78
Distribution. Peninsular Malaysia and Borneo. In Sabah, known only from Kolombuon
R. in Labuk Sugut district, Sosopodon in Ranau district, and Crocker Range in Tambunan
district. In Sarawak, uncommon, and documented from Miri, Kapit, Limbang and Sri
Aman districts. Also in Brunei and Kalimantan.
Ecology. In primary hill mixed dipterocarp forest to montane forest, rarely in limestone
habitat, at 8002300 m altitude.
45. Lithocarpus porcatus Soepadmo

(Latin, porcatus = ridged; the cupule)
Tantra & Sutisna l.c. 158. Type: Carr SFN 26677, Borneo, Sabah, Mt. Kinabalu, Menetudok Gorge
(holotype SING).

greyish green; inner bark brown. Sapwood yellowish. Twigs densely tomentose, lateglabrescent, smooth. Stipules linear, c. 9 2 mm. Leaves coriaceous, not rigid, glabrous
above, densely greyish tomentose with appressed hairs below; broadly ovate to elliptic,
(16)1820(25) (6.5)910(13) cm, base rounded to acute, margin revolute, apex
obtuse to short-acuminate, acumen up to 5 mm long; midrib raised on both surfaces,
stronger below, glabrous; lateral veins thin, 1013(16) pairs, lax, flat above, strongly
raised below, disappearing towards the leaf margin, forming an angle of 4560 with
the midrib; intercostal venation scalariform, dense, obscure on both surfaces or slightly
prominent below; petiole 515 mm long. Inflorescences male or androgynous. Male
inflorescences 1520 cm long; bracts and bracteoles ovate, c. 1 0.5 mm. Male flowers
solitary along the rachis; perianth lobes obtuse, c. 1.5 1 mm; stamen filaments c. 5
mm long; pistillode subglobose, c. 2 mm diameter. Androgynous inflorescences 811.5
cm long; bracts narrowly ovate, c. 1.8 0.6 mm; bracteoles broadly ovate, c. 0.8
0.6 mm. Female flowers solitary or in clusters of 23 along the rachis; perianth lobes
acute, c. 1 0.9 mm; staminodes 10; styles conical, straight, 11.2 mm long. Cupules
in clusters of 23 when young, later solitary along the rachis, with stalk 510 mm long,
depressed globose, 23.5 2.54 cm, densely tomentose, lamellate and ridged; wall woody,
thick, densely hairy, enclosing the acorn completely; lamellae distinct, edge entire or
undulate, sometimes folded inwardly, set in 58 lines. Acorns depressed subglobose, 1
2 1.52 cm, free apical part sparsely tomentose, brown; base rounded, apex flat, umbonate;
Vernacular name. Sarawaktekalet kapuong (Punan).
Distribution. Endemic to Borneo. In Sabah, widespread in Tambunan, Kota Belud, Keningau,
Sipitang, Tenom, and Ranau districts. In Sarawak, known only from Bt. Batu Tiban in
Kapit district, Long Amok in Limbang district, and in Kuching district. Also in Kalimantan.
Ecology. In primary hill mixed dipterocarp forest, at 9001000 m altitude.
79
46. Lithocarpus pseudokunstleri A. Camus

(resembling Lithocarpus kunstleri)
100; Anderson l.c. 202; Whitmore, Tantra & Sutisna l.c. 158; PROSEA l.c. 300; Coode et al. (eds.)
l.c. 126. Type: Beccari PB 3328, Borneo, Sarawak, Marop (holotype FI; isotype K).
Tree up to 25 m tall, 45 cm diameter; buttresses 0.51 m tall. Bark scaly or lenticellate

or smooth, greyish brown; inner bark granular, brownish. Sapwood yellowish. Twigs
sparsely tomentose, densely to sparsely minute- or large-lenticellate, rarely fissured. Stipules
narrowly ovate, c. 2 1 mm. Leaves thin-coriaceous, glabrous on both surfaces; elliptic
to oblanceolate, (8)9.515(20) (2.5)45.5(8.5) cm, base attenuate, margin revolute,
apex acute to acuminate, acumen c. 10 mm long; midrib slightly raised above, strongly
raised below; lateral veins thin, 68(10) pairs, lax, flat or impressed above, raised below,
clearly or faintly joining near the leaf margin, forming an angle of 4560 with the
midrib; intercostal venation subscalariform or reticulate, dense, obscure above, prominent
or rarely obscure below; petiole 36 mm long. Inflorescences male or androgynous. Male
inflorescences 1115 cm long; bracts and bracteoles ovate, 0.81 0.40.5 mm. Male
flowers in clusters of 3 along the rachis; perianth lobes obtuse, c. 1.3 0.8 mm; stamen
filaments c. 2.5 mm long; pistillode subglobose, c. 1 mm diameter. Androgynous
inflorescences 510 cm long; bracts and bracteoles ovate to triangular, 11.2 0.8
1 mm. Female flowers solitary along the rachis; perianth lobes acute, c. 1.2 1 mm;
staminodes 10; styles conical, recurved, c. 1.5 mm long. Cupules solitary along the rachis,
sessile, deeply cup-shaped, 11.8 1.52 cm, densely appressed tomentose with stellate
hairs, scaly; wall woody, thin, enclosing less than half of the acorn; scale-like appendages
slightly distinct, appressed, imbricate, irregularly set. Acorns cylindrical, 2.53.7 1.2
1.5 cm, most part exserted, sparsely simple and long hairy especially at the base and
apex, brownish; base flat, apex acute; scar flat or shallowly concave, 0.71 cm diameter;
Vernacular name. Sarawakempenit johari (Malay).
Distribution. Peninsular Malaysia and Borneo. In Sabah, documented from Sipitang,
Beaufort, Tawau, Tenom, and Kinabatangan districts. In Sarawak, common in Miri, Kapit,
Sibu, Sri Aman, and Kuching districts. Also in Kalimantan.
m altitude, on sandy clay and sandy loam soils.
47. Lithocarpus pulcher (King) Markgr.

(Latin, pulcher = beautiful or excellent; the cupule)
Bot. Jahrb. 59 (1924) 67; Soepadmo l.c. (1970) 269, l.c. (1972) 333; Cockburn l.c. (1976) 103;
Anderson l.c. 202; Whitmore, Tantra & Sutisna l.c. 159; PROSEA l.c. 300; Coode et al. (eds.) l.c.
127; Argent et al. (eds.) l.c. 251. Basionym: Quercus pulchra King l.c. (1889) 85. Lectotype
(Soepadmo, 1970): Beccari PB 3123, Borneo, Sarawak (hololectotype FI; isolectotypes K, L, NY, P).
Synonym: Synaedrys pulchra (King) Koidzumi l.c. 189.
80
Tree up to 36 m tall, 80 cm diameter; buttresses 1.52.5 m tall. Bark irregulary fissured

or scaly, chocolate-brown; inner bark fibrous, greyish. Sapwood yellowish or brownish.
Twigs stellate-hairy, late-subglabrescent, smooth or sparsely lenticellate. Stipules narrowly
ovate, 79 23 mm. Leaves thick-coriaceous, rigid, subglabrous above, densely appressed
brownish tomentose with stellate hairs below; broadly elliptic to oblong, (10)1520
(30) (4)68(12.5) cm, base acute to cuneate, margin revolute, apex acute to
acuminate, acumen 510 mm long; midrib flat or slightly raised above, strongly raised
below; lateral veins (12)1518(22) pairs, dense, flat or impressed above, strongly raised
below, disappearing near the leaf margin, forming an angle of 5070 with the midrib;
intercostal venation scalariform, dense, obscure above, thin but clearly visible below;
petiole 1020 mm long. Inflorescences male or female. Male inflorescences 815 cm
long; bracts and bracteoles ovate, c. 1.5 0.5 mm. Male flowers solitary along the rachis;
perianth lobes acute, c. 1.5 0.5 mm; stamen filaments c. 1.5 mm long; pistillode globose,
c. 2 mm diameter. Female inflorescences c. 10 cm long; bracts broadly ovate, c. 2
1.5 mm; bracteoles subrounded to broadly ovate, c. 1.5 mm across. Female flowers solitary
along the rachis; perianth lobes acute, 12 0.5 mm; staminodes 12; styles conical,
straight, c. 1.5 mm long. Cupules solitary along the rachis, sessile, obconical, 2.54
36 cm, tomentose, tuberculate; wall woody, thick, hairy, enclosing most part of the acorn
except for an apical opening c. 1.54 cm diameter; tubercles distinct, sturdy, irregularly
and densely set at the upper part of the cupule and becoming sparse towards the base.
Acorns obconical, 23 45 cm, free apical part sparsely tomentose, brown; base rounded,
apex flat or convex; scar deeply convex; wall woody, thick, most part adnate to the cupule.
Vernacular names. Sarawakkeraki (Iban), salad urong (Kelabit).
Distribution. Endemic to Borneo. Widely distributed in Sabah. In Sarawak, very common
in Kuching district but uncommon in Kapit and Miri districts. Also in Brunei and
Kalimantan.
Ecology. In primary and secondary mixed dipterocarp, kerangas, to lower montane forests,
up to 1500 m altitude, on sandy clay soils.
48. Lithocarpus pusillus Soepadmo

(Latin, pusillus = very small, tiny; the cupule)
Tantra & Sutisna l.c. 159; PROSEA l.c. 300. Type: Anderson 4560, Borneo, Sarawak, Miri district,
G. Mulu NP (holotype L; isotypes A, K, S).
Tree up to 30 m tall, 80 cm diameter; buttresses up to 0.9 m tall. Bark scaly or flaky,

greyish brown; inner bark reddish or brownish. Sapwood whitish. Twigs glabrescent,
sparsely to densely minute- or usually large-lenticellate. Stipules linear, c. 3 1 mm.
Leaves thin-coriaceous, glabrous above, sparsely appressed brownish tomentose below;
narrowly elliptic or ovate, (5.5)68.5(10.5) 24 cm, base obtuse, margin strongly
revolute, apex caudate or long-acuminate, acumen 1025 mm long; midrib thin, flat above,
strongly raised below; lateral veins thin, 611 pairs, lax, flat above, slightly raised below,
faintly joining near the leaf margin, forming an angle of 4060 with the midrib; intercostal
81
venation reticulate, dense, obscure on both surfaces or thinly prominent below; petiole
59 mm long. Inflorescences male, androgynous or mixed. Male inflorescences 710
cm long; bracts subrounded, 0.30.4 mm across; bracteoles ovate, c. 0.3 0.1 mm. Male
flowers solitary along the rachis; perianth lobes acute, c. 0.8 0.6 mm; stamen filaments
c. 2 mm long; pistillode globose, 11.5 mm diameter. Androgynous or mixed inflorescences
(3)810 cm long; bracts ovate, c. 0.4 0.2 mm. Female flowers solitary along the
rachis; perianth lobes acute, c. 0.7 0.3 mm; staminodes 10; styles conical, straight,
c. 1 mm long. Cupules solitary along the rachis, sessile, saucer-shaped, 0.30.5 0.8
1(1.2) cm, densely appressed tomentose, lamellate; wall woody, thin, enclosing less than
half of the acorn; lamellae slightly distinct, edge entire or minutely denticulate, set in
45 regular lines. Acorns ovoid to conical, (0.7)1.51.8 0.71.5 cm, most part exserted,
glabrous, brown; base flat, apex rounded to acute; scar concave, 0.30.7 cm diameter;
wall bony, thin, most part free from the cupule.
Vernacular names. Sarawakberangan bris (Iban), empenit daun halus, empenit gading
(Malay), languwai (Kedayan), pili (Kenyah), saled (Murut).
Distribution. Endemic to Borneo. In Sabah, recorded for Sipitang, Beaufort, Tawau, Tenom,
and Kinabatangan districts. In Sarawak, common in Miri, Kapit, Sibu, Sri Aman, and
Kuching districts. Also in Kalimantan.
Ecology. In primary hill mixed dipterocarp, peatswamp, kerangas to lower montane forests,
at 4001600 m altitude, on sandy loam or sandy clay soils.
49. Lithocarpus rassa (Miq.) Rehder

(A Malay plant namerasa)
PROSEA l.c. 301; Kochummen l.c. 248. Basionym: Quercus rassa Miq. l.c. (1861) 350. Type: Teijsmann
HB. 675, Sumatra, Padang Residency, Pajakumbuh (holotype BO; isotypes K, U). Synonyms:
Cyclobalanus rassa (Miq.) Oerst. l.c. (1871) 376; Pasania rassa (Miq.) Gamble l.c. (1915) 436; Synaedrys
rassa (Miq.) Koidzumi l.c. (1916) 192; Q. rassa Miq. var. montana King ex Hook. f. l.c. (1888) 613; Q.
wenzigiana King ex Hook. f. l.c. (1888) 613; P. wenzigiana (King ex Hook. f.) Gamble l.c. (1915) 435;
S. wenzigiana (King ex Hook. f.) Koidzumi l.c. 193; L. wenzigiana (King ex Hook. f.) A. Camus l.c.
(1932) 42; Q. rassa Miq. var. lanuginosa Ridl., J. Str. Br. Roy. As. Soc. 61 (1912) 37; L. rangeriana A.
Camus l.c. (1932) 913; L. ridleyana A. Camus l.c. (1932) 913; L. symingtonia A. Camus l.c. (1932) 913.
Tree up to 24 m tall, 45 cm diameter. Bark fissured or scaly or smooth, greyish brown;

inner bark reddish. Sapwood whitish. Twigs glabrescent, sparsely to densely largelenticellate. Stipules linear, 34 1 mm. Leaves thin-coriaceous, not rigid, glabrous
above, sparsely appressed, greyish tomentose with short, simple hairs below; narrowly
elliptic, (3)69(12) (1.5)24(5.5) cm, base acute to cuneate, margin strongly revolute,
apex acute to acuminate or caudate, acumen 1018 mm long; midrib raised on both
surfaces; lateral veins thin, (8)1013(15) pairs, dense, flat above, slightly raised below
or obscure on both surfaces, faintly joining near the leaf margin, forming an angle of
4050 with the midrib; intercostal venation reticulate, dense, obscure on both surfaces
82
or thinly prominent below; petiole (8)1015(20) mm long. Inflorescences male, androgynous

or mixed. Male inflorescences 412 cm long; bracts and bracteoles ovate, 0.91 0.5
0.6 mm. Male flowers in clusters of 23 along the rachis; perianth lobes obtuse, c. 1
0.5 mm; stamen filaments c. 2 mm long; pistillode subglobose, 0.81 mm diameter.
Androgynous or mixed inflorescences 511 cm long; bracts and bracteoles ovate, 0.6
1 0.40.6 mm. Female flowers solitary along the rachis; perianth lobes acute, c. 0.8
0.6 mm; staminodes 10; styles conical, straight, 0.51 mm long. Cupules solitary along
the rachis, sessile or with stalk 34 mm long, cup- or saucer-shaped, 1.31.7 23
(3.5) cm, sparsely appressed tomentose, lamellate; wall woody, thick, enclosing less than
half of the acorn; lamellae clearly distinct, edge entire or denticulate, set in 68 regular
lines. Acorns subglobose, 1.21.8 1.52.5 cm, most part exserted, glabrous, rarely
sparsely tomentose, dark brown; base rounded, apex acute; scar deeply concave, 11.5
cm diameter; wall woody, thick, most part free from the cupule.
Vernacular name. Sarawakkeraki padi (Bidayuh).
Distribution. Sumatra, Peninsular Malaysia and Borneo (Sabah and Sarawak). In Sabah,
known only from Keningau, Tawau and Sipitang districts. In Sarawak, widely distributed
throughout Kapit, Kuching, Miri, and Sri Aman districts.
Ecology. In primary lowland and hill mixed dipterocarp, kerangas, swamp and limestone
forests, up to 1000 m altitude.
50. Lithocarpus revolutus Hatus. ex Soepadmo

(Latin, revolutus = rolled back from edge; the leaf margin)
Tantra & Sutisna l.c. 159. Type: RSNB 4171, Borneo, Sabah, Mt. Kinabalu, Mesilau R. (holotype L;
isotype K).
Tree up to 25 m tall, 50 cm diameter. Bark flaky, brownish; inner bark yellowish brown.
Sapwood yellowish. Twigs glabrous, sparsely large-lenticellate. Stipules narrowly ovate,
46 23 mm. Leaves thick-coriaceous, rigid, not bullate, glabrous on both surfaces;
obovate to elliptic, 1014(21.5) 35(7) cm, base cuneate, margin strongly revolute,
apex rounded to obtuse, tip emarginate; midrib broad, flat or impressed above, strongly
raised; lateral veins thick, (5)79 pairs, lax, flat to impressed above, strongly raised
below, disappearing near the leaf margin, forming an angle of 4560 with the midrib;
intercostal venation subscalariform, dense, obscure on both surfaces or thinly prominent
below; petiole 56 mm long. Inflorescences male or female. Male inflorescences 10
15 cm long; bracts ovate, 22.2 0.51 mm; bracteoles narrowly ovate, c. 1.2 0.3
0.6 mm. Male flowers solitary but arranged closely together along the rachis; perianth
lobes obtuse, c. 1.8 0.81 mm; stamen filaments 23 mm long; pistillode subglobose,
0.81.2 mm diameter. Female inflorescences 814 cm long, sturdy; bracts and bracteoles
narrowly ovate to linear, 12 0.30.5 mm. Female flowers solitary along the rachis;
perianth lobes acute, 1 0.5 mm; staminodes 12; styles cylindrical, straight, c. 2 mm
long. Cupules solitary along the rachis, sessile, cup-shaped, 1.52.5 3.75 cm, densely
tomentose, scaly; wall woody, thick, enclosing up to half of the acorn; scale-like appendages
distinct, rarely obscure, appressed, imbricate, irregularly set. Acorns depressed ovoid,
83
22.3(5) 34.5 cm, most part exserted, glabrous, shining, dark chocolate-brown; base
rounded, apex rounded to acute; scar concave, 23.5 cm diameter; wall woody, thick,
Vernacular name. Sarawaksaled urong (Kelabit).
Distribution. Endemic to Borneo, uncommon. In Sabah, recorded for Keningau, Kinabatangan
and Ranau districts. In Sarawak, known only from Baleh in Kapit district, Sg. Masia
in Lawas district, and in Bario district. Also in Kalimantan.
Ecology. In primary hill mixed dipterocarp, kerangas, and lower montane forests, at 1000
1800 m altitude.
51. Lithocarpus rigidus Soepadmo

(Latin, rigidus = stiff or rigid; the leaf)
159. Type: RSNB 4789, Borneo, Sabah, Mt. Kinabalu, Mesilau Cave (holotype L; isotypes K, SAN,
SING).
Tree up to 18 m tall, 30 cm diameter. Bark greyish brown. Twigs sparsely appressed

tomentose or subglabrous, sparsely lenticellate. Stipules lanceolate, 68 34 mm. Leaves
thick-coriaceous, rigid, glabrous above, sparsely brownish appressed tomentose below;
obovate, (7)1416(18) 6.58(10) cm, base acute to cuneate, decurrent, margin strongly
revolute, apex acute; midrib broad, strongly raised above, slightly raised below; lateral
veins thick and broad, 811 pairs, lax, flat or impressed above, slightly raised below,
disappearing or faintly joining near the leaf margin, forming an angle of 5070 with
the midrib; intercostal venation subscalariform or irregular, lax, obscure on both surfaces;
petiole 610 mm long. Inflorescences male or female. Male inflorescences 46 cm long;
bracts linear, c. 2.7 0.8 mm; bracteoles elliptic, c. 1.3 0.6 mm. Male flowers solitary
along the rachis; perianth lobes acute, c. 2 1 mm; stamen filaments c. 5 mm long;
pistillode subglobose, 22.5 mm diameter. Female inflorescences 810 cm long; bracts
and bracteoles ovate, 35 23 mm. Female flowers in clusters of 3 along the rachis;
perianth lobes acute, 45 mm across; staminodes 12; styles conical, recurved, 12 mm
long. Cupules in clusters of 23 or solitary along the rachis, sessile or with stalk c.
3 mm long, saucer- to cup-shaped, 0.50.7 11.5 cm, sparsely appressed tomentose,
lamellate; wall woody, thick, enclosing less than half of the acorn; lamellae distinct, edge
minutely denticulate, set in irregular lines. Acorns ovoid to conical, 0.81 11.5 cm,
most part exserted, glabrous, dark chocolate-brown; base flat, apex rounded or acute;
scar deeply concave, 0.51 cm diameter; wall woody, thick, most part free from the cupule.
Distribution. Endemic to Borneo. In Sabah, only known from Mesilau Cave, Mt. Kinabalu
and Mt. Tambuyokon in Ranau district. Not known in Sarawak. Also occurs in Kalimantan.
Ecology. In montane forest up to 2700 m altitude.
84
52. Lithocarpus rotundatus (Blume) A. Camus

(Latin, rotundatus = rounded; the upper part of the cupule and acorn)
Riviera Scientific 18 (1931) 41; Soepadmo l.c. (1970) 275, l.c. (1972) 331; Whitmore, Tantra & Sutisna
l.c. 159; PROSEA l.c. 301. Basionym: Quercus rotundata Blume, Verh. Bat. Genootsch. 9 (1823) 219.
Type: Blume s.n., Java, Mt. Salak (L). Synonyms: Pasania rotundata (Blume) Oerst. l.c. (1867) 83;
Synaedrys rotundata (Blume) Koidzumi l.c. 189; Q. clathrata Seemen l.c. (1900) 15; S. clathrata
(Seemen) Koidzumi l.c. (1916) 190; L. clathrata (Seemen) Rehder l.c. (1919) 123; Q. pyriformis Seemen
l.c. (1900) 17; S. pyriformis (Seemen) Koidzumi l.c. (1916) 189; L. pyriformis (Seemen) Rehder l.c.
(1919) 127.
Tree up to 40 m tall, 100 cm diameter. Twigs densely tomentose with simple and stellate
hairs, late-subglabrescent, sparsely to densely warty lenticellate. Stipules linear, 45
0.71 mm. Leaves thin-coriaceous, glabrous above, densely greyish tomentose with
appressed stellate hairs below; ovate to elliptic, (10)1315(20) (4)56(9) cm, base
rounded to acute, margin slightly revolute, apex acute to sharply acuminate, acumen 5
10 mm long; midrib thin above, raised below; lateral veins thin, 911 pairs, impressed
above, raised below, disappearing near the leaf margin, forming an angle of 4560 with
the midrib; intercostal venation subscalariform, dense, obscure on both surfaces; petiole
515 mm long. Inflorescences male or female. Male inflorescences 1015 cm long; bracts
and bracteoles ovate, 0.61 0.6 mm. Male flowers in clusters of 37 along the rachis;
stamen filaments 34 mm long; pistillode subglobose, 1.52 mm diameter. Female
inflorescences 1015 cm long; bracts and bracteoles ovate, c. 1 0.7 mm. Female flowers
solitary along the rachis; staminodes 12; styles conical, recurved, 11.5 mm long. Cupules
solitary along the rachis, obovoid or pear-shaped, 1.52 34 cm, densely stellate-hairy,
lower part smooth or ridged, upper part rotundate, set with imbricate scale-like appendages
or woody tubercules; stalk 520 mm long; wall thick, enclosing most part of the acorn
except for an apical opening c. 1 cm diameter. Acorns subglobose, c. 2 33.5 cm;
Distribution. Java, Borneo and the Philippines. In Borneo, so far only known from Sabah
and Kalimantan.
Ecology. In lowland mixed dipterocarp forest to lower montane forest, up to 1500 m
altitude.
53. Lithocarpus ruminatus Soepadmo

(Latin, ruminatus = ruminate, intricately folded; the inner wall of the acorn)
Tantra & Sutisna l.c. 159; Coode et al. (eds.) l.c. 127; Argent et al. (eds.) l.c. 251. Type: Clemens 29164,
Borneo, Sabah, Mt. Kinabalu (holotype L; isotypes A, BM, BO, K, NY, UC).
Tree up to 20 m tall, 30 cm diameter; buttresses 0.30.6 m tall. Bark smooth or flaky,

greyish brown; inner bark fissured, red-brown. Sapwood brownish or whitish. Twigs
subglabrous, smooth or sparsely minute-lenticellate. Stipules linear, 24 1 mm, caducous
or persistent. Leaves thin-coriaceous to coriaceous, glabrous or subglabrous above, sparsely
appressed greyish tomentose below; elliptic, 1013(16) (3)46 cm, base rounded,
85
acute or cuneate, margin revolute, apex acute to shortly acuminate, acumen c. 10 mm

long; midrib thin, slightly raised above, strongly raised below; lateral veins thin, 57
(9) pairs, lax, impressed or flat above, raised below, disappearing near the leaf margin,
forming an angle of 4560 with the midrib; intercostal venation subscalariform, lax,
obscure on both surfaces or thin and weakly prominent below; petiole 515 mm long.
Inflorescences male, androgynous or mixed. Male inflorescences 612 cm long; bracts
and bracteoles narrowly ovate, c. 11.5 0.5 mm. Male flowers in clusters of 3 along
the rachis; perianth lobes obtuse, c. 1.5 1 mm; stamen filaments c. 1 mm long; pistillode
subglobose, c. 1.5 mm diameter. Androgynous or mixed inflorescences 810 cm long;
bracts narrowly ovate, c. 1 0.5 mm; bracteoles obtuse, c.1 mm across. Female flowers
in clusters of 3 along the rachis; perianth lobes acute, c. 1 0.5 mm; staminodes 12;
styles cylindrical, straight, c. 2 mm long. Cupules solitary along the rachis, sessile, ovoid
to ellipsoid 34.5 45 cm, sparsely tomentose, lamellate; wall woody, c. 0.5 cm thick,
densely hairy, enclosing most part of the acorn except for an apical opening 11.5 cm
diameter; lamellae distinct, edge entire, set in 710 concentric lines. Acorns ovoid to
ellipsoid, 34 45 cm, free apical part sparsely tomentose, brown; base rounded, apex
convex, depressed and umbonate; wall thick, most part adnate to the cupule, inner part
(endocarp) ruminate.
Vernacular name. Sarawaksalud kup (Murut).
Distribution. Endemic to Borneo, uncommon. In Sabah, recorded for Sandakan, Ranau,
Tawau, Tenom, Kota Belud, Sipitang, Beaufort, and Lahad Datu districts. In Sarawak,
known only from Lanjak-Entimau, Ulu Segan in Bintulu district and Ulu Lawas in Lawas
district. Also in Brunei and Kalimantan.
Ecology. In primary hill mixed dipterocarp, swamp, to lower montane forests, at 100
1300 m altitude, on clay soils.
54. Lithocarpus sandakanensis S. Julia & Soepadmo

(of Sandakan, Sabah)
Gard. Bull. Sing. 50 (1998) 143. Type: Wood SAN A 4697, Borneo, Sabah, Sandakan district, Sepilok
FR, Compartment 17 (holotype KEP; isotypes AA, L, MEL, SING).
Tree up to 45 m tall, 90 cm in diameter; buttresses small. Bark smooth, greyish or brownish;

inner bark yellowish. Sapwood whitish or purplish. Twigs glabrescent, sparsely lenticellate.
Leaves coriaceous, subglabrous above, sparsely appressed brownish tomentose below;
elliptic to oblong, 2639(46) 7.511.5 cm, base rounded to acute, margin revolute,
apex sharply acute to acuminate, acumen c. 15 mm long; midrib flat or slightly raised
above, strongly raised below; lateral veins thick, 1114 pairs, dense, flat or impressed
above, strongly raised below, disappearing near the leaf margin, forming an angle of 30
60 with the midrib; intercostal venation subscalariform or reticulate, lax, prominent on
both surfaces, stronger below; petiole 818 mm long. Male inflorescences c. 10 cm long;
bracts narrowly ovate to linear, 11.1 0.4 mm; bracteoles linear, c. 0.6 0.2 mm.
Male flowers solitary along the rachis; perianth lobes obtuse, 1.11.3 0.60.8 mm;
stamen filaments c. 2 mm long; pistillode subglobose, 0.61 mm in diameter. Pistillate
86
inflorescences and flowers not known. Cupules solitary along the rachis, with stalk 0.5
0.7 cm long, cup-shaped, 1.52 33.5 cm, densely tomentose, lamellate; wall woody,
thick, enclosing up to half of the acorn; lamellae distinct, edge minutely denticulate or
wavy, set in 78 regular lines. Acorns depressed ovoid, 3.54 cm across, most part exserted,
densely tomentose with simple hairs, dark brown; base flat, apex rounded to acute; scar
deeply concave, c. 2 cm in diameter; wall woody, thick, most part free from the cupule.
Distribution. Endemic to Sabah. All known specimens were collected from Sepilok FR
in Sandakan district.
Ecology. In primary lowland mixed dipterocarp forest, on yellow clay soils, up to 15 m
altitude.
55. Lithocarpus sericobalanus E.F. Warb.

(Greek, serikos = silky with long, straight, closely pressed glossy hairs, balanos = acorn)
Kew Bull. 20 (1936) 20 ; Soepadmo l.c. (1970) 279, l.c. (1972) 340; Cockburn l.c. (1976) 102; Anderson
l.c. 203; Whitmore, Tantra & Sutisna l.c. 159; Kessler & Sidiyasa l.c. 142; PROSEA l.c. 302; Argent et
al. (eds.) l.c. 251. Type: Richards 1241, Borneo, Sarawak, Mt. Dulit, Ulu Tinjar, Long Kapa (holotype
K; isotypes A, L).
Tree up to 50 m tall, 100 cm diameter; buttresses up to 1.8 m tall. Bark deeply fissured
or scaly, greyish brown; inner bark fibrous, yellowish brown. Sapwood yellowish or
brownish. Twigs densely yellowish tomentose, sparsely lenticellate. Leaves coriaceous,
glabrous above, sparsely yellowish tomentose below; elliptic, (9)1213.5 (3)46 cm,
base acute, margin revolute, apex bluntly acute to acuminate, acumen 510(15) mm
long; midrib raised on both surfaces, stronger below; lateral veins thin, (9)1011(12)
pairs, dense, flat above, raised below, disappearing near the leaf margin, forming an angle
of 4060 with the midrib; intercostal venation scalariform, dense, obscure above,
prominent below; petiole 1015(20) mm long. Inflorescences male or androgynous. Male
inflorescences 615 cm long; bracts elliptic, c. 0.8 0.4 mm; bracteoles ovate, 0.3
0.4 mm across. Male flowers solitary along the rachis; perianth lobes acute, c. 1 0.5
mm; stamen filaments 22.5 mm long; pistillode globose, c. 1 mm diameter. Androgynous
inflorescences 1015 cm long; bracts ovate, c. 0.5 0.3 mm; bracteoles subrounded,
0.20.3 mm across. Female flowers solitary along the rachis; perianth lobes acute, c.
0.7 0.5 mm; staminodes 12; styles conical, straight, 0.81 mm long. Cupules solitary
along the rachis, with stalk 510 mm long, saucer-shaped, 0.51.5 2.54 cm, densely
sericeous, lamellate; wall woody, thick, enclosing up to half of the acorn; lamellae slightly
distinct or obscure, edge entire or denticulate, set in 610 regular lines. Acorns depressed
ovoid to subglobose or discoid, 12 2.54 cm, most part exserted, densely sericeous,
brownish; base rounded, apex rounded to acute; scar deeply concave, 23 cm diameter;
Distribution. Endemic to Borneo. Common and distributed throughout Sabah and Sarawak.
Also in Kalimantan.
Ecology. Common in primary hill mixed dipterocarp forest, up to 1000 m altitude.
Uses. The timber is widely used by local people for house and bridge construction and
also for making furniture and tool-handles.
87
56. Lithocarpus stonei S. Julia & Soepadmo
Fig. 13.
(B.C. Stone, 19331994, former Reader in Botany, Department of Botany, University of

Malaya, Kuala Lumpur)
Gard. Bull. Sing. 50 (1998) 145. Type: Yap & Khairuddin SAN 106051, Borneo, Sabah, Tambunan
road (holotype SAN; isotype KEP).
Tree up to 25 m tall, 60 cm in diameter. Bark fissured or smooth or lenticellate, dark

grey; inner bark fibrous, yellowish brown or whitish. Sapwood whitish. Twigs densely
tomentose, sparsely to densely large-lenticellate. Leaves thick-coriaceous, rigid, sparsely
appressed yellowish tomentose on both surfaces; elliptic to oblong, 2836(40) 914
cm, base rounded to acute, margin revolute, apex acuminate, acumen c. 15 mm long;
midrib strongly raised on both surfaces, stronger below; lateral veins thick, (9)1012
pairs, lax, flat or impressed above, strongly raised below, disappearing near the leaf margin,
forming an angle of 4060 with the midrib; intercostal venation reticulate, lax, obscure
above, prominent below; petiole 1015 mm long. Male inflorescences and flowers
unknown. Female inflorescences c. 20 cm long; bracts narrowly ovate, 22.5 0.2
0.5 mm; bracteoles broadly ovate, c. 0.7 0.5 mm. Female flowers in clusters of 2
3 along the rachis; perianth lobes acute, c. 1 0.5 mm; staminodes 12; styles conical,
straight, c. 1.5 mm long. Cupules in clusters of 23 or rarely solitary along the rachis,
sessile, deeply cup-shaped, 23 45 cm, densely tomentose, lamellate; wall woody,
35 mm thick, enclosing half or more of the acorn; lamellae strongly raised, edge folded
inwards, undulate, set in 79 regular or irregular lines. Acorns depressed ovoid to
subglobose, 34 35 cm, most part exserted, glabrous and shining, brownish; base
flat, apex rounded; scar deeply concave, 22.5 cm in diameter; wall woody, thick, most
Vernacular name. Sarawaksaled urong (Kelabit).
Distribution. Endemic to Borneo. In Sabah, recorded for Crocker Range in Tenom district,
and G. Alab and Tambunan road in Tambunan district. In Sarawak, known only in Ulu
Baram in Marudi district. No record for Brunei and Kalimantan.
Ecology. In hill mixed dipterocarp forest, on recent alluvium and clay soils, up to 1000
m altitude.
57. Lithocarpus sundaicus (Blume) Rehder

(of the Sundalands)
PROSEA l.c. 304; Argent et al. (eds.) l.c. 251. Basionym: Quercus sundaica Blume l.c. (1823) 216.
Type: Blume s.n. (= RHL Sheet No. 90131341), W Java, Banten Residency, Sadjira, Rangkas Bitung
(holotype L; isotype CGE). Synonyms: Pasania sundaica (Blume) Oerst. l.c. (1867) 83; Q.
pseudomolucca Blume var. sundaica (Blume) Wenzig, Jahrb. Bot. Gart. Berl. (1886) 227; Synaedrys
sundaica (Blume) Koidzumi l.c. (1916) 198; Q. pruinosa Blume l.c. (1823) 217, t. 5; P. pruinosa (Blume)
Wenzig l.c (1886) 227; S. pruinosa (Blume) Koidzumi l.c. 197; L. pruinosa (Blume) Rehder l.c. (1919)
130; Castanea latifolia Blume, Bijdr. Fl. Ned. Ind. (1826) 526; Q. muricata Roxb. l.c. (1832) 635;
88
vi
Cyclobalanus muricata (Roxb.) Oerst. l.c. (1871) 379; Q. mappacea Korth. l.c. (1844) 202; Q. korthalsii
Blume var. mappacea (Korth.) Blume l.c. (1850) 292; Q. hystrix Korth. var. mappacea (Korth.) Merr.
l.c. (1921) 213; Q. korthalsii Blume l.c. (1850) 292; P. korthalsii (Blume) Oerst. l.c. (1867) 85; Q.
pseudomolucca Blume var. korthalsii (Blume) Wenzig l.c. (1886) 227; Q. lamponga Miq. l.c. (1861)
348; Cyclobalanus lamponga (Miq.) Oerst. l.c. (1867) 81; P. lamponga (Miq.) Gamble l.c. (1915) 423;
S. lamponga (Miq.) Koidzumi l.c. 195; Q. grandifrons King ex Hook. f. l.c. (1888) 610; P. grandifrons
(King ex. Hook. f.) Gamble l.c. (1915) 421; S. grandifrons (King ex. Hook. f.) Koidzumi l.c. 195; L.
grandifrons (King ex. Hook. f.) A. Camus l.c. (1932) 40.
Tree up to 36 m tall, 90 cm diameter. Bark smooth or fissured, greyish brown; inner

bark yellowish. Sapwood yellowish. Twigs densely appressed tomentose with stellate and
simple hairs, smooth or sometimes sparsely minute-lenticellate. Stipules linear, c. 4
1 mm. Leaves coriaceous, densely or sparsely stellate-hairy (sometimes mixed with simple
hairs) above, densely appressed greyish tomentose with simple and stellate hairs below;
elliptic to oblong, (10)1216(24) (4)56(10) cm, base obtuse, rarely acute, margin
revolute, apex acute to acuminate, acumen 520 mm long; midrib strongly raised on both
surfaces; lateral veins thick, (10)1214(16) pairs, lax, slightly raised above, strongly
raised below, faintly to clearly joining near the leaf margin, forming an angle of 40
60 with the midrib; intercostal venation scalariform, lax, obscure above, prominent or
obscure below; petiole 512 mm long. Inflorescences male, androgynous or mixed. Male
inflorescences 515 cm long; bracts ovate, c. 1 0.7 mm. Male flowers solitary along
the rachis; perianth lobes obtuse, c. 1 1 mm; stamen filaments c. 2.5 mm long; pistillode
globose, c. 1 mm diameter. Androgynous or mixed inflorescences 912 cm long; bracts
and bracteoles broadly ovate, 0.81 0.60.8 mm. Female flowers solitary along the
rachis; perianth lobes acute, c. 0.5 0.3 mm; staminodes 10; styles conical, straight,
c. 1 mm long. Cupules solitary along the rachis, sessile or with stalk 510 mm long,
saucer-shaped, 0.30.7 1.42.5 cm, densely tomentose with simple hairs, scaly; wall
woody, thin, enclosing less than half of the acorn; scale-like appendages distinct, appressed,
set in regular lines. Acorns depressed ovoid, 1.32 1.62 cm, most part exserted, glabrous
except for a small part at the apex which is densely tomentose with stellate and simple
hairs, purplish brown; base rounded, apex acute; scar concave or flat, 11.5 cm diameter;
Distribution. Thailand, Sumatra, Peninsular Malaysia, Singapore, Java, Borneo, and the
Philippines. Widely distributed in Sabah. In Sarawak, recorded for Kapit, Miri, Kuching,
and Sri Aman districts. Also in Brunei and Kalimantan.
Ecology. In primary and secondary mixed dipterocarp, kerangas, limestone, riparian to lower
montane forests, up to 1500 m altitude.
58. Lithocarpus tawaiensis S. Julia & Soepadmo

(of Bt. Tawai, Sabah)
Gard. Bull. Sing. 50 (1998) 148. Type: Berhaman et al. SAN 134267, Borneo, Sabah, Bt. Tawai (holotype
KEP).
Stunted treelet up to 1.5 m tall, 5 cm in diameter. Bark pale brown. Twigs sparsely appressed
tomentose, smooth. Stipules broadly ovate, 1013 1018 mm. Leaves sessile, thick90
vi
coriaceous, rigid, glabrous above, sparsely appressed yellowish tomentose or glabrescent

below; ovate to suborbicular, 6.510.5 69 cm, base cordate, margin revolute, apex acute
to acuminate; midrib raised on both surfaces, slightly stronger below; lateral veins thin, 10
12 pairs, dense, flat and almost invisible on both surfaces, faintly joining near the leaf margin, forming an angle of 3060 with the midrib; intercostal venation reticulate, dense, obscure above, slightly prominent below. Male inflorescences 615 cm long; bracts and
bracteoles ovate, 0.60.8 0.40.6 mm. Male flowers solitary or in clusters of 23 along the
rachis; perianth lobes obtuse to acute, c. 1.2 0.7 mm; stamen filaments c. 2 mm long;
pistillode globose, c. 0.5 mm in diameter. Pistillate inflorescences and flowers unknown.
Young cupule solitary along the rachis, sessile, saucer-shaped, 11.3 0.30.4 cm, scaly,
enclosing less than half of the acorn; scale-like appendages distinct, set irregularly. Acorns
exserted, most part free from the cupule.
Distribution. Endemic to Borneo. Twice collected from Bt. Tawai, Telupid, Sandakan district, Sabah.
Ecology. In hill forest on ultrabasic soils, at about 400 m altitude.
59. Lithocarpus turbinatus (Stapf) Forman
Fig. 14.
(Latin, turbinatus = shape like a top or inverted cone; the cupule and acorn)
Kew Bull. 18, 3 (1966) 423; Soepadmo l.c. (1970) 284, l.c. (1972) 329; Cockburn l.c. (1976) 100;
Anderson l.c. 203; Whitmore, Tantra & Sutisna l.c. 160. Basionym: Castanopsis turbinata Stapf, FMK
(1894) 232. Type: Haviland 1102, Borneo, Sabah, Mt. Kinabalu (holotype K).
Tree up to 30 m tall, 50 cm diameter. Bark flaky, reddish; inner bark granular, yellowish.
Sapwood yellowish. Twigs sturdy with thick nodes, densely tomentose, late-subglabrescent,
smooth or sparsely large-lenticellate. Stipules narrowly ovate to linear, c. 4 2 mm. Leaves
thick-coriaceous, rigid, glabrous above except the midrib, densely yellowish tomentose with
stellate-hairs below; ovate to elliptic, (6)812(13.5) 35(8) cm, base rounded or acute,
margin strongly revolute, apex acute to acuminate, acumen 1015 mm long; midrib flat
or impressed above, strongly raised below; lateral veins thick, 69 pairs, lax, impressed
or rarely flat above, strongly raised below, clearly joining near the leaf margin, forming
an angle of 3060 with the midrib; intercostal venation scalariform, lax or rarely dense,
obscure above, thin and clearly prominent below; petiole 813 mm long. Inflorescences
male, female, androgynous or mixed. Male inflorescences 57 cm long; bracts and bracteoles
ovate, 0.51 mm across. Male flowers solitary along the rachis; perianth lobes acute, c.
1.5 1 mm; stamen filaments c. 1.5 mm long; pistillode subglobose, c. 2 mm diameter.
Female, androgynous or mixed inflorescences sturdy, 46 cm long; bracts and bracteoles
ovate, c. 1 0.5 mm. Female flowers solitary or in clusters of 3 along the rachis; perianth
lobes acute, 1.52 mm across; staminodes 12; styles conical, straight, 2.53 mm long. Cupules
solitary along the rachis, sessile or with stalk c. 5 mm long, turbinate to ellipsoid, 5
7 3.55 cm, sparsely tomentose, lamellate; wall woody, c. 0.5 cm thick, sparsely hairy
and sometimes lenticellate, enclosing most part of the acorn except for an apical opening
0.51 cm diameter; lamellae distinct but not raised or rarely obscure, edge denticulate
especially the upper ones, set in regular 810 lines. Acorns ellipsoid, 56 45 cm, free
apical part sparsely tomentose, brown; base rounded to acute, apex obtuse to flat; wall
woody, thick, most part adnate to the cupule.
92
Distribution. Endemic to Borneo. Uncommon in Sabah and Sarawak. In Sabah, known

from Ranau (especially Kinabalu Park), Tenom, and Tambunan districts. In Sarawak, reported
for Bario, Ulu Sg. Kayan in Belaga district and G. Mulu NP in Miri district. Also in
Kalimantan.
Ecology. In primary and secondary hill mixed dipterocarp forest to montane mossy forest,
at 8002600 m altitude, on sandy and ultrabasic soils.
60. Lithocarpus urceolaris (Jack) Merr.

(Latin, urceolaris = pitcher-shaped; the cupule)
PROSEA l.c. 304; Coode et al. (eds.) l.c. 127; Argent et al. (eds.) l.c. 252. Basionym: Quercus urceolaris
Jack l.c. 86. Neotype (Merrill, 1952): Grashoff 951, Sumatra, Palembang, Banjumasin (holoneotype L;
isoneotype BO). Synonyms: Pasania urceolaris (Jack) Oerst. l.c. (1867) 83; Q. oligoneura Korth. l.c.
(1844) 203; P. oligoneura (Korth.) Oerst. l.c. (1867) 84; P. craterophora Fischer, Kew Bull. (1932) 319;
L. craterophora (Fischer) A. Camus l.c. (1935) 75.
Tree up to 35 m tall, 70 cm diameter. Bark fissured, lenticellate, pale reddish brown;

inner bark fibrous, brownish. Sapwood brownish. Twigs sparsely to densely tomentose,
sparsely or densely lenticellate. Stipules narrowly ovate to linear, 23 1 mm. Leaves
coriaceous, sparsely to densely tomentose or sometimes glabrous above, densely tomentose
with appressed stellate or sometimes mixed with stiff hairs below; elliptic, obovate, oblong
or lanceolate, 1216.5(35) 59(10) cm, base acute to cuneate, rarely rounded, margin
revolute, apex sharply acute to acuminate, acumen 1025 mm long; midrib broad, flat
or slightly raised above, strongly raised below; lateral veins thin, (6)712 pairs, lax,
flat or impressed or sometimes obscure above, raised below, disappearing near the leaf
margin, forming an angle of 4060 with the midrib; intercostal venation scalariform,
dense, obscure above, thin and prominent below; petiole 58 mm long. Inflorescences
male or androgynous. Male inflorescences 1520(25) cm long; bracts elliptic, c. 1
0.3 mm; bracteoles linear, c. 0.5 0.2 mm. Male flowers solitary along the rachis; perianth
lobes obtuse, c. 1 0.9 mm; stamen filaments 23 mm long; pistillode globose, c. 0.5
mm diameter. Androgynous inflorescences 1225 cm long; bracts ovate, c. 0.8 0.5
mm. Female flowers solitary along the rachis; perianth lobes acute, c. 1 0.6 mm;
staminodes 10; styles conical, recurved, 11.5 mm long. Cupules solitary or rarely in
clusters of 3 along the rachis, with stalk 58 mm long, cup-shaped, (0.5)1.53.5 2.5
3.5 cm, densely tomentose, minutely scaly or lamellate; wall woody, thick, densely hairy,
enclosing up to half of the acorn; lamellae obscure, or if scaly, the scale-like appendages
distinct, set in irregular lines. Acorns depressed globose, 24 23.5 cm, most part
exserted, densely tomentose, brownish; base flat, apex acute or rounded; scar concave,
11.5 cm diameter; wall woody, thick, most part free from the cupule.
Distribution. Sumatra, Peninsular Malaysia and Borneo. Common and widespread throughout
Ecology. In primary and secondary mixed dipterocarp, peatswamp, riparian, kerangas to
lower montane forests, up to 1200 m altitude, on various types of soils.
93
61. Lithocarpus wallichianus (Lindl. ex Hance) Rehder

(N. Wallich, 17861854, Danish botanist at Calcutta)
J. Arn. Arb. (1919) 132; Soepadmo l.c. (1970) 287, l.c. (1972) 368; Cockburn l.c. (1972) 227; Anderson
l.c. 203; Corner l.c. 342; Whitmore, Tantra & Sutisna l.c. 160; Turner l.c. 238; PROSEA l.c. 305;
Kochummen l.c. 249. Basionym: Quercus wallichiana Lindl. ex Hance l.c. (1870) 4. Type: Wallich
Cat. 2778, Peninsular Malaysia, Penang, Penang Hill (holotype K; isotypes BM, CGE). Synonyms:
Pasania wallichiana (Lindl. ex Hance) Gamble l.c. (1915) 425; Synaedrys wallichiana (Lindl. in
Wall.) Koidzumi l.c. 199; Q. hystrix Korth. var. longispica Gamble l.c. (1915) 429.
Tree up to 40 m tall, 90 cm diameter. Bark scaly, greyish; inner bark fibrous, greyish.
Sapwood yellowish or brownish. Twigs densely tomentose, late-glabrescent, fissured or
densely minute-lenticellate. Stipules linear, c. 3 1 mm. Leaves coriaceous, sometimes
rigid, sparsely appressed tomentose above, densely appressed tomentose with simple and
stellate-hairs below; elliptic to oblong, 8.515(19) 25(6) cm, base acute to cuneate,
margin revolute, apex acuminate, acumen 812 mm long; midrib strongly raised on both
surfaces; lateral veins thick, (8)1115(18) pairs, dense, impressed or flat above, strongly
raised below, clearly joining near the leaf margin, forming an angle of 4060 with the
midrib; intercostal venation subscalariform, dense, obscure above, strongly prominent
below; petiole 58 mm long. Inflorescences male, androgynous or mixed. Male inflorescences
612 cm long; bracts and bracteoles ovate to triangular, 1.21.8 1 mm. Male flowers
solitary along the rachis; perianth lobes acute, c. 1.2 1 mm; stamen filaments c. 2
mm long; pistillode subglobose, 11.5 mm diameter. Androgynous or mixed inflorescences
1017 cm long; bracts and bracteoles ovate, 1.52 1.2 mm. Female flowers in clusters
of 3 along the rachis; perianth lobes obtuse, c. 1 0.6 mm; staminodes 10; styles cylindrical,
straight, c. 1 mm long. Cupules in clusters of 23 and closely arranged along the rachis,
sessile, saucer-shaped, 0.30.5(0.7) 11.5 cm, densely tomentose with simple and stellate
hairs, scaly; wall woody, thin, enclosing less than half of the acorn; scale-like appendages
slightly distinct or obscure, appressed, set in irregular lines. Acorns ovoid, 11.7 cm
across, most part exserted, densely tomentose with simple hairs, brownish; base flat, apex
acute or rounded; scar deeply concave, c. 0.5 cm diameter; wall woody, thick, most part
Distribution. Thailand, Peninsular Malaysia, Singapore, and Borneo. In Borneo uncommon.
In Sabah, known only from Nabawan in Keningau district. In Sarawak, recorded for
Semengoh Arboretum and Mt. Santubong in Kuching district, and Ulu Sg. Kayan in Lundu
district. Also in Brunei.
Use. The acorns are edible but somewhat bitter.
3. QUERCUS L.
(Latin, quercus = the oak tree)
empili (Iban); empenit, mempening (Malay)
Gen. Pl., ed. 5 (1754) 431; Hooker f., Fl. Br. Ind. 5 (1888) 600; King, Ann. Roy. Bot. Gard. Calc. 2
94
(1889) 19; Ridley, FMP 3 (1924) 372; A. Camus, Chnes 1 (1938) 311; Burgess, TBS (1966) 289;
Soepadmo, Gard. Bull. Sing. 21 (1966) 379, ibid. 22 (1968) 355, FM 1, 7 (1972) 385; Cockburn,
TFM 1 (1972) 228, TS 1 (1976) 116; Anderson, CLST (1980) 203; Whitmore, Tantra & Sutisna, CLK
2, 1 (1990) 160; Kubitzki in Kubitzki et al. (eds.), Fam. & Gen. Vasc. Pl. 2 (1993) 308; PROSEA 15,
2 (1995) 409; Coode et al. (eds.), CLBD (1996) 127; Argent et al. (eds.), MNDT-CK 1 (1997) 252.
Synonyms: Cyclobalanopsis Oerst., Vid. Medd. Nat. For. Kjbn. 8 (1867) 77; Quercus L. subgen.
Cyclobalanopsis (Oerst.) Schneider, Ill. Handb. Laubh. 1 (1906) 210.
Trees; buttresses and stilt-roots sometimes present. Wood with narrow-uniseriate and broadmultiseriate rays. Bark smooth, flaky or cracked; inner bark of various colour, sometimes
fibrous, with ridges growing into the sapwood. Sapwood of various colour. Twigs initially
densely tomentose by simple or stellate hairs or densely brownish, stiff pubescent,
glabrescent, smooth, finely fissured or lenticellate. Terminal buds ovoid, globose, or conical,
crowded at the end of twigs. Stipules extrapetiolar, linear, acute, densely tomentose or
woolly pubescent, caducous. Leaves spirally arranged, crowded near the end of twigs
or rarely pseudowhorled; margin entire, plane or sometimes revolute, or remotely serrulate
in the apical half; petiole thickened at the base. Inflorescences male or female spikes,
unbranched or rarely branched. Male inflorescences pendent, unbranched, solitary in the
axils of lower (proximal) leaves, or in paniculate clusters on lateral new short shoots;
bracts and bracteoles densely tomentose or minutely hairy, caducous. Male flowers solitary
or in dichasial clusters of 34 along the rachis; perianth (4)6-lobed, densely tomentose,
stamens (4)6(9), anthers basifixed, filaments slender, filiform, glabrous or tomentose
at the base; pistillode absent or rarely replaced by a tuft of stiff simple hairs. Female
inflorescences solitary in the axils of upper (distal) leaves, erect; bracts densely pubescent,
caducous. Female flowers solitary or in pairs along the rachis; perianth (4)6(9)-lobed,
staminodes absent or rarely up to 6; ovary 3(6)-loculed, styles 3(6), cylindrical, recurved,
stigma broadly capitate or globose. Cupules cup- or saucer-shaped, each surrounding/
substending a single fruit, lamellate; edge of lamellae entire, undulate or denticulate,
free or connate, pubescent, tomentose or woolly pubescent on both surfaces. Fruits (acorns)
ovoid, conical, subglobose or cylindrical, rounded in cross-section, glabrous and shining
or densely tomentose, apex depressed, distinctly umbonate; umbo with many rings; base
flat, convex or concave; wall woody, its most part free from the cupule.
Distribution. About 400 species, widely distributed in temperate, subtropical and tropical
regions, mainly of the northern hemisphere. In Malesia, found in Sumatra, Peninsular
Malaysia, Singapore, Java, Borneo, and the Philippines (Palawan). Seventeen species are
recorded for Sabah and Sarawak, of which 4 are endemic.
Ecology. In lowland mixed dipterocarp to montane forest, from sea level up to 3350
m altitude, but more commonly found at 6001500 m altitude, on sandy clay, sandy loam
and ultrabasic soils overlying sandstone or granite.
Use. Source of medium hardwood timber.
Key to Quercus species

(for sterile specimens)
1. Petiole (1)35(10) mm long. Leaf apex rounded, obtuse, or emarginate.....2
Petiole (10)2040(50) mm long. Leaf apex acute, acuminate or caudate.....4
95
2. Leaves pseudowhorled, 617 36.5 cm; base rounded, cordate to auriculate; lateral
veins 815 pairs..................13. Q. pseudoverticillata
Leaves spirally arranged, 1.55.5 0.83.5 cm; base attenuate, acute, or obtuse; lateral
veins 48 pairs...............3
3. Leaves glabrous above, sparsely hairy below; margin entire, apex rounded, obtuse,
or emarginate; lateral veins 45(6) pairs; petiole up to 3 mm long, glabrous....
......2. Q. chrysotricha
Leaves initially densely pubescent on both surfaces, late-glabrescent; margin remotely
serrulate in the apical half, apex obtuse; lateral veins (5)68 pairs; petiole 26 mm
long, densely tomentose............10. Q. merrillii
4. Leaves (7)1018(25) (2)47(9) cm..............5
Leaves (4)510(16) 24(5.5) cm...............13
5. Leaf margin entire and revolute...........6
Leaf margin plane, remotely serrulate, serrate, or crenate in the apical half.......7
6. Older twigs sparsely tomentose with stellate hairs, shallowly fissured. Leaves glaucous
and sparsely tomentose beneath, lateral veins (6)710(12) pairs....11. Q. nivea
Older twigs glabrous, densely warty lenticellate. Leaves densely silvery tomentose
beneath, lateral veins (10)1218 pairs................1. Q. argentata
7. Older leaves glabrous on both surfaces or glabrous above, glabrescent beneath..8
Older leaves variously hairy at least beneath.9
8. Leaves thick-coriaceous; lateral veins 1015 pairs, closely spaced, strongly raised
beneath; intercostal venation dense, scalariform, prominent beneath..
...17. Q. valdinervosa
Leaves thin-coriaceous; lateral veins 710 pairs, distantly spaced, slightly raised
beneath; intercostal venation lax, reticulate, obscure on both surfaces or slightly raised
beneath.......5. Q. gemelliflora
9. Leaves densely greyish brown pubescent with appressed, straight simple hairs beneath;
lateral veins 1520 pairs, closely spaced...........8. Q. lineata
Leaves sparsely or densely tomentose with stellate hairs, or sparsely pubescent with
stiff simple hairs beneath; lateral veins (6)812(15) pairs, distantly spaced...............10
10. Leaves sparsely tomentose with stellate hairs and pale glaucous beneath; base
rounded.........12. Q. percoriacea
Leaves densely greyish brown tomentose, or sparsely tomentose with stellate hairs
or sparsely pubescent with stiff, simple hairs beneath; base attenuate, acute.........11
11. Leaf apex acuminate, acumen 1015 mm long; intercostal venation subscalariform.
....3. Q. elmeri
Leaf apex acute; intercostal venation scalariform...........12
12. Lower surface not glaucous, sparsely tomentose with brownish stellate hairs; intercostal
venation distinct below............4. Q. gaharuensis
96
Lower surface glaucous and sparsely pubescent with stiff, simple hairs; intercostal
venation obscure on both surfaces.............15. Q. sumatrana
13. Leaves linear to narrowly lanceolate, densely pubescent with simple minute hairs
beneath........16. Q. treubiana
Leaves elliptic, lanceolate or ovate, glabrous on both surfaces or glabrous above and
densely to sparsely tomentose with silky stellate hairs beneath......14
14. Petiole grooved on the adaxial side. Intercostal venation reticulate............15
Petiole flat on the adaxial side. Intercostal venation scalariform...16
15. Leaves thick-coriaceous, ovate to elliptic, base rounded or cordate, lower surface
densely tomentose with stellate hairs, late-glabrescent; intercostal venation distinct
above, obscure beneath.......................9. Q. lowii
Leaves thin-coriaceous, elliptic to lanceolate, base attenuate or acute, lower surface
sparsely tomentose with silky stellate hairs to glabrous; intercostal venation obscure
on both surfaces.......14. Q. subsericea
16. Leaves elliptic or lanceolate, glabrous on both surfaces, glaucous below; base attenuate
or acute, symmetrical; margin entire and somewhat undulate.............6. Q. kerangasensis
Leaves ovate, minutely pubescent with golden hairs and not glaucous beneath, lateglabrescent; base rounded to obtuse, asymmetrical; margin entire or remotely serrulate
in the apical half.............7. Q. kinabaluensis
Key to Quercus species

(for fruiting specimens with mature acorns and fully developed cupules)
1. Rim of fully developed cupule up to 0.5 mm thick; edge of lamellae free......2
Rim of fully developed cupule 14 mm thick; edge of lamellae tightly pressed to
the cupule wall or rarely free11
2. Cupule deeply cup-shaped, (1)1.53(3.5) cm deep, (1.5)23(4) cm diameter,
covering 1/33/4 part of the acorn...............................3
Cupule shallowly cup-shaped or flattish saucer-shaped, 0.51.5 cm deep, (0.7)1
1.8(3) cm diameter, covering 1/61/3 part of the acorn..........6
3. Cupule 23.5 cm deep, 2.54 cm diameter. Acorn 35 cm long, 23 cm diameter.
Leaves pseudowhorled; petiole 35 mm long...........13. Q. pseudoverticillata
Cupule 12.5 cm deep and 1.53.2 cm diameter. Acorn 23(3.5) cm long, 1.52(
3) cm diameter. Leaves spirally arranged; petiole 1050 mm long..4
4. Cupule rounded at base. Leaves sparsely tomentose with brownish stellate hairs and
neither glaucous nor silvery beneath; margin remotely serrulate in the apical
half..................4. Q. gaharuensis
Cupule tapering towards the base. Leaves densely or sparsely tomentose with stellate
hairs and glaucous or silvery beneath; margin entire.......5
5. Acorn conical, cylindrical or ovoid, 23 cm long, 1.52 cm diameter. Leaves densely
97
tomentose with silvery stellate hairs beneath..............1. Q. argentata

Acorn obovoid to subglobose, c. 2 cm across. Leaves glaucous and sparsely tomentose
with stellate hairs beneath......11. Q. nivea
6. Leaves 1.55.5 0.83.5 cm; petiole 16 mm long...........7
Leaves (3)1015(24) (1)35(9) cm; petiole 1035 mm long............8
7. Acorn densely tomentose. Leaves glabrous above, sparsely hairy below; margin entire,
apex rounded, obtuse, or emarginate; lateral veins 45(6) pairs..........2. Q. chrysotricha
Acorn glabrous. Leaves initially densely pubescent on both surfaces, late-glabrescent;
margin remotely serrulate in the apical half, apex obtuse; lateral veins (5)68
pairs...............10. Q. merrillii
8. Leaves chartaceous to thin-coriaceous; base attenuate or acute.............9
Leaves thick-coriaceous; base rounded or cordate....10
9. Cupule 1.73 cm diameter, base flat, lamellae 46. Acorn 22.5 cm diameter, base
flat to concave. Leaves glaucous and sparsely pubescent with simple hairs below.....
........15. Q. sumatrana
Cupule 0.71 cm diameter, base rounded or attenuate, lamellae 78. Acorn 11.5
cm diameter, base rounded. Leaves not glaucous but sparsely, tomentose with silky
stellate hairs beneath..........14. Q. subsericea
10. Cupule 1.31.8 cm diameter, covering 1/41/3 part of the acorn. Acorn 1.52 1.3
1.5 cm, apex obtuse. Leaves (3)610(14) (1)24(5.5) cm, densely tomentose
with stellate hairs and not glaucous beneath; lateral veins 58 pairs.9. Q. lowii
Cupule 22.3 cm diameter, covering 1/61/5 part of the acorn. Acorn 22.5 1.5
2.5 cm, apex depressed, rounded. Leaves (10)1618(20) (3.5)48(9) cm,
glaucous and sparsely tomentose with stellate hairs beneath; lateral veins 1012
pairs............................................................12. Q. percoriacea
11. Cupule 12 cm diameter. Acorn 12(3) 0.81.5 cm.....12
Cupule 24 cm diameter. Acorn 25.5 13 cm...........13
12. Leaves elliptic or lanceolate, glabrous on both surfaces, glaucous below; base attenuate
or acute, symmetrical, margin entire and somewhat undulate...........6. Q. kerangasensis
Leaves ovate, minutely pubescent with golden hairs and not glaucous below; base
rounded to obtuse, asymmetrical; margin entire or remotely serrulate in the apical
half.................7. Q. kinabaluensis
13. Older leaves glabrous. Acorn up to 5.5 cm long.............14
Older leaves densely hairy beneath. Acorn up to 3.2 cm long15
14. Leaf lateral veins 1015 pairs, closely spaced, strongly prominent below; intercostal
venation dense and prominent below.........17. Q. valdinervosa
Leaf lateral veins 710 pairs, distantly spaced, only slightly prominent below; intercostal
venation lax, obscure on both surfaces or only slightly prominent beneath..
......5. Q. gemelliflora
98
15. Leaf lateral veins 1520 pairs, closely spaced; intercostal venation densely scalariform,
prominent beneath....8. Q. lineata
Leaf lateral veins 512 pairs, distantly spaced; intercostal venation fine, scalariform,
reticulate, or subscalariform, obscure on both surfaces or distinct beneath and obscure
above.......16
16. Cupule cup-shaped, 11.5 23 cm; lamellae 1012. Acorn with convex base. Leaves
linear to narrowly lanceolate, (3)710(12) 13 cm; intercostal venation fine,
scalariform.......16. Q. treubiana
Cupule saucer-shaped, 0.51.3 22.5 cm; lamellae 47. Acorn with truncate base.
Leaves elliptic, oblong or lanceolate, (5)813(14) (1)1.54(5) cm; intercostal
venation subscalariform..............3. Q. elmeri
1. Quercus argentata Korth.

(Latin, argentatus = silvered; the silvery indumentum of the lower leaf surface)
Kruidk. (1844) 215; King l.c. 30; Gamble, J. As. Soc. Beng. 75, 2 (1915) 408, p.p.; Merrill, EB (1921)
211; Ridley l.c. 373, p.p.; Camus l.c. (1938) 311; Masamune, EPB (1942) 233; Soepadmo l.c. (1968)
384, l.c. (1972) 389; Cockburn l.c. (1972) 228, l.c. (1976) 118; Anderson l.c. 203; Whitmore, Tantra &
Sutisna l.c.160; Turner, Gard. Bull. Sing. 47 (1995) 238; PROSEA l.c. 413; Argent et al. (eds.) l.c. 252.
Type: Korthals s.n. (= RHL Sheet No. 901309448), Sumatra, Melintang Forest (holotype L; isotype K).
Synonyms: Cyclobalanopsis argentata (Korth.) Oerst. l.c. 79; Lithocarpus argentata (Korth.) Merr.,
Contr. Arn. Arb. 8 (1934) 42; Q. argentata Korth. var. concolor Blume, Mus. Bot. Lugd.-Bat. 1 (1850)
299; Q. pinanga Blume l.c. (1850) 303; Q. wilhelminae Seemen, Bull. Dpt. Agr. Ind. Nerl. 1 (1906) 3;
Merrill l.c. (1921) 216; Masamune l.c. 237.
Tree up to 40 m tall, 100 cm diameter; buttresses up to 1.5 m tall. Bark smooth, lenticellate,
pale grey; inner bark brittle, pale brown. Sapwood yellowish. Twigs glabrous, densely
warty lenticellate, greyish brown. Terminal buds ovoid to globose, 23 22.5 mm,
stellate- or simple-tomentose, glabrescent. Stipules hairy, linear, 35 11.5 mm. Leaves
spirally arranged, coriaceous, glabrous and glossy above, tomentose with dense silvery
stellate hairs below, glabrescent; elliptic, oblong, lanceolate or oblanceolate, (7)1018
(25) (2)47(9) cm, base attenuate, rounded or acute, slightly asymmetrical, margin
entire, revolute, apex sharply acute to acuminate, acumen 515 mm long; midrib prominent
below, impressed above; lateral veins (10)1218 pairs, parallel, arching towards the
leaf margin, forming an angle of 5075 with the midrib; intercostal venation scalariform,
obscure on both surfaces; petiole slender, black at the base, 1040 mm long, adaxially
flat, grooved or sulcate, glabrous. Male inflorescences 510 cm long; bracts ovate, 1.5
2 0.51 mm, densely hairy. Male flowers: perianth 5-lobed, lobes c. 1 1.5 mm,
connate at the base, hairy, stamens 6(8), filaments hairy at the base. Female inflorescences
many-flowered, 23 cm long, densely hairy; bracts and bracteoles linear, acute, 0.51
mm long. Female flowers: perianth 46-lobed, densely tomentose outside; ovary conical,
0.81 mm across, styles 34, pubescent with long golden-brown hairs at the base. Cupules
deeply cup-shaped, 0.81.5 1.52 cm, tapering towards the base into a 0.30.5 cm
long stalk, covering up to 1/3 of the acorn; rim thinner than 0.5 mm; lamellae 812,
edge free, denticulate. Acorns conical, cylindrical, or ovoid, 23 1.52 cm, dark
chocolate-brown when mature, glabrous; apex rounded or acute, base rounded.
99
Vernacular names. Sarawakempili (Iban), mempening (Malay).

Distribution. Sumatra, Peninsular Malaysia, Singapore, Java, and Borneo. In Sabah and
Sarawak scattered. Also in Kalimantan.
Ecology. In mixed dipterocarp forest to montane forest, at altitudes from sea level to
2500 m. Flowering in JulySeptember; fruiting between October and May.
Use. Important source of mempening timber.
2. Quercus chrysotricha A. Camus

(Greek, chrysotrichos = golden-hairy; young twigs and inflorescences)
Chnes 3, Expl. Pl. 5 (1949) 50, t. 347, ibid. Text (1954) 1212; Soepadmo l.c. (1968) 397, l.c. (1972)
394; Anderson l.c. 203; Whitmore, Tantra & Sutisna l.c. 160. Type: Richards 1885, Borneo, Sarawak,
Mt. Dulit, alt. c. 1300 m (holotype K; isotype L). Synonym: Q. arbutifolia (non Hickel & A. Camus)
E.F. Warb., Kew Bull. (1936) 19.
Tree up to 20 m tall, 30 cm diameter; buttresses up to 1 m tall. Bark smooth, finely

fissured or hoop-marked, brown mottled with greyish green patches. Young twigs initially
densely covered with golden, woolly, simple hairs, late-glabrescent, lenticellate. Terminal
buds ovoid to subglobose, c. 3 2 mm. Stipules linear, 810 1 mm, densely woolly
rufous-tomentose with simple hairs. Leaves spirally arranged, thin-coriaceous, glabrous
above, sparsely pubescent below; ovate, elliptic or obovate, 2.55.5 1.53.5 cm, base
attenuate, acute or obtuse, asymmetrical, margin entire, revolute, apex rounded, obtuse
or emarginate; midrib raised below, flat or impressed above; lateral veins 45(6) pairs,
prominent below, flat above, parallel, arching towards the leaf margin, forming an angle
of 6070 with the midrib; intercostal venation fine, scalariform, obscure on both surfaces;
petiole (1)23 mm long, adaxially flat, glabrous. Male inflorescences up to 2 cm long,
few-flowered, densely woolly pubescent with golden-brown hairs; bracts ovate, acute.
Male flowers in dichasial clusters of 3 along the rachis; perianth 5-lobed; stamens 5,
filaments glabrous. Female inflorescences up to 1 cm long, densely woolly pubescent
with golden-brown hairs. Female flowers: perianth 5-lobed, densely woolly pubescent
with golden-brown hairs; ovary conical, c. 1 0.6 mm, pubescent. Cupules shallowly
cup-shaped, 11.5 cm across, covering 1/41/3 of the acorn, base attenuate, pubescent
on both surfaces; rim up to 0.5 mm thick; lamellae (4)5(7), edge free, denticulate. Acorns
ovoid or conical, 11.5 cm across, chocolate-brown, densely tomentose, glabrescent; apex
attenuate, acute, base rounded.
Vernacular name. Sarawakmempening (Malay).
Distribution. Endemic to Borneo. In Sabah uncommon, in Sarawak scattered. Also in
Kalimantan.
Ecology. In kerangas to lower montane forests, at 12001300 m altitude.
100
3. Quercus elmeri Merr.

(A.D.E. Elmer, 18701942, plant collector in Borneo and the Philippines)
Univ. Calif. Publ. Bot. 15 (1925) 43; Soepadmo l.c. (1968) 414, l.c. (1972) 398; Cockburn l.c. (1972)
229, l.c. (1976) 117; Anderson l.c. 203; Whitmore, Tantra & Sutisna l.c. 160; PROSEA l.c. 414; Turner
l.c. 238. Type: Elmer 21213, Borneo, Sabah, Tawau district (holotype K; isotypes A, L, U).
Tree up to 40 m tall, 90 cm diameter; buttresses narrow, up to 3 m tall. Bark scaly,

lenticellate, greyish brown; inner bark fibrous, reddish brown. Sapwood ochre. Twigs
initially densely grey-brown pubescent, glabrescent, densely lenticellate. Terminal buds
ovoid to subglobose, c. 35 mm across, densely tomentose. Stipules linear, acute, 5
7 12 mm, pubescent. Leaves spirally arranged, coriaceous, upper surface glabrous,
lower surface densely tomentose with greyish brown stellate hairs, late-glabrescent; elliptic,
lanceolate to oblong, (5)813(14) (1)1.54(5) cm, base attenuate, acute, usually
asymmetrical, margin plane, remotely serrulate in the upper half, apex acuminate, acumen
1015 mm long; midrib prominent below, flat or slightly raised above; lateral veins (6)
812 pairs, parallel, arching towards the leaf margin, distantly spaced, forming an angle of
6070 with the midrib; intercostal venation fine, subscalariform, distinct beneath, obscure
above; petiole 1030 mm long, terete or flat on the adaxial side, densely pubescent,
glabrescent. Male inflorescences 13 cm long; bracts ovate, acute, densely pubescent.
Male flowers: perianth 34-lobed, densely pubescent; stamens 45, filaments pubescent
at the base. Female inflorescences 34-flowered, 12 cm long, densely hairy; bracts ovate,
acute. Female flowers: ovary conical, c. 1 0.8 mm, styles 34, recurved, 23 mm
long, densely hairy at the base. Cupules saucer-shaped, 0.51.3 22.5 cm, base truncate
or rounded, covering 1/41/3 of the acorn, densely pubescent on both surfaces; rim 1
2 mm thick; lamellae 47, edge tightly pressed towards cupule wall, denticulate. Acorns
densely sericeous, glabrescent, ovoid, conical to cylindrical, 23.2 1.52 cm; apex
depressed, rounded, base truncate.
Vernacular name. Sabah and Sarawakmempening (Malay).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah, known from Bt. Kalong,
Mt. Kinabalu, and Tawau. In Sarawak scattered. Also in Kalimantan.
Ecology. In lower montane forest, up to 1400 m altitude, on sandy loam and ultrabasic
soils. Fruiting between September and April.
4. Quercus gaharuensis Soepadmo

(of G. Gaharu, Sarawak)
Gard. Bull. Sing. 21 (1966) 384, l.c. (1968) 389, l.c. (1972) 391; Cockburn l.c. (1972) 229; Anderson
l.c. 203; Whitmore, Tantra & Sutisna l.c. 160; PROSEA l.c. 414; Turner l.c. 238; Type: Anderson et al.
S. 15534, Borneo, Sarawak, Serian, lower slope of G. Gaharu, alt. c. 100 m (holotype K; isotypes BO, L,
SAN, SAR).
Tree up to 30 m tall, 70 cm diameter; buttresses low, spreading, up to 0.7 m high. Bark
101
smooth to slightly fissured, greyish brown. Twigs initially densely stellate-tomentose,

glabrescent, lenticellate. Terminal buds ovoid to subglobose, c. 2 mm across. Stipules
linear, acute, c. 5 1 mm. Leaves spirally arranged, thin-coriaceous, glabrous above,
not glaucous but sparsely tomentose with brownish stellate hairs below; elliptic, lanceolate
to oblong, (5)1020(23) (2)46 cm, base attenuate, acute or rarely rounded, slightly
asymmetrical, margin plane, remotely serrulate in the apical half, apex acute; midrib
prominent and sparsely hairy with stellate hairs below; lateral veins 815 pairs, distantly
spaced, parallel, arching towards the leaf margin, forming an angle of 4560 with the
midrib; intercostal venation scalariform, distinct below; petiole 1535 mm long, adaxially
flat, initially densely stellate-tomentose, glabrescent. Male inflorescences up to 3.5 cm
long; bracts ovate, acute, 11.5 1 mm, caducous, hairy. Male flowers in dichasial clusters
of 3 along the rachis; perianth 45-lobed, hairy; stamens 46, filaments hairy only at
the base; pistillode replaced by a tuft of stiff hairs. Female inflorescences up to 1.5
cm long, 24-flowered; bracts ovate to linear, 12 0.51 mm, densely hairy. Female
flowers: perianth 56-lobed; ovary conical, c. 0.8 1 mm, styles 34, cylindrical, 1
2 mm long, densely hairy at the base. Cupules deeply cup-shaped, 11.5 2.53 cm,
covering 1/22/3 of the acorn, base rounded, woolly pubescent on both sides, glabrescent;
rim up to 0.5 mm thick; lamellae 68, edge free, slightly recurved and denticulate. Acorns
ovoid to conical, 23 22.5 cm, densely tomentose, glabrescent; apex obtuse, base
flat or convex.
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah, uncommon, known
only from Sipitang district. In Sarawak scattered. Possibly also in Kalimantan (record
doubtful).
Ecology. In hill mixed dipterocarp forest to lower montane forest, up to 1400 m altitude.
Fruiting between September and February.
5. Quercus gemelliflora Blume

(Latin, gemellus = twin-borne; floris = flower; the paired female flowers along the rachis)
Verh. Bat. Gen. K. & W. 9 (1823) 222; A. Camus l.c. (1938) 363; Backer & Bakhuizen f., FJ 2 (1965) 5;
Soepadmo l.c. (1968) 406, l.c. (1972) 395; Cockburn l.c. (1972) 229, l.c. (1976) 118; Whitmore, Tantra
& Sutisna l.c. 161; PROSEA l.c. 414; Turner l.c. 238. Type: Blume s.n. (= RHL Sheet No. 901310215),
Java, Mt. Salak (holotype L). Synonyms: Q. turbinata Blume, Bijdr. Fl. Ned. Ind. (1826) 523; Q. merkusii
Endlicher, Gen. Pl., Suppl. 4, 2 (1847) 28; Q. horsfieldii Miq., Fl. Ned. Ind. 1, 1 (1856) 856;
Cyclobalanopsis gemelliflora (Blume) Oerst. l.c. (1867) 77; Cyclobalanopsis horsfieldii (Miq.) Oerst.
l.c. (1867) 78; Cyclobalanopsis merkusii (Endl.) Oerst. l.c. (1867) 79; Q. lineata Blume var. merkusii
(Endl.) Wenzig, Jahrb. Kn. Bot. Gart. Berlin 4 (1886) 232; Cyclobalanopsis turbinata (Blume) Schottky,
Bot. Jahrb. 47 (1912) 648; Q. turbinata Blume var. crassilamellata Gamble, J. As. Soc. Beng. 75, 2
(1915) 410; Q. crassilamellata (Gamble) A. Camus, Bull. Mus. Hist. Paris 2, 3 (1931) 689.
Tree up to 30 m tall, 60 cm diameter. Bark smooth or cracked, greyish brown; inner

bark orange. Sapwood white. Twigs initially densely tomentose, glabrescent. Terminal
buds ovoid to subglobose. Stipules linear, acute, 510 12 mm, hairy. Leaves spirally
102
arranged, thin-coriaceous, glabrous above, initially densely pubescent below, soon

glabrescent; obovate, elliptic, lanceolate, or oblanceolate, (5)915(24) (2)46(7)
cm, base attenuate or acute, often asymmetrical, margin plane, serrulate in the apical
half, apex acute, acuminate, or caudate, acumen 515 mm long; midrib prominent below,
flat or slightly raised above; lateral veins 710 pairs, distantly spaced, slightly raised
beneath, parallel, arching towards the leaf margin, forming an angle of 4560 with the
midrib; intercostal venation reticulate, lax, obscure on both surfaces or slightly distinct
beneath; petiole (10)2030(40) mm long, adaxially flat, densely tomentose, glabrescent.
Male inflorescences many-flowered, crowded mostly on young shoots, 36 cm long; bracts
ovate, 23 11.5 mm, hairy. Male flowers solitary or in dichasial clusters of 3 along
the rachis; perianth 36-lobed, lobes c. 1 mm across, densely hairy; stamens 36. Female
inflorescences 27-flowered, 24 cm long, densely hairy; bracts ovate, acute, 12 1
mm. Female flowers usually in pairs along the rachis; perianth 46-lobed, densely hairy;
staminodes 06; ovary ovoid, c. 1 mm across, styles 46, 12 mm long, hairy at the
base. Cupules usually in pairs along the rachis, cup-shaped, 1.52.5 1.72.5 cm, covering
1/31/2 of the acorn, base attenuate to rounded, densely woolly pubescent inside, densely
tomentose with short brown hairs outside; rim c. 2 mm thick; lamellae 68, edge tightly
pressed towards cupule wall, denticulate. Acorns ovoid, conical or cylindrical, 25.5
22.5 cm, densely tomentose, glabrescent; apex obtuse, base rounded.
Distribution. Sumatra, Peninsular Malaysia, Borneo, and Java. In Sabah and Sarawak
scattered. Also in Kalimantan.
Ecology. In hill mixed dipterocarp forest to montane forest, up to 2600 m altitude, usually
near streams, on red sandy clay overlying sandstone or granite or on ultrabasic soils.
Flowering and fruiting the whole year round.
6. Quercus kerangasensis Soepadmo

(Iban, kerangas = a type of soil which is not suitable for cultivating rice; referring to its natural
habitat)
Gard. Bull. Sing. 21 (1966) 399, l.c. (1968) 396, l.c. (1972) 395; Anderson l.c. 204; Whitmore, Tantra &
Sutisna l.c. 161; Coode et al. (eds.) l.c. 127. Type: Brunig S. 1065, Borneo, Brunei, Seria, Badas FR
(holotype K; isotypes L, SAR).
Tree up to 30 m tall, 50 cm diameter; buttresses low. Bark greyish, smooth or lenticellate.

Twigs initially densely greyish brown pubescent with stiff hairs, glabrescent, densely warty
lenticellate. Terminal buds ovoid to subglobose, 36 24 mm, densely pubescent. Stipules
linear, acute, 23 0.51 mm, densely pubescent, glabrescent, caducous. Leaves spirally
arranged, coriaceous, glabrous on both surfaces, glaucous below; elliptic or lanceolate,
(4)510(12) 24(5) cm, base attenuate or acute, symmetrical, margin entire and
somewhat undulate, apex acute to acuminate, acumen 515 mm long; midrib prominent
below, flattish above; lateral veins fine on both surfaces, 710 pairs, subparallel, arching
towards the leaf margin, forming an angle of 6065o with the midrib; intercostal venation
103
fine, scalariform, obscure on both surfaces; petiole slender, 1012 mm long, flat on the
adaxial side. Male inflorescences 12 cm long, few-flowered; bracts ovate, acute, c.
1 mm across, membranous, pubescent outside. Male flowers in dichasial clusters of 3
along the densely tomentose rachis; perianth 56-lobed, membranous, densely pubescent;
stamens 56, filaments up to 2 mm long. Female inflorescences unknown. Cupules cupshaped, obconical, 0.81.2 1.52 cm, covering up to 1/3 of the acorn, base attenuate,
densely brownish tomentose on both sides; rim 11.5 mm thick; lamellae 58, edge tightly
pressed towards cupule wall, denticulate or smooth. Acorns ovoid, conical or cylindrical,
densely sericeous with short brown hairs, glabrescent, shining, chocolate-brown, 23
1.21.5 cm; apex rounded or acute, base strongly convex.
Vernacular names. Sabahmempening (Malay). Sarawakempenit (Iban), mempening
(Malay).
Distribution. Endemic to Borneo. No record for Sabah. In Sarawak recorded only from
Ulu Baram in Marudi district. Also in Brunei. The correct identity of Quercus kerangensis
as reported for C Kalimantan (Argent et al. (eds.) l.c. 255) is doubtful.
Ecology. In lowland kerangas forest, up to 200 m altitude. Flowering in June; fruiting
in September.
7. Quercus kinabaluensis Soepadmo

Gard. Bull. Sing. 21 (1968) 401, l.c. (1972) 395; Anderson l.c. 204; Cockburn l.c. (1976) 117;
Whitmore, Tantra & Sutisna l.c. 161. Type: Chew & Corner RSNB 7148, Borneo, Sabah, Mt. Kinabalu,
Mentaki Ridges (holotype K; isotype L).
Tree up to 35 m tall, 70 cm diameter. Bark greyish, smooth or slightly cracked; inner

bark light brown. Sapwood yellowish. Twigs initially densely brownish pubescent with
stiff simple hairs, glabrescent, sparsely lenticellate. Terminal buds ovoid to subglobose,
410 34 mm. Stipules linear, acute, 510 0.51 mm, caducous. Leaves spirally
arranged, thick-coriaceous, glabrous above, minutely pubescent with golden hairs beneath,
late-glabrescent; ovate, 510 24 cm, base rounded to obtuse, often asymmetrical,
margin entire or remotely serrulate in the apical half, slightly revolute, apex acute to
acuminate, acumen up to 15 mm long; midrib slightly prominent below, flat above; lateral
veins 69 pairs, parallel, arching towards the leaf margin, forming an angle of 4560
with the midrib; intercostal venation fine, scalariform, obscure on both surfaces; petiole
1015 mm long, adaxially flat, densely brownish pubescent, glabrescent. Male inflorescences
densely yellowish brown pubescent, 510 cm long; bracts ovate, acute, 23 12 mm,
membranous, densely pubescent outside. Male flowers in dichasial clusters of threes or
solitary along the rachis; perianth 47-lobed, pubescent outside, glabrescent; stamens 4
7, filaments c. 2 mm long. Female inflorescences 13 cm long, 13-flowered, densely
hairy. Female flowers: perianth 46-lobed, densely hairy; ovary ovoid, c. 1 0.6 mm,
styles 35, hairy at the base, recurved, 23 mm long. Cupules obconical, cup-shaped,
12 12 cm, covering up to 1/2 of the acorn, base attenuate, pubescent on both surfaces;
rim c. 1 mm thick; lamellae 68, edge tightly pressed towards cupule wall, smooth to
104
denticulate. Acorns ovoid to conical, densely brown tomentose, glabrescent, 11.5 0.8
1 cm; apex rounded or attenuate, base convex.
Vernacular names. Sabahmempening (Malay). Sarawakempeli (Iban).
Distribution. Endemic to Borneo, uncommon. In Sabah only recorded for Mt. Kinabalu
area, and in Sarawak from G. Pueh in Lundu district and G. Murud in Limbang district.
Not known in Brunei and Kalimantan.
Ecology. In hill mixed dipterocarp forest to montane forest, at 6002600 m altitude, mainly
on ultrabasic soils. Flowering in February; fruiting between March and May.
8. Quercus lineata Blume

(Latin, lineatus = marked by parallel, fine lines; referring to the closely spaced, parallel
lateral veins)
Bijdr. Fl. Ned. Ind. (1825) 523; A. Camus l.c. (1938) 207; Backer & Bakhuizen f. l.c. 5; Soepadmo
l.c. (1968) 408, l.c. (1972) 396; Cockburn l.c. (1972) 230, l.c. (1976) 117; Whitmore, Tantra & Sutisna
l.c. 161; Turner l.c. 238; PROSEA l.c. 415. Type: Blume s.n. (= RHL Sheet No. 90131161), Java
(holotype L; isotypes A, BO, K). Synonyms: Q. polyneura Miq., Pl. Jungh. 1 (1851) 11; Q. lineata
Blume var. heterochroa Miq. l.c. (1856) 855; Q. oxyrhyncha Miq. l.c. (1860) 347; Cyclobalanopsis
lineata (Blume) Oerst. l.c. 78; Q. lineata Blume var. typica King, Ann. Roy. Bot. Gard. Calc. 2 (1889)
33; Q. lineata Blume var. oxyrhyncha (Miq.) Seemen l.c. 4; Q. hendersoniana A. Camus, Bull. Mus.
Paris, Ser. I, 4 (1932) 123.
Tree up to 30 m tall, 60 cm diameter. Bark smooth, greyish; inner bark fibrous, reddish
brown. Sapwood pale, white. Young twigs and buds densely pubescent with golden brown
hairs, glabrescent, lenticellate. Terminal buds ovoid to subglobose, 1.52.5 mm across.
Stipules linear, acute, 1015 12 mm, densely pubescent, caducous. Leaves spirally
arranged, thin-coriaceous, glabrous to sparsely pubescent above, densely greyish brown
tomentose with appressed simple hairs below; elliptic or lanceolate, (5)1520(23)
26 cm, base attenuate, acute, sometimes asymmetrical, margin plane, remotely serrulate
in the apical half, apex acuminate, acumen 1015 mm long; midrib prominent below,
flat or slightly raised above; lateral veins 1520 pairs, closely spaced, straight and parallel
towards the leaf margin, forming an angle of 4560 with the midrib; intercostal venation
densely scalariform, distinct below; petiole 1525 mm long, adaxially flat or shallowly
furrowed, sparsely tomentose. Male inflorescences 510 cm long; bracts ovate, acute,
23 2 mm, membranous, densely sericeous outside, glabrescent. Male flowers solitary
or in dichasial clusters of three along the rachis; perianth 56-lobed, densely tomentose
outside; stamens 6(8), filaments up to 1 mm long, glabrous; pistillode replaced by a
clusters of stiff, simple hairs. Female inflorescences 56-flowered, 1.52 cm long, densely
hairy; bracts ovate, acute, 2 1 mm, densely tomentose. Female flowers: perianth 5
6-lobed, hairy; ovary conical, c. 1.3 1 mm, styles 3, 13 mm long, recurved, sparsely
pubescent at the base. Cupules cup-shaped, obconical, 11.2 22.5 cm, covering c.
1/3 of the acorn, base attenuate or rounded, pubescent outside; rim 12 mm thick; lamellae
510, edge tightly pressed towards cupule wall, denticulate. Acorns ovoid, subglobose
to cylindrical, 23 13 cm, densely tomentose, base rounded; apex attenuate to rounded.
105

Distribution. Sumatra, Peninsular Malaysia, Java, and Borneo. In Sabah known mainly
from Mt. Kinabalu, and in Sarawak from G. Pueh and G. Berumput in Lundu district.
Ecology. In lower to upper montane forest, at 10002000 m altitude, on yellow sandy
or ultrabasic soils. Flowering in April to May; fruiting between August and April.
9. Quercus lowii King
Fig. 15.
(Sir Hugh Low, 18241905; officer of the British East India Company; plant collector)
Ann. Roy. Bot. Gard. Calc. 2 (1889) 28; Merrill l.c. (1921) 214; A. Camus l.c. (1938) 366; Masamune
l.c. 235; Soepadmo l.c. (1968) 394, l.c. (1972) 393; Cockburn l.c. (1976) 117; Whitmore, Tantra &
Sutisna l.c. 161. Type: Low s.n., Borneo, Sabah, Mt. Kinabalu (holotype CAL, n.v.). Synonym:
Cyclobalanopsis lowii (King) Schottky, Bot. Jahrb. 47 (1912) 653.
Tree up to 20 m tall, 40 cm diameter, occasionally with short buttresses. Bark rough;

inner bark pinkish to brownish, fibrous. Sapwood pale ochre to yellow. Twigs dark grey,
shallowly fissured, initially densely pubescent with stellate hairs, late-glabrescent, lenticellate.
Terminal buds ovoid to subglobose, densely tomentose. Stipules linear, acute, 24 0.3
0.5 mm. Leaves spirally arranged, thick-coriaceous, upper surface glabrous and glossy
or dull with sparse stellate hairs on the midrib, lower surface densely tomentose with
brownish stellate hairs, late-glabrescent; ovate to elliptic, (3)610(14) (1)24(5.5)
cm, base rounded or cordate, sometimes asymmetrical, margin entire or remotely serrulate
in the upper half, apex acute to acuminate, acumen (5)1015(20) mm long; midrib
prominent or flattish below, impressed above; lateral veins 58 pairs, parallel, arching
towards leaf margin, forming an angle of 5060 with the midrib; intercostal venation
fine, reticulate, distinct above, obscure below; petiole 1025 mm long, adaxially grooved,
densely tomentose with simple and stellate hairs, glabrescent. Male inflorescences slender,
510 cm long, densely tomentose with stellate hairs, unbranched or rarely branched; bracts
ovate, acute, densely hairy, 12 0.5 mm. Male flowers solitary or in dichasial clusters
of three along the rachis; perianth 56-lobed, densely tomentose, glabrescent; stamens
46, filaments up to 1 mm long; pistillode replaced by fine woolly hairs. Female
inflorescences 25-flowered, 12 cm long, densely hairy; bracts ovate, acute, c. 1 mm
across. Female flowers: perianth (4)56(7)-lobed, densely hairy; ovary conical, c. 1
0.7 mm, styles 36, hairy at the base. Cupules shallowly cup-shaped, 0.81 1.3
1.8 cm, covering 1/41/3 of the acorn, base rounded, densely tomentose on both sides;
rim less than 0.5 mm thick; lamellae 45, edge free, denticulate. Acorns conical, ovoid
to cylindrical, 1.52 1.31.5 cm, densely tomentose with stellate hairs, glabrescent;
apex obtuse, base convex.
Distribution. Endemic to Borneo; so far known from Sabah (mainly on Mt. Kinabalu),
Sarawak and Kalimantan (Mt. Lianggagang).
106
vi
Ecology. In hill mixed dipterocarp forest to montane forest, up to 2500 m altitude, mainly
on blackish ultrabasic soils. Flowering in JuneJuly; fruiting between August and March.
Use. The timber is source of medium hardwood in Sabah.
10. Quercus merrillii Seemen

(E.D. Merrill, American botanist; 19191923 Director of the Bureau of Science in Manila,
19301935 Director of the New York Botanical Garden, U.S.A.)
Fedde Rep. 5 (1908) 21; Merrill, Philip. J. Sc. 3 (1908) Bot. 329; Soepadmo l.c. (1968) 397, l.c.
(1972) 394; Cockburn l.c. (1976) 117; Anderson l.c. 204; Whitmore, Tantra & Sutisna l.c. 161. Type:
Foxworthy 566, the Philippines, Palawan (holotype K). Synonym: Cyclobalanopsis merrillii (Seemen)
Schottky, Bot. Jahrb. 47 (1912) 649.
Tree up to 10 m tall, 20 cm diameter. Bark slightly flaky, greyish brown. Twigs densely
hairy with brownish hairs, glabrescent, lenticellate. Terminal buds ovoid or conical c.
3 2 mm. Stipules linear, acute, 0.51 0.1 mm, caducous. Leaves spirally arranged,
thin-coriaceous, initially densely pubescent on both surfaces, late-glabrescent; obovate,
ovate, or elliptic, (1.5)34(5.5) 0.82.5 cm, base attenuate or acute, margin remotely
serrulate in the apical half, apex obtuse; midrib prominent beneath, impressed to flat
above; lateral veins (5)68 pairs, parallel, arching towards the leaf margin, forming
an angle of 5060 with the midrib; intercostal venation fine, scalariform, obscure on
both surfaces; petiole 26 mm long, adaxially flat, densely tomentose, glabrescent. Male
inflorescences 34 cm long, densely woolly pubescent with golden brown hairs; bracts
ovate. Male flowers solitary along the rachis; perianth 5-lobed, pubescent with soft long
simple golden hairs; stamens 67, filament glabrous. Female inflorescences 14-flowered,
12 cm long, densely hairy; bracts ovate, acute. Female flowers densely hairy; perianth
45-lobed; ovary conical, c. 1 0.7 mm, styles 4, recurved, densely pubescent at the
base. Cupules shallowly cup-shaped, 0.71.4 1.51.8 cm, covering 1/41/3 of the acorn,
pubescent on both surfaces; rim up to 0.5 mm thick; lamellae 48, edge free, finely
denticulate. Acorns ovoid, conical or cylindrical, 22.5(3) 11.5 cm, glabrous; apex
acute, base rounded.
Vernacular names. Sabahmempening (Malay). Sarawakempili (Iban).
Distribution. Borneo and Palawan in the Philippines. In Sabah and Sarawak uncommon.
Ecology. In hill mixed dipterocarp forest, up to 500 m altitude. Fruiting between April
and August.
11. Quercus nivea King

(Latin, niveus = snow white; the leaf undersurface)
Ann. Roy. Bot. Gard. Calc. 2 (1889) 31; Merrill l.c. (1921) 214; A. Camus l.c. (1938) 313; Soepadmo
l.c. (1968) 387, l.c. (1972) 391; Cockburn l.c. (1972) 230; Anderson l.c. 204; Whitmore, Tantra &
Sutisna l.c. 161. Lectotype (Soepadmo, 1968): Beccari PB 2551, Borneo, Sarawak (FI).
108
Tree up to 25 m tall, 40 cm diameter; buttresses up to 2 m tall. Bark pale grey, rough;

inner bark dull brown, fibrous. Sapwood creamy or white. Twigs initially densely tomentose
with brownish stellate hairs, subglabrescent, shallowly fissured, sparsely lenticellate.
Terminal buds ovoid to ellipsoid, 34 23 mm. Stipules linear, acute, 23 0.51
mm, tomentose with stellate hairs, caducous. Leaves spirally arranged, thick-coriaceous,
glabrous and glossy above, sparsely tomentose with minute stellate hairs and glaucous
below; elliptic to oblong, (6)1015(20) (2.5)35(8) cm, base rounded or obtuse,
sometimes asymmetrical, margin entire, revolute, apex acute, acuminate or caudate; midrib
impressed or flat above, strongly raised below; lateral veins (6)710(12) pairs, parallel,
faint or flattish on both surfaces, arching towards the leaf margin, forming an angle of
6070o with the midrib; intercostal venation scalariform, obscure on both surfaces; petiole
slender, 2545 mm long, adaxially deeply sulcate towards leaf blade, densely tomentose
with stellate hairs, glabrescent. Male inflorescences (5)1015(20) cm long, densely
tomentose with stellate hairs. Male flowers in dichasial clusters of three or solitary along
the rachis; perianth 46-lobed; stamens 46, filaments 12 mm long, hairy at the base.
Female inflorescences: bracts ovate, acute, densely hairy with short stiff hairs, caducous.
Female flowers: perianth 56-lobed, densely tomentose with stellate hairs; staminodes
rather well-developed; ovary subglobose, c. 1 mm across, styles 34, 11.2 mm long,
sparsely tomentose with stellate hairs at the base. Cupules deeply cup-shaped, tapering
towards the base, 11.5 33.2 cm, covering 1/31/2 of the acorn, woolly pubescent
inside, glabrescent outside; rim up to 0.5 mm thick; lamellae 47, edge free, undulate.
Acorns obovoid to subglobose, c. 2 cm across, glabrous; apex depressed, base convex.
Distribution. Peninsular Malaysia and Borneo, uncommon. In Borneo, known only from
G. Gaharu in Simunjan district and G. Pueh in Lundu district, Sarawak.
Ecology. In hill mixed dipterocarp and kerangas forests, up to 1000 m altitude. Flowering
in January to February; fruiting in June to July.
12. Quercus percoriacea Soepadmo

(Latin, per- = exceedingly, coriaceous = leathery; the thick-leathery leaves)
Gard. Bull. Sing. 21 (1966) 382, l.c. (1968) 396, l.c. (1972) 394; Anderson l.c. 204; Whitmore, Tantra
& Sutisna l.c. 161; Coode et al. (eds.) l.c. 127. Type: Anderson S. 20226, Borneo, Sarawak, Bario,
Ulu Baram (holotype K; isotypes L, SAN, SAR, SING).
Tree up to 30 m tall, 50 cm diameter; buttresses up to 2 m tall. Bark smooth, grey,

hoop-marked. Twigs grey, glabrous, lenticellate. Terminal buds ovoid to subglobose, c.
3 2 mm, densely tomentose. Leaves spirally arranged, thick-coriaceous, glabrous and
shining above, pale glaucous and sparsely tomentose with fine stellate hairs to glabrescent
below; ovate or elliptic, (10)1618(20) (3.5)48(9) cm, base rounded, margin entire,
plane and undulate in the lower half, distantly serrate or crenate in the upper half, apex
acuminate, acumen 1015 mm long; midrib prominently raised beneath, flattish above;
lateral veins prominent below, impressed above, 1012 pairs, distantly spaced, parallel,
arching towards the leaf margin, forming an angle of 4060 with the midrib; intercostal
109
venation dense, fine, scalariform, obscure above, prominent below; petiole 2035 mm,
slightly grooved or flat on the adaxial side, glabrous. Male inflorescences 56 cm long,
many-flowered, densely tomentose with stellate hairs. Male flowers in dichasial clusters
of 34 along the rachis; perianth 56-lobed, hairy; stamen 56, filament 1.52 mm long.
Female inflorescences 24-flowered, up to 1 cm long, densely covered with brown hairs.
Female flowers: perianth 45-lobed, densely hairy; ovary conical, c. 0.8 1.2 mm, styles
34. Cupules shallowly cup-shaped, 0.71 22.3 cm, covering 1/61/5 of the acorn,
base rounded, densely sericeous on both surfaces; rim up to 0.5 mm thick; lamellae 6
8, edge free, denticulate or smooth. Acorns densely sericeous, ovoid to subglobose, 2
2.5 1.52.5 cm; apex depressed, rounded, base convex.
Vernacular names. Sarawakberangan bu, empili (Iban), mempening (Malay), palan
baboi (Kenyah), salad (Murut).
Distribution. Endemic to Borneo. In Sarawak scattered. Not yet reported for Sabah. Also
in Brunei.
Ecology. In primary kerangas forest on plateau sands, at 10001200 m altitude. Fruiting
in June to July.
13. Quercus pseudoverticillata Soepadmo
Fig. 16.
(Latin, pseudo- = somewhat, verticillatus = whorled; the pseudowhorled leaves)

Gard. Bull. Sing. 21 (1966) 380, l.c. (1968) 384, l.c. (1972) 389; Cockburn l.c. (1976) 116; Whitmore,
Tantra & Sutisna l.c. 161. Type: Chew & Corner RSNB. 4434, Borneo, Sabah, Mt. Kinabalu, Bembangan
R., alt. c. 1600 m (holotype K; isotype L).
Tree up to 30 m tall, 90 cm diameter; buttresses up to 2 m high. Bark brown, scaly;

inner bark pink. Sapwood pink. Twigs glabrous, lenticellate. Terminal buds ovoid to
subglobose, 23 2 mm. Leaves pseudowhorled, thick-coriaceous, glabrous above, sparsely
tomentose below; oblanceolate, elliptic or lanceolate, (6)1012(17) (3)44.5(6.5)
cm, base rounded, cordate to auriculate, margin entire, revolute, apex rounded to obtuse;
midrib flat to slightly sunken above, strongly raised beneath; lateral veins 815 pairs,
parallel, arching towards the leaf margin, forming an angle of 3045 with the midrib;
intercostal venation fine, scalariform, obscure and minutely pubescent on both surfaces,
petiole 35 mm long, adaxially flat, tomentose, glabrescent. Inflorescences and flowers
unknown. Cupules deeply cup-shaped, 23.5 24 cm, covering 1/31/2 of the acorn;
base attenuate, densely pubescent inside, glabrescent outside; rim up to 0.5 mm thick;
lamellae 1012, edge free, dentate or undulate. Acorns ovoid to cylindrical, densely
sericeous, 35 23 cm; apex rounded or depressed, base flat to convex.
Vernacular name. Sabahmempening (Malay).
Distribution. Endemic to Borneo. So far only known from Sabah (Mt. Kinabalu) and
Kalimantan.
Ecology. In lower montane forest, up to 1600 m altitude. Fruiting between October and
February.
110
vi
14. Quercus subsericea A. Camus

(Latin, sub- = almost, sericeus=silky; referring to the indumentum of the leaf undersurface)
Bull. Soc. Bot. France 80 (1933) 354, l.c. (1938) 366; Soepadmo l.c. (1968) 392, l.c. (1972) 393; Cockburn
l.c. (1972) 231, l.c. (1976) 116; Anderson l.c. 204; Whitmore, Tantra & Sutisna l.c. 162; Turner l.c. 238;
Coode et al. (eds.) l.c. 127; Argent et al. (eds.) l.c. 255. Type: Teijsmann HB. 21080, Sumatra, Bangka
(holotype BO; isotype K). Synonyms: Q. seriacea (non Willd. 1805) Scheffer, Nat. Tijdschr. Ned. Ind.
31 (1870) 361, King l.c. 63, Merrill l.c. (1921) 215; Cyclobalanopsis seriacea (Scheff.) Schottky l.c.
656; Synaedrys seriacea (Scheff.) Koidzumi, Bot. Mag. Tokyo 30 (1916) 192.
Tree up to 20 m tall, 40 cm diameter; buttresses up to 2 m tall. Bark greyish brown,

finely fissured to scaly, lenticellate; inner bark fibrous, brownish. Sapwood white. Twigs
initially densely tomentose with stiff simple and stellate hairs, glabrescent, minutely
lenticellate. Terminal buds ovoid to subglobose, 34 23 mm, densely tomentose. Stipules
linear, acute, 57 12 mm. Leaves spirally arranged, thin-coriaceous, glabrous and
shining above, not glaucous but sparsely silky tomentose with stellate hairs to glabrous
below; elliptic to lanceolate, rarely ovate, (4)510(16) (2)34(5) cm, base attenuate
or acute, margin entire or occasionally remotely serrulate towards apex, faintly revolute,
apex acute or acuminate, acumen 515 mm long; midrib prominent below, impressed
or sunken above; lateral veins 612 pairs, slightly prominent to flattish below, impressed
above, parallel, arching towards the leaf margin, forming an angle of 4560 with the
midrib; intercostal venation fine, reticulate, obscure on both surfaces; petiole 1025 mm
long, adaxially grooved, glabrous. Male inflorescences with many well-spaced flowers,
46 cm long; bracts ovate, acute, hairy, c. 12 1 mm. Male flowers solitary or in
dichasial clusters of three along the rachis; perianth 56-lobed, tomentose; stamens 5
7, filaments slender, 11.5 mm long. Female inflorescences 25-flowered, 12 cm long,
densely tomentose; bracts ovate, acute, tomentose outside. Female flowers: perianth 5
6-lobed, densely tomentose; ovary ovoid to globose, c. 0.5 mm across, styles 34, tomentose
at base. Cupules shallowly cup-shaped, 0.51.2 0.71 cm, base rounded or attenuate,
covering 1/41/3 of the acorn; rim up to 0.5 mm thick; lamellae 78, densely tomentose,
edge free, denticulate. Acorns ovoid, conical to subglobose, 12 11.5 cm, densely
sericeous; apex acute, depressed or rounded, base rounded.
Distribution. Sumatra, Peninsular Malaysia, Borneo, and W Java. In Sabah and Sarawak
scattered. Also in Brunei and Kalimantan.
Ecology. In lowland to lower montane forests, up to 1500 m altitude, on sandy clay,
sandy loam or granite-derived soils. Flowering and fruiting irregularly.
15. Quercus sumatrana Soepadmo

(of Sumatra)
Gard. Bull. Sing. 21 (1966) 387, l.c. (1968) 392, l.c. (1972) 393; Cockburn l.c. (1972) 229, l.c. (1976)
117; Anderson l.c. 204; Whitmore, Tantra & Sutisna l.c. 162. Type: Achmat 1501, N Sumatra, Tapah
(holotype BO; isotype L).
112
Tree up to 40 m tall, 130 cm diameter; buttresses up to 2.5 m tall. Bark greyish, rough;
inner bark fibrous. Twigs initially densely pubescent with brown stiff-hairs, glabrescent,
smooth and lenticellate. Terminal buds ovoid to subglobose, 23 1.52 mm. Stipules
linear, acute, membranous, 57 1 mm, densely fulvous tomentose with soft, short, simple
hairs. Leaves spirally arranged, chartaceous to thin-coriaceous, glabrous and pale greybrown above, glaucous and sparsely pubescent with stiff simple hairs below; elliptic or
lanceolate, (8)1218(24) (2)3.57(8) cm, base attenuate, acute, margin plane,
remotely serrulate in the apical half, apex acute; midrib prominent beneath, slightly raised
above; lateral veins (8)1012(13) pairs, distantly spaced, prominent below, slightly
so above, parallel, arching towards the leaf margin, forming an angle of 4560 with
the midrib; intercostal venation scalariform, obscure on both surfaces; petiole slender,
1525 mm long, adaxially flat, sparsely tomentose. Male and female inflorescences
unknown. Cupules saucer-shaped, (0.7)11.5 (1.7)22.5(3) cm, covering 1/61/5
of the acorn, base flat, densely tomentose on both surface; rim up to 0.5 mm thick;
lamellae 46, edge free, denticulate. Acorns ovoid to conical, 1.52 22.5 cm, densely
sericeous, glabrescent; apex depressed with umbo up to 3 mm long, base flat to concave.
Distribution. Sumatra and Borneo. In Sabah and Sarawak uncommon. Also in Kalimantan.
on sandy loam or basalt-derived soils. Fruiting between April and December.
16. Quercus treubiana Seemen

(Melchior Treub, 18511910; former Director of the Bogor Botanic Gardens, Indonesia)
Bull. Dp. Agr. Ind. Nerl. 1 (1906) 3; Merrill l.c. (1921) 216; A. Camus l.c. (1938) 368; Soepadmo l.c.
(1968) 412, l.c. (1972) 398; Cockburn l.c. (1976) 117; Anderson l.c. 204; Whitmore, Tantra & Sutisna
l.c. 162. Type: Hallier 2915, Borneo, Kalimantan, Mt. Lianggagang (holotype L; isotypes K, SAR).
Synonym: Cyclobalanopsis treubiana (Seemen) Schottky l.c. 648.
Tree up to 30 m tall, 60 cm diameter; buttresses up to 1 m tall. Bark greyish brown,

rough, peeling off into rectangular pieces; inner bark fibrous, yellowish or pale ochre.
Sapwood brownish white. Twigs initially densely hairy with brownish simple hairs,
glabrescent, lenticellate. Terminal buds ovoid to ellipsoid. Stipules linear, acute, 510
12 mm, densely pubescent outside, caducous. Leaves spirally arranged, thin-coriaceous,
glabrous above, densely pubescent with simple minute hairs below; linear to narrowly
lanceolate, (3)710(12) 13 cm, base attenuate, acute, sometimes asymmetrical, margin
entire or remotely serrulate in the apical half, apex acute to acuminate, acumen 510
mm long; midrib prominent on both surfaces; lateral veins 510 pairs, distantly spaced,
parallel, arching towards the leaf margin, forming an angle of 4560 with the midrib;
intercostal venation fine, scalariform, obscure on both surfaces; petiole slender, 1020
mm long, adaxially flat, tomentose, glabrescent. Male inflorescences up to 3 cm long,
densely pubescent; bracts linear, acute. Male flowers solitary or in dichasial clusters of
three along the rachis; perianth 56-lobed, densely sericeous outside; stamens 56, filaments
hairy at base, up to 2 mm long; pistillode replaced by a tuft of stiff simple hairs. Female
113
inflorescences 13-flowered, 1.53 cm long, densely hairy; bracts ovate, acute. Female
flowers: perianth 5-lobed, densely hairy; ovary conical, c. 0.5 1.3 mm, styles 3, 1.5
2 mm long, densely pubescent at base. Cupules cup-shaped, 11.5 23 cm, covering
1/31/2 of the acorn, base attenuate or rounded, woolly pubescent on both sides; rim
11.5 mm thick; lamellae 1012, edge tightly pressed towards cupule wall, undulate. Acorns
ovoid, subglobose or cylindrical, 22.5 23 cm, glabrous; apex depressed, base convex.
Vernacular names. Sabahmempening (Malay). Sarawakempili (Iban).
Distribution. Sumatra and Borneo. In Sabah, known from Mt. Kinabalu and Mt. Tombuyokan.
In Sarawak recorded for Hose Mts. in Kapit district, Bt. Lumut in Bintulu district, and
Merurong Plateau. Also in Kalimantan.
Ecology. In hill mixed dipterocarp forest to montane forest, at 6002100 m altitude, on
sandy waterlogged soils. Fruiting between July and August.
17. Quercus valdinervosa Soepadmo

(Latin, valde = strongly or prominently, nervosus = nerved; the distinctly veined leaves)
Gard. Bull. Sing. 21 (1968) 404, l.c. (1972) 395; Cockburn l.c. (1976) 117; Anderson l.c. 204; Coode
et al. (eds.) l.c. 127. Lectotype (Soepadmo, 1968): Clemens 31317, Borneo, Sabah, Mt. Kinabalu
(holotype K; isotypes A, B, K, L, SING, UC). Synonym: Q. mespilifolia Wall. ex A. DC. var. borneensis
Heine, Feddes Rep. 54 (1951) 54.
Tree up to 35 m tall, 60 cm diameter. Bark smooth, grey-brown; inner bark red-brown.

Sapwood pale ochre. Twigs initially densely hairy with simple and stellate golden brown
hairs, glabrescent, lenticellate. Terminal buds densely tomentose, ovoid to subglobose,
0.51 0.5 cm. Stipules densely hairy, linear, acute, 1015 12 mm. Leaves spirally
arranged, thick-coriaceous, glabrous above, densely pubescent beneath, soon glabrescent;
elliptic, oblong or obovate, (5)613(14) (2.5)35(6) cm, base attenuate, rounded,
or acute, margin plane, remotely serrate in the apical half, apex acute to acuminate,
acumen 510 mm long; midrib flat or impressed above, prominent beneath; lateral veins
1015 pairs, closely spaced, strongly prominent beneath, parallel, arching towards the
leaf margin, forming an angle of c. 60 with the midrib; intercostal venation dense,
scalariform, prominent beneath; petiole 1225 mm long, flat on the adaxial side, densely
hairy, soon glabrescent. Male inflorescences many-flowered, 410 cm long; bracts triangular,
acute, 35 23 mm, densely hairy with stiff golden brown hairs. Male flowers: perianth
56-lobed, densely hairy; stamens 69, filaments 34 mm long. Female inflorescences
46-flowered, 1.52 cm long, densely hairy; bracts narrowly ovate to linear, 1.52
1 mm. Female flowers: perianth 6-lobed, densely hairy; ovary conical, c. 0.6 0.8 mm,
styles 3, recurved, 1.52 mm long. Cupules cup-shaped to bowl-shaped, 1.52 33.5
(4) cm, covering up to 1/3 of the acorn, base attenuate, rounded, densely hairy on both
surfaces; rim 24 mm thick; lamellate 710, free edge crenate. Acorns ovoid, cylindrical
or ellipsoid, densely tomentose, 34 1.53(4) cm; apex acute, base convex.
114

Distribution. Endemic to Borneo. In Sabah known from Mt. Kinabalu. In Sarawak reported
for Kelabit Highlands and G. Mulu. Also in Brunei and Kalimantan.
Ecology. In lower to upper montane forests, at 13002300 m altitude, on brownish ultrabasic
and organic soils. Fruiting between February and August.
4. TRIGONOBALANUS Forman
(Greek, trigonos = 3-angled, balanos = acorn)
berangan beluyan (Iban), mempening babi (Malay).
Taxon 11 (1962) 140; Kew Bull. 17 (1964) 381, ibid. 21 (1967) 331; Soepadmo, FM 1, 7 (1972) 398;
Cockburn, TFM 1 (1972) 232, TS 1 (1976) 119; Lozano-C, Caldesia 12 (1979) 517; Corner, WSTM
3rd. edition 1 (1988) 343; Nixon & Crepet, Am. J. Bot. 76 (1989) 828; Kubitzki in Kubitzki et al.
(eds.), Fam. & Gen. Vasc. Pl. 2 (1993) 308.
Trees with coppicing habit. Terminal bud solitary. Stipules interpetiolar. Leaves (in Sabah
and Sarawak) in whorls of 3; margin toothed above the middle; petiole slightly thickened
at the base. Inflorescences spikes, male, female, androgynous or mixed. Male inflorescences
pendent or patent, simple or branched, in the axils of normal lower (proximal) leaves
or in the axils of reduced leaves and crowded together in lateral or subterminal paniculate
clusters; bracts and bracteoles small, caducous. Male flowers in dichasial clusters of (1
)37(12) along the rachis; perianth membranous, 6-lobed, the lobes imbricate; stamens
6, filaments free, exserted; anthers basifixed, ovoid, 2-loculed, pistillode replaced by a
cluster of erect simple hairs. Female, androgynous or mixed inflorescences erect, simple
or branched, solitary in the axils of upper (distal) leaves; bracts and bracteoles small,
caducous. Female flowers in dichasial clusters of (1)37(15); perianth with 6 imbricate
lobes, the lower part adnate to the ovary; staminodes 6, well-developed and exceeding
the perianth or remaining shorter than the perianth, sometimes polliniferous; styles 3,
recurved or connate at the base; stigmas capitate. Cupules in condensed clusters of 2
3 or solitary along the rachis, partly enclosing 17(12) nuts (fruits), 312-lobed, outside
covered with transversely dentate lamellae. Nuts trigonous in cross-section, ovoid to
triangular in outline; wall thinner than 0.5 mm, most part free from the cupule. Seed
Distribution. Three species with disjunct distribution: Trigonobalanus doichangensis
occurs in N Thailand; T. verticillata in Peninsular Malaysia, Borneo and C Sulawesi;
and T. excelsa in Colombia, S America. In Borneo, T. verticillata occurs in Sabah and
Sarawak.
Ecology. Confined to hill and lower montane forests, at 7001800 m altitude.
115
Trigonobalanus verticillata Forman
Fig. 17.
(Latin, verticillatus = whorled; the leaves)

Taxon 11 (1962) 140, l.c. (1964) 383, l.c. (1967) 331; Soepadmo l.c. (1972) 400; Cockburn l.c. (1972)
232, l.c. (1976) 119; Corner l.c. 343; Whitmore, Tantra & Sutisna l.c. 162; Turner l.c. 238. Type: RSNB
2732A, Borneo, Sabah, Mt. Kinabalu, Liwagu R. (holotype K; isotypes L, LE, US).
Tree up to 36 m tall, 70 cm diameter; producing many sucker-shoots at the base; low

buttresses or stilt-roots occasionally present. Bark smooth, shallowly fissured to scaly,
pale grey-brown; inner bark granular, reddish brown. Sapwood pale reddish. Twigs
subglabrous, densely or sparsely minute-lenticellate. Stipules ovate, acute, 35 2 mm.
Leaves rigid, coriaceous, glabrous above, sparsely appressed tomentose with stellate hairs
below; elliptic or obovate, 58(10.5) (2.2)2.54.5(5.5) cm, base acute to cuneate,
margin strongly revolute, toothed on the apical half, apex obtuse to rounded and emarginate;
midrib raised on both surfaces, stronger below; lateral veins thin, 810(11) pairs, lax
or dense, prominent on both surfaces, stronger or sometimes flat below, arching and
disappearing towards the leaf margin, forming an angle of 4060 with the midrib;
intercostal venation reticulate, dense, obscure above, thinly prominent below; petiole
(3) 410 mm long. Male inflorescences 410 cm long; bracts and bracteoles broadly
ovate, 11.5 0.8 mm. Male flowers solitary but arranged closely together along the
rachis; perianth 56-lobed, lobes coriaceous, elliptic, c. 1 0.8 mm; stamens 56, filaments
1.52 mm long. Androgynous or mixed inflorescences 510 cm long; bracts and bracteoles
broadly ovate, 0.81 0.50.8 mm. Female flowers in dichasial clusters of (2)3
7(15) along the rachis; perianth 56-lobed, lobes coriaceous, elliptic c. 1 0.8 mm;
staminodes 57; styles 3, cylindrical, straight, 1.52 mm long. Cupules sessile, open, 3
12-lobed, the lobes acute, 48 36 mm, densely appressed tomentose on both sides; wall
bony, thin, usually containing 23 nuts; scale-like appendages minute, distinct, set irregularly.
Nuts ovoid to conical, 57 35 mm, sparsely tomentose, brownish; base truncate, apex
acute; scar flat, triangular, c. 2 mm diameter; wall bony, thin, most part free from the cupule.
Vernacular names. Sabahmempening babi (Malay). Sarawakberangan beluyan (Iban),
salad rettan (Kelabit), mengkusub (Tagar).
Distribution. Peninsular Malaysia, Borneo and C Sulawesi. In Sabah, common in Ranau
district (Kundasang, Kinasaraban, Sosopodon, and Kinabalu Park) and once collected from
Bt. Rimau, Long Pasia in Sipitang district. In Sarawak, common in Lawas area, and in Bario
and Mt. Hose in Kapit district. Not known in Brunei and Kalimantan.
Ecology. In primary and old secondary hill to lower montane forests, including kerangas
forest, at 7001600 m altitude, on ultrabasic or loamy soils overlying Tertiary sandstone;
often gregarious along streams. Flowering in SeptemberFebruary, fruiting in AprilNovember.
116
vi
118
LEGUMINOSAE-CAESALPINIOIDEAE (DING HOU)
LEGUMINOSAE
(Subfamily CAESALPINIOIDEAE)
Ding Hou
Rijksherbarium/Hortus Botanicus
Leiden, the Netherlands
Backer in Hooker f., Fl. Brit. Ind. 1 (1876) 60 (as suborder Caesalpinieae); Prain, J. As. Soc. Beng. 66,
2 (1897) 152 (as suborder Caesalpinieae); Ridley, FMP 1 (1922) 616 (as suborder Caesalpinieae); Backer
& Bakhuizen f., FJ 1 (1964) 523 (as Caesalpiniaceae); Hutchinson, Gen. Fl. Pl. 1 (1964) 221 (as
Caesalpiniaceae); Keng, OFMSP (1969) 65; Whitmore, TFM 1 (1972) 245; Cockburn, TS 1 (1976)
156; Anderson, CLTS (1980) 231 (as part of Leguminosae); Cowan in Polhill & Raven (eds.), Adv. Leg.
Syst. 1 (1981) 57; Watson & Dallwitz, Gen. Leg.-Caesalp. (1983) 1; Corner, WSTM 3rd. edition 1
(1988) 418 (as Caesalpiniaceae); Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 193 (as part of
Leguminosae); Kessler & Sidiyasa, Trees of the Balikpapan-Samarinda Area (1994) 71 (as
Caesalpiniaceae); Turner, Gard. Bull. Sing. 47, 1 (1995) 289 (as part of Leguminosae); Ding Hou, K. &
S.S. Larsen, FM 1, 12 (1996) 409 (as Caesalpiniaceae); Coode et al. (eds.), CLBD (1996) 160; Loo &
Tan, Gard. Bull. Sing. 49, 1 (1997) 55 (as Caesalpinaceae); Argent et al. (eds.), MNDT-CK 2 (1997)
342 (as part of Leguminosae).
Trees, shrubs, or woody lianas; sometimes armed with thorns or spines; bole sometimes
buttressed. Stipels usually lacking, or minute when present. Stipules usually present, often
caducous. Leaves often spiral, compound (once pinnate or bipinnate, rarely unifoliolate)
or simple, petiolate. Inflorescences axillary and/or terminal, or borne on older branches
and stems, solitary or fascicled, commonly paniculate, through reduction and condensation
leading to racemose, spicate, rarely 1-flowered; bracts and bracteoles usually present,
often caducous. Flowers usually bilaterally symmetrical or less often radially symmetrical,
bisexual and/or unisexual, usually 5-merous except for the gynoecium; hypanthium (or
receptacle) usually present, cupular, tubular, or turbinate, sometimes obscure or absent,
inside often containing nectariferous tissue; calyx lobes or sepals generally 5, sometimes
4, usually imbricate; petals usually 5, sometimes reduced to 4 (or 1), or 0, often unequal;
disk sometimes present and extrastaminal; stamens basically 10, mostly unequal, rarely
fewer, or 0 (in female flowers), all fertile or some reduced; filaments free or variously
connate at the base; anthers variously developed, basi- or dorsifixed, often versatile,
usually longitudinally dehiscent, rarely dehisce by apical or basal pores; pistil stipitate
and often partly adnate to the hypanthium, rarely sessile; ovary (1)fewmany-ovuled,
style short or long, often curving inward, stigma large or small, capitate or peltate, sometimes
obscure. Fruit variable, usually a pod, oblong to linear or strap-shaped, laterally compressed,
straight or curved, dehiscent and 2-valved, valves chartaceous, fleshy, coriaceous, or woody,
thick, indehiscent and drupaceous, or winged; with or without pulp; glabrous, hairy, or
spiny. Seeds varying in form, often flattened; testa membranous, coriaceous, or crustose;
sometimes arillate; endosperm present or lacking; cotyledons fleshy or foliaceous, radicle
straight or slightly oblique, never folded.
119
Distribution. Predominantly tropical plants comprising about 160 genera and over 2000
species. In Sabah and Sarawak, the subfamily is represented by 42 species in 17 genera.
Ecology. In tropical primary and secondary forests mostly at low and medium altitudes
to 600(1500) m, growing on hillsides, ridges, river banks, swampy or marshy places,
or just behind the mangroves, occasionally on limestone or sandstone.
Uses. Members of some genera, e.g., Afzelia, Cynometra, Intsia, etc., are sources for
important (commercial) timbers. Some species produce resin or wood-oil, medicine, or
tannin. Trees of some species are cultivated for ornamental purpose.
Key to genera
(for flowering specimens)
1. Leaves simple, bilobed, or deeply divided (as 2 free leaflets).........
Bauhinia L.
Sp. Pl. 1 (1753) 374; Prain, J. As. Soc. Beng. 66, 2 (1897) 175; Ridley, FMP 1 (1922) 624;
Backer & Bakhuizen f., FJ 1 (1964) 531; Corner, WSTM 3rd. edition 1 (1988) 419; Coode et
al. (eds.), CLBD (1996) 160; K. Larsen & S.S. Larsen, FM 1, 12 (1996) 442; Loo & Tan, Gard.
Bull. Sing. 49, 1 (1997) 57. Synonyms: Phanera Lour., Fl. Coch. (1790) 37, de Wit, Reinwardtia
3 (1956) 435; Bracteolanthus de Wit l.c. (1956) 415; Lysiphyllum (Benth.) de Wit l.c. (1956)
431.
A pantropical genus consisting of about 300 species. About 28 species are found
Woody tendrilled climbers, shrubs or trees. Leaves simple, bilobed, or deeply
divided (as 2 free leaflets); venation palmate (or fan-like), or parallel. Flowers
bisexual, solitary, or in racemes, corymbs or panicles; calyx cup-shaped, 25lobed; petals usually 5, subequal; stamens 010 fertile, staminodes often present;
ovary 1many-ovuled. Fruits usually dehiscent. In various habitats, from lowland
to lower montane forest up to 1500 m altitude.
Leaves compound, rarely unifoliolate (as a simple leaf)...............2
2. Leaves bipinnate.................3
Leaves once pinnate (paripinnate or imparipinnate), rarely unifoliolate.............6
3. Plants armed with spines or prickles................4
Plants not armed.................5
4. Floral parts often punctate; calyx lobes and petals usually bilaterally symmetrical;
ovary 110(13)-ovuled...............................
Caesalpinia L.
Sp. Pl. (1753) 380; Prain, J. As. Soc. Beng. 66, 2 (1897) 225; Ridley, FMP 1 (1922) 648;
Backer & Bakhuizen f., FJ 1 (1964) 544; Whitmore, TFM 1 (1972) 246; Hattink, Reinwardtia
9 (1974) 9; Corner, WSTM 3rd. edition 1 (1988) 426; Ding Hou, FM 1, 12 (1996) 535; Coode
et. al. (eds.), CLBD (1996) 161; Loo & Tan, Gard. Bull. Sing. 49, 1 (1997) 60. Synonym:
Poinciana L., Sp. Pl. (1753) 380, Backer & Bakhuizen f. l.c. 546.
A pantropical genus consisting of about 100 species. In Sabah and Sarawak

represented by 6 indigenous species.
120
Lianas, shrubs, or small trees, usually spiny or prickly. Leaves bipinnate; rachises
on the underside often with prickles; leaflets few to many pairs, no terminal leaflet.
Inflorescences axillary and/or terminal, racemose or paniculate. Flowers usually
bisexual, bilaterally symmetrical, all parts often punctate (secretory cavities);
hypanthium usually obliquely short-cupular or funnel-shaped; calyx lobes 5,
unequal, the lowest one hood-shaped; petals 5, unequal; stamens 10, free, anthers
uniform; ovary flat, 110(13)-ovuled. Fruits very variable, dehiscent or indehiscent,
more or less flattened, wingless or winged along the upper suture, 110(13)seeded. Seeds orbicular or globose, ellipsoid or reniform, sometimes flattened,
often smooth, usually without endosperm.
Mostly in secondary vegetation, sometimes in coastal, rarely in primary forest,
on various types of soils from sea level to c. 1700(2000) m altitude.
Floral parts not punctate; calyx lobes and petals slightly unequal; ovary 12-ovuled......
Pterolobium R. Brown ex Wight & Arn., nom. cons.
Prod. Fl. Ind. Orient. 1 (1834) 283; Prain, J. As. Soc. Beng. 66, 2 (1897) 235; Ridley, FMP 1
(1922) 648; Backer & Bakhuizen f., FJ 1 (1964) 543; Ding Hou, FM 1, 12 (1996) 654.
About 11 species in the Old World. Only 1 species (P. borneense Merr.) in Sarawak:
Baram district; also known in Kalimantan.
Tall climbers or climbing shrubs armed with recurved prickles. Leaves bipinnate;
leaflets numerous, small. Flowers bisexual, in terminal or axillary panicles of
racemes; calyx 5-lobed; petals 5, the upper one (standard) usually different in
shape and size; stamens 10, free; ovary free, 12-ovuled. Fruits samaroid, basal
seed-bearing part flattened, indehiscent, usually 1-seeded; upper wing-like part
membranous.
In lowland forest up to 500 m altitude.
5. Leaflets (1)2(36) pairs per pinna. Petals 2.55 mm long; stamens with glabrous
filaments..........13. Sympetalandra (in part)
Leaflets often 427 pairs per pinna. Petals 1225 mm long; stamens with filaments
pilose or villous at the base or basal part................10. Peltophorum
6. Leaves imparipinnate, rarely unifoliolate..................7
Leaves paripinnate.....................11
7. Petals 5...................8
Petals absent or greatly reduced................9
8. Leaves with only 1 leaflet..............14. Uittienia
Leaves with 512(17) leaflets.................8. Koompassia
9. Calyx lobes 5 or 3; stamens 2 or 6............6. Dialium
Calyx lobes 4; stamens usually 10..............10
10. Inflorescences racemose. Ovary 16-ovuled.....................4. Crudia
Inflorescences paniculate. Ovary 2-ovuled...................3. Copaifera
11. Petals absent...................11. Saraca
Petals present, 15..................12
121
12. Petal only 1 developed, the others rudimentary or wanting..........13

Petals often 25 developed..................15
13. Calyx lobes often spiny outside; fertile stamens often 10..............12. Sindora
Calyx lobes smooth outside; fertile stamens 38...............14
14. Stamens often (5)7 fertile.................1. Afzelia
Stamens often 3 fertile...............7. Intsia
15. Petals 2 (or 1); fertile stamens 2.....................9. Leucostegane
Petals usually 5; fertile stamens usually 10.......................16
16. Calyx and corolla pellucid-glandular; stamens 10, alternately long and short.....
........13. Sympetalandra (in part)
Calyx, corolla and stamens not as above................17
17. Leaves with 412 pairs of symmetrical leaflets. Stamens 3 + 7, unequal (3 almost
twice as long as the others); ovary many-ovuled..........2. Cassia
Leaves usually with only 1 or 2 pairs of asymmetrical leaflets. Stamens (8)10
(15), equal in length; ovary 12( 4)-ovuled........5. Cynometra
Key to genera
(for fruiting specimens)
1. Leaves simple, bilobed, or deeply divided (as 2 free leaflets)...............Bauhinia
Leaves compound or unifoliolate (as a simple leaf)................2
2. Leaves bipinnate...................3
Leaves once pinnate (paripinnate or imparipinnate), or unifoliolate....................7
3. Plants armed with spines or prickles................4
Plants not armed.................6
4. Fruits not winged...............Caesalpinia (in part)
Fruits winged......................5
5. Fruits samaroid, winged above the seed-bearing part..........Pterolobium
Fruits winged along one suture side.............Caesalpinia (in part)
6. Fruits indehiscent....................10. Peltophorum
Fruits dehiscent, 2-valved..............................13. Sympetalandra (in part)
7. Leaves imparipinnate.........................8
Leaves paripinnate....................12
8. Fruits surrounded by a papery wing..................8. Koompassia
Fruits not winged....................9
122
9. Fruit a pod, dehiscent..................10

Fruit drupaceous, indehiscent.......................11
10. Fruits 14(6)-seeded; seeds without aril......................4. Crudia
Fruits 12-seeded; seeds enclosed by 2-lobed aril...........3. Copaifera
11. Fruits with a thin-crustaceous, usually hairy exocarp, easily crushed or sometimes
detached from mesocarp. Leaves each usually with (1 or) 3 or more leaflets.......
........6. Dialium
Fruits with a crustaceous, glabrous exocarp united with a thick (45 mm) and hard
mesocarp, uncrushable when pressed hard between two hands. Leaves with only 1
leaflet..............14. Uittienia
12. Fruits often spiny outside................12. Sindora
Fruits smooth or rugose outside..............13
13. Fruits indehiscent.................14
Fruits dehiscent.....................15
14. Fruits elongate, cylindrical, 2060 cm long, many-seeded. Leaves with 412 pairs
of leaflets....................2. Cassia
Fruits broad ovoid or broadly ellipsoid, or obovoid, less than 8 cm long, 1(4)-seeded.
Leaves with 1 or 2 pairs of leaflets, rarely unifoliolate......5. Cynometra
15. Stipules foliaceous, often (5)12.540 mm long, persistent......9. Leucostegane
Stipules minute, caducous................16
16. Seeds distinctly arillate.................1. Afzelia
Seeds not arillate.................17
17. Valves of fruits twisting and enrolling, or coiling during dehiscence...........11. Saraca
Valves of fruits often straight or slightly twisting during dehiscence...............18
18. Leaflets usually with 1 or 2 small crateriform glands at the basal part on lower surface.
Valves of fruits leathery or slightly woody..................7. Intsia
Leaflets without the glands as above. Valves of fruits distinctly woody......
......13. Sympetalandra (in part.)
1. AFZELIA Sm., nom. cons.

(A. Afzelius, 17501837, Sweden, one of the best students of Linnaeus, Professor of
Botany at Upsala, collected plant specimens in the west coasts of Africa)
Trans. Linn. Soc. 4 (1798) 221, non J.E. Gmelin, l791 (= Seymeria Pursh, Scrophulariaceae); Prain, J.
As. Soc. Beng. 66, 2 (1897) 207; Ridley, FMP 1 (1922) 639; Cockburn, TS 1 (1976) 157; Anderson,
CLTS (1980) 232; Cowan & Polhill in Polhill & Raven (eds.), Adv. Leg. Syst. 1 (1981) 128; Whitmore,
Tantra & Sutisna, CLK 2, 1 (1990) 194; Ding Hou, FM 1, 12 (1996) 438; Argent et al. (eds.), MNDTCK 2 (1997) 346. Synonym: Pahudia Miq., Fl. Ned. Ind. l, l (1855) 85.
123
Trees. Stipules interpetiolarly connate at base, minute, caducous. Leaves compound, once
pinnate and paripinnate. Leaflets chartaceous, shortly petiolulate, petiolules often turned
or twisted. Inflorescences terminal or axillary, racemose, often fasciculate or paniculate;
bracts and bracteoles small, often caducous. Flowers bisexual, bilaterally symmetrical,
pedicelled; hypanthium cupular, narrowly infundibuliform or cylindrical, puberulous
outside, glabrous inside; calyx lobes 4, imbricate, puberulous and smooth on both surfaces;
petals only one fully developed, large, flabellate, lower half narrowed into a claw, the
others small or absent; stamens 9, usually (5)7 fertile, almost equal; ovary 38(or more)ovuled. Fruits smooth outside, dehiscent, oblong, obliquely oblong, or slightly rhomboid,
compressed, black when ripe, glabrous, 2-valved; valves thick, woody; often 3(or more)seeded. Seeds ellipsoid, ovoid-oblong or broadly ellipsoid, smooth, without endosperm;
aril yellow, orange, or red, often 2-lobed, covering the seed for up to half or more of
its length.
Distribution. About 12 species, in tropical Africa, Asia and S China; 1 species in Sabah
and Sarawak.
Ecology. In forest at low and medium altitudes, up to 400(1400) m.
Afzelia rhomboidea (Blanco) Vidal
Fig. 1.
(Latin, rhomboideus = oval, a little angular in the middle; the fruits)

Cat. Pl. Prov. Manila (1880) 28; Whitmore, Tantra & Sutisna l.c. 194; Ding Hou l.c. (1994) 316, l.c.
(1996) 441; Argent et al. (eds.) l.c. 348. Basionym: Eperua rhomboidea Blanco, Fl. Filip., ed. 2 (1845)
260. Neotype (Ding Hou, 1994): Merrill Sp. Blanc. no. 862, the Philippines, Luzon, Laguna Prov.
(holotype L; isotype BO). Synonyms: Pahudia rhomboidea (Blanco) Prain, Sc. Mem. Med. Off. Army
Ind. 12 (1901) 46, Merrill, Spec. Blanc. (1918) 171; Intsia acuminata Merr., Publ. Gov. Lab. Philip. 17
(1904) 20; A. borneensis Harms in Fedde, Rep. Sp. Nov. 14 (1916) 256, Cockburn l.c. 157, Anderson
l.c. 232, Whitmore, Tantra & Sutisna l.c. 194; P. acuminata (Merr.) Merr., Philip. J. Sc. 11 (1916) Bot.
86; A. acuminata (Merr.) Harms in Fedde, Rep. Sp. Nov. 15 (1917) 19; P. borneensis (Harms) Merr., J.
Str. Br. R. As. Soc. 76 (1917) 84, EB (1921) 297, Masamune, EPB (1942) 349; P. rhomboidea (Blanco)
Prain var. praetermissa de Wit, Bull. Bot. Gard. Buitenz. 3, 17 (1941) 151; A. rhomboidea (Blanco)
Vidal var. praetermissa Lonard, Reinwardtia 1 (1950) 63.
Tree up to 36 m tall, 80(120) cm diameter; bole cylindrical; buttresses sometimes present,

up to 2 m high. Bark dark grey, deeply pock-marked; inner bark deep green-brown.
Sapwood pale yellow, heartwood bright orange-red. Twig often hollow; young twig, rachis,
petiole, petiolules, and leaflets often glabrous, sometimes sparsely, very rarely densely
pubescent, especially on the midrib of the lower (sometimes also on the upper) leaf surface.
Leaves 1014(19) cm long, each with 35 pairs of leaflets. Leaflets ovate, elliptic,
3.510(15) 2.54.5(6) cm, base acute or obtuse, rarely more or less truncate or shallowly
concave, apex (shortly)acuminate; lateral veins 58 per side, intercostal venation rather
closely reticulate; petiolules 1.54.5 mm long. Inflorescences 1214(15) cm long, both
main axis and branches puberulous; pedicel 4.55.5 mm long. Flowers: calyx lobes broadly
ovate or subrotundate, 512 59 mm; developed petals 710 59 mm; stamen filaments
slightly hairy near the base; ovary puberulous along the margin, style of two kinds: long
(4050 mm), or very short (12 mm), slightly hairy at the base. Fruits 912(20) 5
124
vi
7(10) 1.5 cm, 26-seeded. Seeds 1.73 1.52.5 1 cm, black with a rich orange
aril.
Vernacular names. Sabahipil darat (preferred name), merbau tanduk (Malay, Tawau).
Distribution. Sumatra, Borneo and the Philippines. In Sabah scattered, known for Kalabakan
and Tawau in Tawau district, and Mt. Kinabalu in Ranau district. In Sarawak, known
by a single record from Semengoh Arboretum in Kuching district. Also in Kalimantan.
Ecology. In primary forest, occasionally occurring on limestone or sandstone, up to 350
m altitude, once found at c. 1400 m on Mt. Kinabalu. Flowering and fruiting almost
all year round.
Use. Valuable timber tree, fresh logs sink.
2. CASSIA L.
(from the Greek name kassia)
Sp. Pl. 1 (1753) 376; Prain, J. As. Soc. Beng. 66, 2 (1897) 153; Ridley, FMP 1 (1922) 616; Backer &
Bakhuizen f., FJ 1 (1964) 535; Whitmore, TFM 1 (1972) 246; Cockburn, TS 1 (1976) 158; Anderson,
CLTS (1980) 232; Irwin & Barneby in Polhill & Raven (eds.), Adv. Leg. Syst. 1 (1981) 105; Mem. N.Y.
Bot. Gard. 35 (1982) 4; K. Larsen & S.S. Larsen, Fl. Thail. 4 (1984) 102, p.p.; Corner, WSTM 3rd.
edition 1 (1988) 428; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 197; Rudd, Rev. Handb. Fl. Ceylon
7 (1991) 59, p.p., excl. syn.; K. Larsen & Ding Hou, FM 1, 12 (1996) 556; Coode et al. (eds.), CLBD
(1996) 162; Argent et al. (eds.). MNDT-CK 2 (1997) 352.
Trees or large shrubs. Leaves spirally arranged, often distichous, compound, once pinnate
and paripinnate, each with 412 pairs of symmetrical leaflets; extrafloral nectaries absent.
Inflorescences terminal on main branches or on short side shoots, racemose. Flowers:
hypanthium variable; calyx lobes 5; petals 5; stamens 10, unequal (3 + 7): 3 with filaments
curved, usually longer than the others, and anthers dehiscing by slits, the remaining 7
filaments straight and short with anthers mostly dehiscing by basal pores; ovary manyovuled. Fruits smooth outside, elongate, cylindrical or compressed, indehiscent, 2060
cm long. Seeds many, 12-seriate, without aril.
Distribution. About 30 species, in tropical America, Africa, Asia, Malesia and Australia.
In Sabah and Sarawak, only one indigenous species is recorded.
Cassia javanica L.
(of Java)
Sp. Pl. (1753) 379; Prain l.c. 156; Merrill l.c. (1921) 299; Masamune l.c. 338; de Wit, Webbia 11 (1955)
214; Backer & Bakhuizen f. l.c. 537; Burkill, EPMP 1 (1966) 476; Verdcourt, Lae Bot. Bull. 11 (1979)
47; K. Larsen & S.S. Larsen, Fl. Cambodia, Laos & Vietnam 18 (1980) 84; Irwin & Barneby, Mem. N.Y.
Bot. Gard. 35 (1982) 46; K. Larsen & S.S. Larsen, Fl. Thail. 4 (1984) 107; Corner l.c. 431; Whitmore,
126
vi
Tantra & Sutisna l.c. 198; K. Larsen & Ding Hou, FM 1, 12 (1996) 560; Argent et al. (eds.) l.c. 352.
Type (de Wit, 1955): Commelin s.n., Hort. Med. Amst. 1 (1697) 217, t. 111.
Distribution. A very polymorphous species with a wide distribution ranging from N India
through the Indo-Chinese Peninsula and the Malesian region to New Guinea; with 5
subspecies widespread in the Malesian area, wild and cultivated; one of them in Sabah
and Sarawak.
subsp. nodosa (Buch.-Ham. ex Roxb.) K. Larsen & S.S. Larsen

(Latin, nodus = node; with nodes or joints)
Fig. 2.
Nat. Hist. Bull. Siam Soc. 25 (1974) 205, l.c. (1980) 85, l.c. (1984) 108; Rudd l.c. 68; Turner, Gard.
Bull. Sing. 47, 1 (1995) 296; K. Larsen & Ding Hou l.c. (1996) 564. Basionym: C. nodosa Buch.-Ham.
ex Roxb., Fl. Ind. ed. 2, 2 (1832) 336, Ridley l.c. (1922) 616, de Wit l.c. (1955) 223, Backer & Bakhuizen
f. l.c. 537, Cockburn l.c. 158, Anderson l.c. 233, Irwin & Barneby l.c. (1982) 50, Corner l.c. 431, Keng,
Conc. Fl. Sing. (1990) 33; Whitmore, Tantra & Sutisna l.c. 198, Ng, MFFSS 2 (1992) 427. Type (Irwin
& Barneby, 1982): Authentic collection from Hort. Bot. Calcutta., distributed as Wallich Herb. no. 5331
(K-W), a native of Chittagong, India.
Deciduous tree, up to 35 m tall, 36 cm diameter; trunk unarmed. Leaves each with 4

12 pairs of leaflets. Leaflets elliptic-ovate to oblong, 410 1.53.5 cm, base obtuse
or rounded, apex acute; lower surface pubescent, glabrescent. Inflorescences axillary,
racemose, corymbose, 410 cm long; peduncle 23 cm long; pedicel 23.5 cm long. Flowers
white, later pink; calyx lobes green in living state, ovate, c. 6 mm long; petals obovate,
1.52 mm long, contracted into a short claw; stamens: 3 long ones with the filament
recurved, 1520 mm long, with a spherical enlargement near the middle of the 20 mm
long filament, 7 shorter ones 1012 mm long. Fruits dangling, round, black, 3070
1.22 cm. Seeds many, pale brown, flat, more or less orbicular, 0.81 cm diameter.
Vernacular names. Sabahbarung-barung (Malay), busuk-busuk (preferred name), langgong
(Kalabakan).
Distribution. E Himalayas, southern Thailand, through Peninsular Malaysia, Borneo to
Java and Lesser Sunda Islands. In Sabah scattered in most districts; in Sarawak, rare,
reported from Lundu district.
Ecology. In primary forests at low altitude.
3. COPAIFERA L., nom. cons.

(Derived from copaiba, the Brazilian name for the balsam of capevi, and
Latin, -fer = to bear or to carry; tree producing the balsam of capevi)
Sp. Pl. ed 2 (1762) 557; Anderson, CLTS (1980) 233; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990)
198; Ding Hou, Blumea 38 (1994) 320, FM 1, 12 (1996) 570; Coode et al. (eds.), CLBD (1996) 162.
128
Synonym: Pseudosindora Symington, Proc. Linn. Soc. Lond. 155 (1944) 285, Cowan & Polhill in
Polhill & Raven (eds.), Adv. Leg. Syst. 1 (1981) 132.
Trees. Leaves compound, once pinnate and imparipinnate, petiolate; leaflets alternate,
petiolulate. Stipules caducous. Inflorescences axillary, paniculate; bracts and bracteoles
small, caducous. Flowers bisexual, radially symmetrical, pedicelled; hypanthium unknown;
calyx lobes 4, narrowly imbricate or subvalvate; petals absent; stamens 10, free; anthers
dorsifixed; ovary 2-ovuled, style slender, stigma small, capitellate. Fruits not winged,
dehiscent pods, ellipsoid, rather smooth, 2-valved; valves thick-coriaceous; 12-seeded.
Seeds without endosperm, oblong, enclosed by 2-lobed aril.
Distribution. A genus comprising 2530 species, mostly in tropical America, 4 in tropical
Africa, and one in Borneo. In Sabah and Sarawak one species; also in Brunei and
Kalimantan.
Copaifera palustris (Symington) de Wit
Fig. 3.
(Latin, palustris = swampy, marshy; plants growing in marshy habitats)

Webbia 9 (1954) 462; Anderson l.c. 233; Whitmore, Tantra & Sutisna l.c. 198; Ding Hou l.c. (1994)
321, l.c. (1996) 573; Coode et al. (eds.) l.c. 162. Basionym: Pseudosindora palustris Symington, Proc.
Linn. Soc. Lond. 155 (1944) 285, Browne, FTSB (1955) 242. Type: Flemmich FMS 32600, Borneo,
Brunei (holotype KEP).
Tree up to 40 m tall, 180 cm diameter, not buttressed. Bark minutely fissured. Stipules
auriculate to subfalcate, up to 15 10 mm. Leaves 4.512 cm long, glabrous. Leaflets
(2)4(6) for each leaf, coriaceous, glossy, pellucid-dotted, glabrous except sometimes
sparsely hairy on the midrib or at the base on the lower surface, glabrescent; ellipticoblong, often unequally sided, 59(14) 3.56(8) cm, base rounded, apex acute or
shortly acuminate; lateral veins many, distinct or visible on both surfaces; often with
13 elliptic, slightly depressed glands towards the basal part of the margin; petiolules
47 mm long. Inflorescences 411 cm long; pedicel 34 mm long. Flowers: calyx lobes
elliptic, 47.5 24 mm, puberulous outside, densely tomentose inside; stamens alternately
long and short. Fruits 4.57.5 3.54 cm. Seeds 2.42.7 c. 1.5 cm, glossy brown
or black; aril narrowly 2-lobed, enclosing the seed.
Vernacular names. Sabahsepetir paya (preferred name). Sarawaksepetir paya (preferred
name), tepih (Melanau, Rejang).
Distribution. Endemic to Borneo. In Sabah uncommon; in Sarawak known from Betong,
Bintangor, Marudi, Miri, and Sibu districts. Also in Brunei and Kalimantan.
Ecology. In peatswamp forest near the coast, sometimes also found in well-drained lowland
forest, once found in secondary heath forest, up to 30 m altitude. Flowering in April and
October; fruiting in July.
Uses. The timber of this species is similar to that of Sindora and can be used for the same
purposes. (cf.: PROSEA 5, 1 (1993) 437).
129
vi
4. CRUDIA Schreber, nom. cons.

(Dr. Crudy, collected plants before 1810 in Bahama Is., St. Thomas and St. Lucia,
who communicated the specimen of the tree to Schreber)
Gen. 1 (1789) 282; Prain, J. As. Soc. Beng. 66, 2 (1897) 218; Ridley, FMP 1 (1922) 643; de Wit, Bull.
Bot. Gard. Buitenz. 3, 18 (1950) 411; Backer & Bakhuizen f., FJ 1 (1964) 527; Whitmore, TFM 1
(1972) 248; Cockburn, TS 1 (1976) 160; Anderson, CLST (1980) 233; Cowan & Polhill in Polhill &
Raven (eds.), Adv. Leg. Syst. 1 (1981) 131; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 198; Ding
Hou, FM 1, 12 (1996) 573; Coode et al. (eds.), CLBD (1996) 162; Argent et al. (eds.), MNDT-CK 2
(1997) 353. Synonym: Pryona Miq., Fl. Ned. Ind. 1, 1 (1858) 1081.
Trees or shrubs, rarely scramblers. Stipules usually intrapetiolar, often connate at the
basal parts, sometimes interpetiolar and free (in taxa outside Sabah and Sarawak), caducous
or persistent. Leaves compound, once pinnate and imparipinnate, each with 19(13)
leaflets; petiole and rachis often prolonged beyond the upper petiolule into a very short
(deciduous) tip. Leaflets alternate, very rarely opposite; lateral veins ascending and
archingly anastomosing towards the margins; intercostal venation often faintly, loosely
reticulate; petiolules short. Inflorescences axillary or terminal, racemose, solitary or
fasciculate; bracts entire or lobed (sometimes 3-lobed with the central lobe redivided),
often caducous, sometimes persistent; bracteoles 2, often minute, caducous; pedicel rather
short. Flowers bisexual; hypanthium shortly cupular or campanulate; calyx lobes 4; petals
absent; stamens 10 or fewer, filaments slightly equal or unequal in length, anthers small;
ovary densely hairy, 16-ovuled, style filiform, stigma terminal, small knob-like. Fruits
(pods) not winged, obliquely orbicular, ellipsoid or ovoid, compressed or somewhat swollen,
dehiscing into 2 coriaceous or slightly woody valves, 14(6)-seeded. Seeds suborbicular
or subreniform, compressed, without aril; cotyledons concave.
Distribution. A tropical genus consisting of about 50 species distributed in America,
Africa, Asia, and Malesia. Ten species occur in Sabah and Sarawak.
Ecology. In lowland forest, sometimes occurring up to 400 m altitude. De Wit (1950:
411) found that the cotyledons are concave with a large inner cavity (also found, e.g.,
in Entada), which may cause buoyancy and promote dispersal.
Key to Crudia species

1. Leaflets usually glabrous, rarely sparsely puberulous on the lower surface.....2
Leaflets pubescent to densely or minutely puberulous, rarely glabrescent and seemingly
glabrous, on the lower surface...........5
2. Stipules falcate, 56 mm long. Bracts at the base of peduncle usually subulate and
long-drawn, sometimes many, aggregate, and brush-like..........5. C. reticulata
Stipules ovate, small or minute, up to 3.5 mm long. Bracts at the base of peduncle
triangular, ovate to lanceolate or linear, not long-drawn or brush-like as above............3
3. Leaves each with 1 or 2 leaflets; petiole and rachis up to 1 cm long. Pedicel 2
4 mm long...............6. C. subsimplicifolia
131
Leaves each with 26(8) leaflets; petiole and rachis 1.514 cm long. Pedicel usually
1522 mm long.....................4
4. Leaves each with 46(8) leaflets; petiole and rachis (6)914 cm long. Fruits oblong,
elliptic, or oblanceolate, 1014 4.55 cm. Seeds irregularly suborbicular, 2.53
3.54 cm..............4. C. ornata
Leaves each with 24 leaflets; petiole and rachis 1.57 cm long. Fruits ellipsoid
or fusiform, 58 2.53.5 cm. Seeds transverse-oblong or slightly reniform, 22.2
33.7(4.7) cm....................7. C. tenuipes
5. Leaves each with (1)2(3) leaflets. Inflorescences or infructescences pendulous, up
to 45 cm long.........1. C. bantamensis
Leaves each with 28 leaflets. Inflorescences or infructescences erect, 415.5(20)
cm long.....6
6. Leaves with petiole and rachis prolonged beyond the upper petiolule into a short
tip...............2. C. beccarii
Leaves with petiole and rachis not prolonged as above....................7
7. Leaves with petiole and rachis 0.51 cm long.....................8. C. velutina
Leaves with petiole and rachis 1.510 cm long.......8
8. Stamens alternately long and short........................9. C. venenosa
Stamens almost equal in length....................9
9. Leaves each with (4)68 leaflets. Pedicel 1015 mm long. Fruits elliptic, ovateoblong or obovate, 710 2.53.5 cm, velvety...........3. C. caudata
Leaves each with 35(7) leaflets. Pedicel 34 mm long. Fruits ovate-oblong or broadly
elliptic, 58 3.54.5 cm, short-hairy....................10. C. wrayi
1. Crudia bantamensis (Hassk.) Benth.

(of Bantam, Java)
Trans. Linn. Soc. 25 (1868) 315; Masamune l.c. 339; Fischer, Kew Bull. (1932) 73; de Wit l.c. (1950)
415; Backer & Bakhuizen f. l.c. 527; Ding Hou l.c. (1996) 577. Basionym: Touchiroa bantamensis
Hassk., Retzia 1 (1885) 202. Lectotype (de Wit, 1950): Hasskarl s.n. (Cult. in Hort. Bog. sub no. I-I-38,
coll. in March 1925), Java, Bantam (holotype BO; isotype L). Synonyms: Pryona bantamensis (Hassk.)
Miq., Fl. Ned. Ind. 1, 1 (1858) 1081; Apalatoa bantamensis (Hassk.) Baill., Hist. Pl. 2 (186970) 104.
Tree up to 16 m tall, 35 cm diameter. Stipules 3.54.5 mm long. Leaves each with

(1)2(3) leaflets (usually one leaflet early shed and two persistent); petiole and rachis
0.52.5 cm long, prolonged beyond the upper petiolule into a very short tip. Leaflets
glabrous above, minutely, loosely, appressedly puberulous, glabrescent (or almost glabrous
except on the midrib) beneath; obovate-oblong or elliptic-oblong, or oblanceolate, 12
24 46.5(9) cm, base acute, obtuse or rotundate, often symmetrical, apex acute, abruptly
or gradually, shortly acuminate, sometimes rounded to subemarginate; lateral veins 5
7 per side; petiolules 1.55 mm, glabrous. Inflorescences axillary, up to 45 cm long,
132
pendulous; rachis puberulous; bracts 23.5 mm long, ovate or triangular, or 3-lobed at

about the upper 1/3 (the middle lobe redivided into a 3-pointed fork), puberulous,
glabrescent; bracteoles 2, small, usually persistent; pedicel 12 mm long, articulated at
the apex, puberulous. Flowers loosely puberulous outside, glabrescent, glabrous inside;
hypanthium shortly cupular, c. 1.5 mm long; calyx lobes elliptic or ovate, 34 1.5
2 mm; stamens 810, filaments up to 12 mm long, glabrous; ovary c. 3 mm long, 3
5-ovuled, style 68 mm long. Fruits oblong, ovate or broadly elliptic, 49 24 cm,
rugose, slightly beaked, dehiscing into 2 crooked, rather thin valves, 14-seeded. Seeds
slightly reniform, or irregularly rounded, 33.75 1.753 cm, rather smooth.
Distribution. W Java (Bantam) and Borneo. In Sabah (once reported, Fischer, 1932);
not yet found in Sarawak. Also in Kalimantan.
Ecology. In lowland forest, up to 400 m altitude, scattered, sometimes locally common.
Flowering in JanuaryFebruary, JulySeptember, and December; fruiting in July, September,
and December.
2. Crudia beccarii Ridl.

Kew Bull. (1929) 257; de Wit l.c. (1950) 418; Anderson l.c. 233; Whitmore, Tantra & Sutisna l.c. 198;
Ding Hou l.c. (1996) 579; Coode et al. (eds.) l.c. 162. Type: Beccari PB 3903, Borneo, Sarawak (holotype
K; isotype FI).
Tree up to 25 m tall, 40 cm diameter. Stipules c. 3 mm long. Leaves each with 3 or

4 leaflets; petiole and rachis 1.53 cm long, prolonged beyond the upper petiolule into
a very short tip. Leaflets glabrous above, minutely appressedly hairy, sometimes glabrescent,
or almost glabrous beneath; obovate, obovate-oblong or elliptic-oblong, or elliptic, 6
14 2.55 cm, base obtuse or rounded, often symmetrical, apex abruptly acuminate;
lateral veins 57 per side; petiolules c. 5 mm long, glabrous. Inflorescences axillary
or terminal, up to 14 cm long, erect; bracts triangular or subulate, 0.754 mm long,
puberulous outside; bracteoles 2, minute, triangular, c. 0.5 mm long; pedicel 23 mm
long, articulated at the base. Flowers slightly puberulous outside; hypanthium c. 1 mm
long, glabrous inside; calyx lobes ovate, broadly elliptic or slightly obovate, 33.5
2 mm, glabrous inside; stamens (9 or)10, filaments 57 mm long; ovary 4(6)-ovuled.
Fruits broadly ellipsoid or slightly obovoid, 3.54.3 2.5 cm, flattened. Seeds subreniform,
c. 1.5 2.5 cm.
Distribution. Endemic to Borneo. In Sarawak, so far known only from two collections:
Beccari PB 3351 and PB 3903, without precise localities; not yet reported in Sabah.
Ecology. In lowland forest, up to 50 m altitude. Flowering in July, fruiting in October.
3. Crudia caudata Prain

(Latin, caudatus = tail-shaped; the leaf apex)
133
J. As. Soc. Beng. 66, 2 (1897) 219 & 488; Merrill l.c. (1921) 296; Ridley l.c. (1922) 644; Masamune l.c.
339; de Wit l.c. (1950) 422; Whitmore l.c. 249; J.E. Vidal, Fl. Thail. 4 (1984) 92; Whitmore, Tantra &
Sutisna l.c. 198; Turner, l.c. 298; Ding Hou l.c. (1996) 581. Type: Ridley 6399, Peninsular Malaysia,
Johore (holotype K, isotype BM). Synonym: C. bracteolosa de Wit l.c. (1950) 421.
Tree up to 25 m tall. Bark grey-brown, only slightly peeling, c. 7 mm thick, red-brown

but inside dull yellow. Stipules triangular to lanceolate, up to 7(13) mm long. Leaves
each with (4)68 leaflets; petiole and rachis up to 10 cm long, not prolonged into a
short tip. Leaflets glossy and glabrous above, rusty pubescent beneath; elliptic-oblong
or obovate-oblong, 79(15) 23(4) cm, base obtuse, rounded, or cuneate, symmetrical,
apex acuminate or caudate; lateral veins 58 per side; petiolules 12 mm long. Inflorescences
axillary or terminal, usually erect, up to 15 cm long; rachis rusty tomentose, rather densely
flowered; bracts ovate to lanceolate, sometimes oblanceolate, (5)79 mm long, pubescent
outside and glabrous inside, some of them 3-lobed and the central lobe imparipinnately
relobed; bracteoles lanceolate or spoon-shaped, 69 mm long, almost enclosing the bud;
pedicel 1015 mm long, pubescent, articulated at the apex. Flowers pubescent outside;
hypanthium 22.5 mm long; calyx lobes ovate, unequally sized, 57 3.55 mm, glabrous
inside; stamens 10, or rarely 9, filaments almost equal in length, up to 15 mm long;
ovary (4)6-ovuled, style 817 mm long. Fruits (unripe) elliptic, ovate-oblong, or obovate,
710 2.53.5 cm, flat, rusty, velvety, usually 1-seeded, valves thin-coriaceous. Seeds
shortly oblong, broadly elliptic, or rounded, flat, 34 23 cm.
Distribution. Peninsular Thailand, Peninsular Malaysia and Borneo. In Sarawak (no precise
localities) known from the following collections: Beccari PB 1763 and PB 3711; Foxworthy
306; not yet found in Sabah. Also in Brunei and W Kalimantan.
Ecology. In riverine and freshwater swamp forest, at low altitude. Fruiting in JuneJuly.
4. Crudia ornata de Wit
Fig. 4.
(Latin, ornatus = adorned or embellished; inference unknown)

Bull. Bot. Gard. Buitenz. 3, 18 (1950) 425; Cockburn l.c. 160; Whitmore, Tantra & Sutisna l.c. 199;
Ding Hou l.c. (1996) 587; Argent et al. (eds.) l.c. 352. Type: Elmer 20708, Borneo, Sabah, near Tawau
(holotype BO; isotypes BM, K, L). Synonym: C. reticulata auct. non Merr. (1918): Merrill l.c. (1929)
99.
Tree up to 20 m tall, 30 cm diameter, occasionally as shrub-like tree or treelet up to

1.5 m high. Stipules ovate, 23 mm long. Leaves each with 46(8) leaflets; petiole
and rachis (6)914 cm long, minutely puberulous. Leaflets glabrous above, rarely minutely
puberulous, glabrescent, or glabrous beneath; elliptic-oblong, obovate-oblong or ovateoblong, or elliptic, (7.5)1420 (3.5)4.87.5 cm, base cuneate, acute, obtuse, or rounded,
symmetrical, apex cuspidate or rather abruptly acuminate; lateral veins 58 per side;
petiolules 59 mm long. Inflorescences axillary, sometimes also terminal, up to 15.5
cm long, erect; rachis tomentose; bracts triangular, ovate, lanceolate, or linear, 1015
mm long, puberulous, glabrescent; bracteoles elliptic, lanceolate, or oblanceolate, 68
mm long, often at the upper half of pedicel, covering the bud at its very young stage;
pedicel 1520 mm long, puberulous. Flowers puberulous outside; hypanthium 23 mm
long, glabrous inside; calyx lobes unequal, broadly elliptic or elliptic, or obovate, 6
134
vi
7 34 mm, slightly loosely, appressedly puberulous often on the lower half inside,
glabrescent, or almost glabrous; stamens 10(or 9), filaments up to 15 mm long, glabrous;
ovary (4)6-ovuled, style up to 8 mm long. Fruits oblong, elliptic, or oblanceolate, 10
14 4.55 cm, shortly hairy, 1several-seeded, valves hard, coriaceous. Seeds irregularly
suborbicular or subreniform, 2.53 3.54 cm, slightly finely rugose.
Vernacular names. Sabahkelat (Malay, Timbun Mata Is.), sansag (Kinabatangan).
Distribution. Endemic to Borneo. In Sabah known from Sandakan, Kota Kinabalu, Tenom,
Labuk Sugut, Kinabatangan, Semporna (Timbun Mata Is.), and Tawau districts. In Sarawak
recorded from the 5th Division. Also in Kalimantan.
Ecology. In primary forests, from lowland up to 200 m altitude, sometimes occurring
on sandy soils and limestone. Flowering in JanuaryFebruary, April, JulyAugust, and
November; fruiting in April and JulyAugust.
5. Crudia reticulata Merr.

(Latin, reticulatus = reticulate; the intercostal venation of the leaflets)
Philip. J. Sci. 13 (1918) 73, l.c. (1921) 296; Masamune l.c. 340; de Wit l.c. (1950) 426; Cockburn l.c.
160; Anderson l.c. 233; Whitmore, Tantra & Sutisna l.c. 199; Ding Hou l.c. (1996) 589; Argent et al.
(eds.) l.c. 353. Type: Villamil 393, Borneo, Sabah, Sandakan (isotype K).
Tree up to 18 m tall, 20 cm diameter, sometimes shrub-like up to 4 m tall, rarely a

climber (?). Stipules falcate, 56 mm long, caducous. Leaves each with (1)23(4) leaflets;
petiole and rachis 0.52(4) cm long. Leaflets glabrous or rarely sparsely minutely hairy
below; elliptic, elliptic-oblong, ovate-oblong or obovate-oblong, 1029(36) 38(12.5)
cm, base cuneate, acute, obtuse, or rounded, symmetrical or slightly asymmetrical, apex
acuminate or rather abruptly caudate-acuminate, acumen slender, 1.52.5(3.5) cm long;
lateral veins 59 (or more) per side; petiolules 37 mm long. Inflorescences axillary,
up to 10(28) cm long, rachis puberulous; bracts subulate and long-drawn, or 3-lobed
with the central lobe divided again and bristle-like, slightly hairy, up to 8 mm long,
sometimes many, aggregate, surrounding the base of the peduncle and brush-like,
(semi-) persistent, puberulous outside, slightly ciliate on the margins, those at the
base of pedicel rather small, triangular, 0.31.5 mm long; bracteoles minute; pedicel
35 mm long. Flowers puberulous outside, glabrous inside; hypanthium c. 1 mm long;
calyx lobes ovate or broadly ovate, 3.54 22.5 mm; stamens (8)10, filaments usually
46 mm long; ovary usually 4-ovuled, style 4.55.5(10) mm long. Fruits (young) ellipsoid
or ovoid, 8.512.5 34 cm, densely shortly hairy. Seeds broadly ellipsoid or suborbicular,
2.63.3 22.4 cm; testa rusty scurfy, finely rugose.
Vernacular names. Sabahangar-angar (preferred name), maguwau (Dusun, Lahad Datu),
selangapid (Sandakan), tanggar-angor (Kinabatangan).
Distribution. Endemic to Borneo. In Sabah known for Sandakan (Sepilok), Lahad Datu,
Kinabatangan (Lamag) and Tawau districts. In Sarawak recorded for northern region,
Bintulu and Tatau districts. Also in Kalimantan.
136
Ecology. In lowland primary mixed dipterocarp, periodically inundated freshwater swamp

and peatswamp forests on sandy, loamy soils with coral limestone, occasionally in secondary
forest, up to 250 m, once recorded at 1000 m altitude (Berau, Kalimantan). Flowering
in JanuaryApril and JulySeptember; fruiting in April and JuneOctober.
6. Crudia subsimplicifolia Merr.

(Latin, sub- = somewhat, simplici- = simple, -folius = -leaved; tree with somewhat simple
leaves)
Philip. J. Sci. 5 (1910) Bot. 39; de Wit l.c. (1950) 431; Ding Hou l.c. (1996) 592. Type: Klemme FB
4287, the Philippines, Luzon (isotype K).
Tree up to 15 m tall, 40 cm diameter. Stipules ovate, c. 3.5 mm long, caducous. Leaves

each with 1 or 2 leaflets, when 2 never opposite; petiole and rachis up to 1 cm long.
Leaflets glabrous or rarely sparsely minutely hairy below; elliptic, elliptic-oblong or
obovate-oblong, or lanceolate, 9.521(29) 3.57(9) cm, base acute or obtuse, symmetrical,
apex shortly acuminate or acuminate; lateral veins 69 per side; petiolules 23.5 mm
long. Inflorescences axillary or sometimes terminal, up to 11(16) cm long, glabrous;
bracts triangular or 3-lobed with the central lobe relobed and bristle-like, up to 5 mm
long, glabrous except ciliate at the margins; bracteoles often at the lower half or at the
base of the pedicel, minute; pedicel 24 mm long, glabrous. Flowers glabrous outside
except ciliate at the margins of the calyx lobes; hypanthium c. 1 mm long, sparsely hairy
inside; calyx lobes ovate or broadly elliptic, c. 4 2.53 mm, glabrous inside; stamens
10, filaments 46 mm long; ovary 34-ovuled, style up to 10 mm long. Fruits (very
young) elliptic, c. 0.6 0.3 cm.
Vernacular names. Sabahmaid maid, sarason (Lahad Datu).
Distribution. Borneo, the Philippines and Aru Is. In Sabah known in Kinabatangan (Pin
R. and Tongod), Sandakan, Keningau, Lahad Datu, Semporna, Tawau, and Tenom districts.
In Sarawak recorded for the 3rd Division. Also in E Kalimantan.
Ecology. In primary mixed dipterocarp forest on ridges, low undulating lands, or near
the rivers, up to 300(900) m altitude. Flowering in JanuaryApril, June, and October;
fruiting in April.
7. Crudia tenuipes Merr.

(Latin, tenui- = slender, thin, pes = foot; flower with slender pedicel)
Philip. J. Sc. 11 (1916) Bot. 83, l.c. (1921) 296; Masamune l.c. 340; de Wit l.c. (1950) 430; Cockburn
l.c. 160; Anderson l.c. 233; Whitmore, Tantra & Sutisna l.c. 199; Ding Hou l.c. (1996) 593; Coode et al.
(eds.) l.c. 162; Argent et al. (eds.) l.c. 354. Type: Native Collector (Bur. Sci.) 1358 (err. as 1385),
Borneo, Sarawak (isotype K).
Tree up to 27 m tall, 40 cm diameter; buttresses steep, up to 1.5 m high and 30 cm

extending outwards from the base. Stipules ovate, minute, caducous. Leaves each with
24 leaflets; petiole and rachis 1.57 cm long. Leaflets glabrous on both surfaces; elliptic137
oblong or ovate-oblong, 720 37 cm, base cuneate, acute, obtuse, or rounded,

symmetrical or slightly asymmetrical, apex acuminate or rather abruptly acuminate; lateral
veins 57 per side; petiolules c. 5 mm long, glabrous. Inflorescences axillary, up to
24 cm long, glabrous; bracts triangular, up to 1.5 mm long, glabrous, caducous; bracteoles
minute, caducous; pedicel (7)1522(30) mm long, glabrous. Flowers glabrous outside;
hypanthium c. 2 mm long; calyx lobes elliptic or ovate-oblong, 44.5 22.5 mm, inner
surface often puberulous on the vertical, central part, glabrescent, sometimes almost
glabrous; stamens (8)10, filaments up to 12 mm long, glabrous; ovary (2)4(5)-ovuled,
style up to 14 mm long. Fruits ellipsoid or fusiform, 58 2.53.5 1.82.5 cm, swollen,
gradually narrowed from the middle towards both ends, shortly hairy, 12(more)-seeded;
valves coriaceous, hard. Seeds transverse-oblong or slightly reniform, 22.2 33.7
(4.7) 1.82.3 cm, slightly swollen, smooth, shining.
Distribution. Endemic to Borneo. In Sabah known in Sandakan, Beaufort, Kota Kinabalu,
Lahad Datu, Tenom, and Tawau districts. In Sarawak recorded for Kapit, Kuching, Marudi
(Mt. Dulit), and Miri districts. Also in Brunei and Kalimantan.
Ecology. Widespread in forests; sometimes found on limestone, coral rock, sandy yellow
loam soils. From lowland up to 500 m altitude. Flowering in FebruaryMay, and October
November; fruiting in AprilMay, JulySeptember, and December.
8. Crudia velutina Ridl.

(Latin, velutinus = velvety; some parts of the plant covered with fine short hairs)
Kew Bull. (1929) 257; de Wit l.c. (1950) 432; Whitmore l.c. 252; Anderson l.c. 233; Whitmore, Tantra
& Sutisna l.c. 199; Turner l.c. 299; Ding Hou l.c. (1996) 594. Type: Haviland 3071, Borneo, Sarawak,
Sri Aman district (holotype K; isotypes BM, L).
Tree up to 30 m tall, 70 cm diameter; buttresses up to 2.4 m high. Bark smooth to

dippled; inner bark brown, laminated. Sapwood yellowish brown. Stipules subulate, 2
3 mm long, deciduous. Leaves each with 23(4) leaflets; petiole and rachis 0.51 cm
long, brown puberulous, glabrescent. Leaflets upper surface puberulous when young,
glabrescent, or almost glabrous when old, lower surface glaucous, finely, uniformly,
appressedly, rather densely puberulous (when young); obovate, obovate-oblong or ellipticoblong, or elliptic, 8.513.5(18) 3.56(8) cm, base obtuse or rounded, often symmetrical,
apex shortly bluntly acuminate, or acuminate; lateral veins c. 5 per side; petiolules c.
5 mm long. Inflorescences axillary or terminal, up to 12 cm long, erect; rachis densely
puberulous; bracts small, triangular, c. 1 mm long, puberulous outside, ciliate on the margins;
bracteoles minute; flowers subsessile or pedicel 12 mm long. Flowers minutely, appressedly
puberulous outside; hypanthium c. 1 mm long, glabrous inside; calyx lobes ovate or elliptic,
35 22.5 mm, glabrous inside, ciliate on the margins; stamens (9 or)10, filaments
37(10) mm long; ovary (3)4(5)-ovuled, style 67 mm long. Fruits not seen.
Distribution. Peninsular Malaysia and Borneo. In Sarawak known only from Bt. Kalong
in Sri Aman district; not yet found in Sabah.
Ecology. In forests on undulating lands, a few meters above sea level. Flowering in April
and May.
138
9. Crudia venenosa de Wit

(Latin, venenosus = poisonous; poisonous plant)
Bull. Bot. Gard. Buitenz. 3, 18 (1950) 433; Whitmore, Tantra & Sutisna l.c. 199; Ding Hou l.c. (1996)
595. Type: Maidin (SAN A) 1692, Borneo, Sabah, Sg. Kinabatangan (holotype BO; isotype K).
Tree up to 8 m tall. Stipules caducous. Leaves each with 46 leaflets; petiole and rachis
6.510 cm long, or longer. Leaflets glabrous above, sparsely minutely puberulous below;
elliptic, elliptic-oblong or obovate-oblong, 1118 56.8 cm, base rounded, symmetrical,
apex abruptly shortly acuminate; lateral veins 58 per side; petiolules distinct, c. 7 mm
long. Inflorescences axillary or terminal, erect, up to 10 cm long; rachis puberulous;
bracts triangular, 0.52.5 mm long, glabrous except ciliate at the margins; bracteoles minute;
pedicel 26 mm long. Flowers puberulous outside; hypanthium 12 mm long; calyx lobes
elliptic or ovate, 34 2 mm; stamens 10, alternately long and short; ovary 34-ovuled.
Fruits not seen.
Vernacular name. Sabahsansag (Kinabatangan).
Distribution. Endemic to Borneo. In Sabah known only by the type collection from Sg.
Kinabatangan; not yet found in Sarawak.
Ecology. In swamp forest. Flowering in March.
10. Crudia wrayi Prain

(L. Wray, Jr., 18531942, sometime Superintendent of the Larut Hill Garden at Taiping,
Curator of the Perak State Museum, and Director of Museums of the Federated Malay
States)
J. As. Soc. Beng. 66, 2 (1897) 222 & 488; Ridley l.c. (1922) 646; Whitmore l.c. 252; Anderson l.c. 233;
Whitmore, Tantra & Sutisna l.c. 199; Turner l.c. 299; Ding Hou l.c. (1996) 596. Type: Wray Jr. 2874,
Peninsular Malaysia, Perak, Larut (holotype K). Synonym: C. havilandii Prain, J. As. Soc. Beng. 73, 2
(1904) 199, de Wit l.c. (1950) 423.
Tree up to 32 m tall, 45 cm diameter, with fluted bole; buttresses steep up to 3.6 m

high, 300 cm wide, 5 cm thick. Bark greyish brown, smooth to flaky; inner bark light
brownish green. Stipules lanceolate, 23.5 mm long, deciduous. Leaves each with 3
5(7) leaflets; petiole and rachis 1.54(6.5) cm long, loosely puberulous, glabrescent,
or glabrous. Leaflets sparsely puberulous or glabrous above, loosely, minutely puberulous
beneath; elliptic, ovate-elliptic or obovate-elliptic, (3.5)5.513 (1.3)2.55 cm, base
obtuse, or cuneate, symmetrical, apex acuminate or caudate; lateral veins 48 per side;
petiolules 25 mm long. Inflorescences axillary or terminal, up to 9(20) cm long, erect;
rachis puberulous; bracts deltoid or 3-lobed, up to 4 mm long, sparsely puberulous outside,
caducous; bracteoles minute; pedicel 34 mm long. Flowers sparsely puberulous outside;
hypanthium c. 0.6 mm long, glabrous inside; calyx lobes unequal, broadly elliptic, elliptic,
or oblong, 3.54 2.53 mm, glabrous inside; stamens (8)10, filaments almost equal
in length, up to 11 mm long; ovary c. 3.5 mm long, 2(4)-ovuled, style c. 8 mm long,
139
glabrous. Fruits ovate-oblong or broadly elliptic, 58 3.54.5 cm, short-hairy, 1 (more)seeded. Seeds broadly obovate, c. 3.5 2.5 cm.
Distribution. Peninsular Malaysia and Borneo. In Sarawak known from Kuching district;
not yet found in Sabah. Also in Kalimantan.
Ecology. In primary lowland mixed dipterocarp forest, riverine and swamp forest, and
once recorded at the base of limestone hill at Bau (near Kuching), up to 90 m altitude.
Flowering in SeptemberOctober and December; fruiting in January, March and September.
5. CYNOMETRA L.
(Greek, kuon = dog, metra = female genitalia; referring to the shape of the fruit)
katong-katong, kekatong (Malay)
Sp. Pl. (1753) 382; Prain, J. As. Soc. Beng. 66, 2 (1897) 196; Ridley, FMP 1 (1922) 635; Backer &
Bakhuizen f., FJ 1 (1964) 525; Knaap-van Meeuwen, Blumea 18 (1970) 12; Whitmore, TFM 1 (1972)
253; Cockburn, TS 1 (1976) 160; Verdcourt, Lae Bot. Bull. 11 (1979) 77; Anderson, CLTS (1980) 233;
Cowan & Polhill in Polhill & Raven (eds.), Adv. Leg. Syst. 1 (1981) 124; Corner, WSTM 3rd. edition 1
(1988) 433; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 200; Ding Hou, FM 1, 12 (1996) 597; Loo &
Tan, Gard. Bull. Sing. 49, 1 (1997) 72; Argent et al. (eds.), MNDT-CK 2 (1997) 354.
Trees or shrubs. Buds small, with numerous brown scales in two ranks. New leaves
developing in bright pink tassels. Stipules caducous. Leaves compound, once pinnate
and paripinnate, each with 1 or 2 pairs of asymmetrical leaflets. Leaflets opposite,
asymmetrical, entire and mostly glabrous. Inflorescences axillary, paniculate, racemose,
or spike-like, solitary or sometimes 2 per axil, rarely borne on older branches and stems
and fasciculate, mostly spherical in outline, sometimes elongated, sessile, densely flowered;
bracts scale-like, mostly persistent, appressed hairy, often glabrescent. Flowers bisexual;
hypanthium usually shortly campanulate, sometimes obscure or absent; calyx lobes 4(
5); petals 5(or 4), glabrous; fertile stamens (8)10(15), equal in length, filaments free,
rarely united at the base, usually glabrous; ovary 12(4)-ovuled. Fruits usually thickened,
broad ovoid or broad ellipsoid, less than 8 cm long, indehiscent, often flattened especially
when young, smooth or rugose, sometimes warty, 1(4)-seeded. Seeds without endosperm.
Distribution. A pantropical genus comprising about 70 species; 3 species in Sabah and
Sarawak.
Ecology. In forests, sometimes growing on banks of rivers or streams, or in coastal areas,
from lowland up to 1300 m altitude.
Key to Cynometra species

1. Leaves with only 1 pair of leaflets. Rachis of inflorescence c. 4.5 cm long........1. C. elmeri
Leaves with 1 or 2 pairs of leaflets. Rachis of inflorescence less than 4 cm long.....2
140
2. Inflorescences borne on the trunk; pedicel of flowers 715 mm long. Fruits deeply
rugose............3. C. ramiflora
Inflorescences axillary; pedicel of flowers 1826 mm long. Fruits smooth.
....2. C. mirabilis
1. Cynometra elmeri Merr.
Fig. 5AB.
(A.D.E. Elmer, 18701942, plant collector in the Philippines and Borneo)

PEB (1929) 98; Knaap-van Meeuwen l.c. 29; Meijer, Field Guide Tree W. Mal. (1974) 181; Cockburn
l.c. 162; Whitmore, Tantra & Sutisna l.c. 200; Ding Hou l.c. (1996) 602. Type: Elmer 21386 (err. as
21368), Borneo, Sabah, Tawau (isotypes L, U).
Tree up to 25 m tall. Leaves each with 1 pair of leaflets; petiole 0.40.8 cm long. Leaflets
ovate-oblong or oblong, more or less falcate, 3.515 1.34 cm, base cuneate, apex
acuminate, acumen blunt and emarginate; lateral veins 57 pairs. Flowers unknown. Rachis
of infructescence c. 4.5 cm long. Fruits pendulous, flat, 46 2.53.5 cm, inner suture
more or less straight, outer suture widely curved, with a small beak on the inner suture
side of the apex, surface rough.
Distribution. Borneo and the Philippines. In Sabah found in Tawau and Kinabatangan
districts, not yet reported in Sarawak.
Ecology. In back mangrove and freshwater swamp forests, up to 300 m altitude.
2. Cynometra mirabilis Meeuwen
Fig. 5CD.
(Latin, mirari = wonderful, extraordinary; inference unknown)

Blumea 18 (1970) 25; Cockburn l.c. 162; Whitmore, Tantra & Sutisna l.c. 200; Ding Hou l.c. (1996)
605. Type: Miranda FB 20528, the Philippines (holotype L; isotypes BO, UC, US).
Tree up to 35 m tall, 80 cm diameter. Bark pale yellow-grey, smooth, later lenticellate;

inner bark pink-brown, thin. Leaves each with 1 or 2 pairs of leaflets. Leaflets (ob)ovateoblong to elliptic or oblong, 6.512.5 35 cm, base cuneate, apex acuminate, sometimes
slightly emarginate; midrib 1018 mm from the upper margins; lateral veins c. 7 pairs;
petiole 510 mm long. Inflorescences axillary, rachis 0.61 cm long, glabrous or with
a few hairs; bracts up to 11 mm long; pedicel 1826 mm long, glabrescent. Flowers:
hypanthium 11.5 mm long; calyx lobes 56 mm long, glabrous; petals c. 7 mm long;
stamens occasionally with a few hairs; ovary woolly or only with a few hairs outside,
inside wall lined with curly appressed hairs. Fruits (immature) obovoid, 3 1.7 0.2
cm, smooth.
Vernacular name. Sabahkatong-katong (preferred name).
Distribution. Borneo and the Philippines. In Sabah known from Sandakan, Lahad Datu,
Semporna, Tawau, Kinabatangan, and Kudat (Banggi Is.) districts; not known in Sarawak.
141
Ecology. Often found in inland forest, also occurring near the seashore or along rivers,
up to 150(600) m altitude. Flowering in May, August and October; fruiting in April.
3. Cynometra ramiflora L.
Fig. 5EF.
(Latin, ramus = branch, flos = flower; flowers borne on branches)

Sp. Pl. (1753) 382, excl. syn. Iripa Rheede; Prain l.c. (1897) 197; Merrill, Int. Rumph. Herb. Amb.
(1917) 254; Ridley l.c. (1922) 635; Browne, FTSB (1955) 223); Backer & Bakhuizen f. l.c. 526; Knaapvan Meeuwen l.c. 23; Whitmore l.c. 254; H. Keng l.c. (1974) 258; Cockburn l.c. 162; Verdcourt l.c. 82;
Anderson l.c. 233; K. Larsen & S.S. Larsen l.c. (1980) 118, l.c. (1984) 48; Corner l.c. 434; Keng l.c.
(1990) 34; Whitmore, Tantra & Sutisna l.c. 200; Turner l.c. 299; Ding Hou l.c. (1996) 606; Argent et al.
(eds.) l.c. 354. Type: Cynomorium sylvestre Rumph., Herb. Amb. 1 (1741) t. 63. Synonym: C. bijuga
Span. ex Miq., Fl. Ind. Bat. 1, 1 (1855) 78, Merrill l.c. (1921) 295, Masamune l.c. 340.
Distribution. A species consisting of two varieties distributed in India, throughout SE

Asia and Malesia to the Pacific. In Sabah and Sarawak only var. ramiflora is known.
Another one, var. bifoliolata (Merr.) Meeuwen, occurs in the Philippines.
var. ramiflora
Ding Hou, FM 1, 12 (1996) 606; Loo & Tan l.c. 73. Synonyms: C. ramiflora L., Sp. Pl. (1753) 382,
excl. syn. Iripa Rheede; C. bijuga Span. ex Miq., Fl. Ned. Ind. 1, 1 (1855) 78, Merrill l.c. (1921) 295; C.
whitfordii Elmer, Leafl. Philipp. Bot. 8 (1915) 2734.
Tree up to 26 m tall. Leaves each with 1 and/or 2 pairs of leaflets, rarely with 2 pairs
only; petiole 0.31.5 cm long, rachis 1.54 cm long, both sometimes patent-hairy,
glabrescent, or glabrous. Leaflets ovate, ovate-oblong, oblong, elliptic, lanceolate, rarely
orbicular, (1.2)4.5(6.5)20 (0.5)2.5(3.5)7 cm, base cuneate, apex acute or acuminate;
lateral veins (4)816 pairs; petiolules indistinct. Inflorescences borne on the trunks,
1.32.5 cm long, usually glabrescent, sometimes glabrous; pedicel 715 mm long. Flowers:
hypanthium 11.25 mm long; calyx lobes 46 mm long; petals lanceolate, 58 mm long;
stamens 10; ovary densely, often curly hairy, 1(2)-ovuled. Fruits broadly ovoid or broadly
ellipsoid, the tip pointing up, thick-woody, deeply rugose, shortly brown scurfy, 2.25
1.34 cm.
Vernacular name. Sabahkatong-katong (preferred name).
Distribution. From India throughout SE Asia and Malesia to the Pacific. In Sabah in
coastal area and Mt. Silam in Lahad Datu district; in Sarawak throughout.
Ecology. In forest just behind mangrove; also occurring in inland forests, up to 400 m
altitude.
Uses. The dark brown timber is heavy and hard but not very durable. It is only available
in small quantity and is used in local construction.
142
vi
143
6. DIALIUM L.
(Latinised old Greek plant name dialion, used for a kind of heliotrope)
keranji (Malay)
Mant. 1 (1767) 3; Prain, J. As. Soc. Beng. 66, 2 (1897) 167; Ridley, FMP 1 (1922) 621; Backer
& Bakhuizen f., FJ 1 (1964) 535; Whitmore, TFM 1 (1972) 255; Cockburn, TS 1 (1976) 164;
Anderson, CLTS (1980) 233; Irwin & Barneby in Polhill & Raven (eds.), Adv. Leg. Syst. 1 (1981)
101; Corner, WSTM 3rd. edition 1 (1988) 436; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990)
200; Rojo, FM 1, 12 (1996) 608; Coode et al. (eds.), CLBD (1996) 162; Loo & Tan, Gard. Bull.
Sing. 49, 1 (1997) 75; Argent et al. (eds.), MNDT-CK 2 (1997) 354. Synonyms: Dansera Steenis,
Bull. Bot. Gard. Buitenz. 3, 17 (1948) 413; Dialium subgen. Dansera (Steenis) Steyaert, Reinwardtia
2 (1953) 355.
Trees; bole columnar, often slightly twisted especially above; buttresses steep, plank-like
spreading. Bark usually pale brown or grey, often rugose. Stipules small, caducous. Leaves
compound, once pinnate and imparipinnate, each with (1)3 or more leaflets, rarely
unifoliolate. Leaflets shortly stalked, often alternate, entire, more or less glabrous above,
glabrous or velvety beneath. Inflorescences terminal or axillary panicles, often in fascicles,
usually hairy. Flowers bisexual, bilaterally symmetrical; hypanthium concave or narrow,
sometimes flattish; calyx lobes 5 or 3, minutely hairy on both surfaces; petals 0; stamens
generally 2 or 6, anthers large, basifixed, sparsely hairy on both sides; ovary densely
hairy, (1)2-ovuled. Fruits drupaceous, not winged, ellipsoid, ovoid, obovoid, subglobose,
or globose, sometimes slightly compressed, indehiscent, often brown to black, shining;
exocarp thin crustaceous, usually hairy. Seeds usually 1 per fruit, smooth, surrounded
by a sweet edible pulp; endosperm present.
Distribution. A genus consisting of 27 species, distributed in tropical Africa, S America,
Asia, and Malesia (not found in the eastern parts of Borneo and Java). Five species in
Sabah and Sarawak.
Ecology. In well-drained primary and sometimes also in old secondary forests, scattered,
at low and medium altitudes up to 1200 m.
Uses. The timber is very strong and hard, and suitable for making excellent tool handles,
planes, furniture and for use in heavy construction. A little pulp surrounding the seed
is edible, and the fruits are sold in village markets throughout Borneo. See Browne, FTSB
(1955) 224226; Burkill, EPMP (1966) 810813; PROSEA 5, 1 (1993) 161166.
Key to Dialium species

1. Leaves unifoliolate.......................5. D. procerum
Leaves pinnate, each with 3 or more leaflets.....................2
2. Leaves each usually with 35 leaflets; often mixed with unifiolate leaves.....
......3. D. kunstleri
Leaves each with 717 leaflets.....................3
144
3. Leaves each usually with 1517 leaflets. Ovary inserted on a flattish hypanthium..
......1. D. hydnocarpoides
Leaves each often with 713 leaflets. Ovary inserted on a narrow or concave
hypanthium..............4
4. Hypanthium concave. Anthers V-channelled on the abaxial side; filaments flattish.
......4. D. platysepalum
Hypanthium narrow. Anthers not as above; filaments more or less subulate...
....2. D. indum
1. Dialium hydnocarpoides de Wit

(Greek, hudnon = truffle, karpos = fruit, -oides = like; with truffle-like fruit)
Blumea 7 (1953) 320; Whitmore, Tantra & Sutisna l.c. 200; Rojo l.c. (1996) 609; Argent et al. (eds.) l.c.
355. Type: NIFS 185T. 3P. 541, Sumatra, Palembang (holotype L; isotype BO). Synonym: D.
praetermissum de Wit l.c. (1953) 321.
Tree up to 30 m tall, 45 cm diameter; buttresses relatively high. Leaves up to 20(24)

cm long, pinnate, each usually with 1517 leaflets. Leaflets subcoriaceous, brownish
hairy beneath, glabrescent; elliptic-oblong, ovate, 3.56(8.5) 23(3.5) cm, base usually
rounded, apex rounded, obtuse, or acute; lateral veins 69 pairs, rather fine; intercostal
venation reticulate, indistinct. Inflorescences terminal; rachis usually 915 cm long, densely
brownish hairy; pedicel 12 mm long. Flowers: calyx lobes 5, ovate-lanceolate, 3.5
4 1.52.5 mm; stamens 2, rarely 3, filaments 11.5 mm long, anthers 2.53 mm long,
connective hairy on both sides; ovary inserted on a narrow, patent-hairy, flattish hypanthium,
1.52 mm long. Fruits subglobose, sometimes slightly compressed, 1.52 cm diameter,
densely hairy. Seeds orbicular, c. 0.9 0.8 cm, shining.
Distribution. Sumatra and Borneo. In Sabah, rare, known in Tawau district; not recorded
in Sarawak. Also in Kalimantan.
Ecology. In primary forest at low altitude, rather rare and scattered. Flowering in January
July; fruiting in AprilJune.
2. Dialium indum L.
(Latin of the Indies)
Mant. 1 (1767) 24; Prain l.c. (1897) 169; Ridley l.c. (1922) 622; Backer & Bakhuizen f. l.c. 535; Whitmore
l.c. 259; Cockburn l.c. 167; Anderson l.c. 234; Whitmore, Tantra & Sutisna l.c. 201; Turner l.c. 302;
Rojo l.c. 609; Coode et al. (eds.) l.c. (1996) 162; Loo & Tan, l.c. 76; Argent et al. (eds.) l.c. 355. Type:
Linn. Herb. 23.1, no exact locality (holotype LINN).
Tree up to 40 m tall, 100 cm diameter; buttresses up to 3 m high. Leaves pinnate, each

usually with 79 leaflets; petiole and rachis usually 715 cm long. Leaflets subcoriaceous
or stiff-coriaceous, glabrous above, slightly hairy, glabrescent beneath; ovate-oblong, ovatelanceolate, broadly elliptic, sometimes broadly lanceolate, 610 35(7) cm, base rounded
145
or cuneate, apex long-acuminate to cuspidate, obtuse or acute; lateral veins 810(14)

pairs; intercostal venation fine, distinct beneath. Inflorescences terminal, rachis 1020
cm long; pedicel 26 mm long. Flowers: calyx lobes 5, ovate-oblong or elliptic, c. 5
2.5 mm; stamens 2, filaments more or less subulate, anthers oblong, c. 4.5 mm long,
connective sparsely hairy on both sides; ovary white hairy, hypanthium narrow. Fruits
globose or ovoid, 1.52 11.5 cm, hairy. Seeds squarish or reniform, 0.71.2 0.5
cm.
Vernacular name. Sabahkeranji (preferred name).
Distribution. A species with 2 varieties distributed in Southernmost Thailand, Sumatra,
Peninsular Malaysia, Singapore, Borneo, and Java. Both of the varieties are known for
Sabah and Sarawak.
Key to varieties
Leaflets ovate-oblong or ovate-lanceolate; apex usually long-acuminate to cuspidate.....
var. indum
Turner l.c. 302; Rojo l.c. 611; Coode et al. (eds.) l.c. 162; Loo & Tan l.c. 77. Synonyms: D. indum
L., Mant. 1 (1767) 24; D. patens Baker in Hooker f., Fl. Brit. Ind. 2 (1878) 270, Ridley l.c. (1922)
622, Masamune l.c. 344, Whitmore l.c. 260; D. marginatum de Wit l.c. (1953) 320; D. turbinatum
de Wit l.c. (1953) 320.
Southern Thailand, Sumatra, Peninsular Malaysia, Singapore, Borneo, and Java.

Common in Sabah and Sarawak; also in Brunei. In lowland well-drained forests,
sometimes in swampy areas, up to 1200 m altitude.
Leaflets broadly elliptic, sometimes broadly lanceolate; apex obtuse or abruptly
acuminate...........
var. bursa (de Wit) Rojo
(Latin, bursa = purse; inference unknown)
FM 1, 12 (1996) 612; Turner l.c. 302; Coode et al. (eds.) l.c. 162; Loo & Tan l.c. 77. Basionym:
Dialium laurinum Baker var. bursa de Wit l.c. (1953) 320. Type: NIFS. 186T. 1P. 197, Sumatra,
Palembang (holotype L; isotype K).
Sumatra, Peninsular Malaysia, Singapore, and Borneo. Known in Sabah, Sarawak and
Brunei. In low, well-drained areas and hills, up to 1200 m altitude.
3. Dialium kunstleri Prain

(H.H. Kunstler, Kings Collector in Perak c. 1880)
J. As. Soc. Beng. 65, 2 (1897) 168 & 484; Ridley l.c. (1922) 621; Whitmore l.c. 259; Whitmore, Tantra
& Sutisna l.c. 201; Turner l.c. 302; Rojo l.c. 613; Coode et al. (eds.) l.c. 163; Argent et al. (eds.) l.c. 355.
Type: Kunstler 4415, Peninsular Malaysia (holotype K; isotype BM).
Tree up to 40 m tall, 90 cm diameter. Leaves pinnate, each with 35(7) leaflets, often
mixed with unifoliolate leaves; petiole and rachis up to 5 cm long. Leaflets coriaceous,
sparsely hairy, glabrescent beneath; usually elliptic, elliptic-oblong, or ovate to lanceolate,
515(19) 36(7.5) cm, base cuneate or rounded, apex long-acuminate or acute; lateral
veins often 68 pairs, rather distinct; petiolules 34(7) mm long. Inflorescences usually
146
axillary, rarely terminal, up to 12(20) cm long; pedicel 34 mm long. Flowers: calyx

lobes 3, elliptic-oblong, c. 3.5 22.5 mm, minutely hairy inside; stamens 6, anthers
linear, 33.5 mm long, sometimes hairy along both sides of the connective; ovary c. 1.5
mm long, whitish hairy. Fruits ellipsoid or ovoid, 34 2.53 cm, slightly hairy, becoming
glabrous. Seeds suborbicular, flattened, c. 1.5 cm across.
Distribution. A species with 2 varieties distributed in Peninsular Malaysia and Borneo.
Both of the varieties occur in Sabah and Sarawak.
Key to varieties
Leaflets usually 5, sometimes mixed with 3 or 7......................
var. kunstleri
Turner l.c. 302; Rojo l.c. 614; Coode et al. (eds.) l.c. 163. Synonyms: D. silvestre de Wit l.c. (1953)
321, Cockburn l.c. 167.
Peninsular Malaysia and Borneo. Common in Sabah and Sarawak; also in Brunei.
In primary forests, up to 250 m altitude, occurring in kerangas forest in Sarawak.
Leaflets 3, often associated with unifoliolate leaves.....................
var. trifoliolatum (de Wit) Rojo
(Latin, with three leaflets)
l.c. 614; Turner l.c. 302. Basionym: D. trifoliolatum de Wit l.c. (1953) 321, Cockburn l.c. 167,
Whitmore Tantra & Sutisna l.c. 201. Type: NIFS bb 19826, Kalimantan (holotype BO; isotypes L,
SING).
Peninsular Malaysia and Borneo. In Sabah known from Keningau district; in Sarawak
scattered. Also in Brunei and Kalimantan. In lowland forests below 450 m altitude.
4. Dialium platysepalum Baker

(Greek, platus = broad, sepalum = sepal; with broad sepals)
In Hooker f., Fl. Brit. Ind. 2 (1878) 270; Prain l.c. (1897) 172; Ridley l.c. (1922) 623; Whitmore l.c. 261;
Cockburn l.c. 165; Anderson l.c. 234; Whitmore, Tantra & Sutisna l.c. 201; Ng, MFFSS (1992) 428;
Turner l.c. 302; Rojo l.c. (1996) 612; Coode et al. (eds.) l.c. 163; Loo & Tan l.c. 78; Argent et al. (eds.)
l.c. 355. Lectotype (Rojo, 1996): Griffith 1847, Peninsular Malaysia, Malacca (holotype K). Synonyms:
D. maingayi Baker in Hooker f. l.c. 269; D. ambiguum Prain l.c. (1897) 172; D. kingii Prain l.c. (1897)
175; D. havilandii Ridl., Kew Bull. (1929) 259; D. triste de Wit l.c. (1953) 321.
Tree up to 45 m tall, 120 cm diameter. Leaves pinnate, each usually with 713 leaflets;
petiole and rachis 1524(28) cm long. Leaflets opposite, (sub)coriaceous, hairy beneath;
oblong-elliptic to oblong-lanceolate, 68(15) 24(7) cm, base rounded or cuneate,
apex abrupt to long acuminate; lateral veins usually 1012 pairs, distinct beneath; petiolules
24.5 mm long. Inflorescences terminal; rachis 1015(30) cm long; pedicel 2.54
(7) mm long. Flowers: calyx lobes 5, triangular, c. 6 4 mm; stamens 2, filaments
flattish, anthers c. 4 mm long, with both slits drawn towards the abaxial side, forming
a V-channel on that side of hairy connective; ovary 23.5 mm long; hypanthium concave.
Fruits subglobose or obovoid, sometimes slightly compressed, usually 22.5 cm long,
densely brown velvety. Seeds 1(or 2) per fruit, roundish or reniform, 0.31.3 0.9 cm.
147
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Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo. In Sabah rare; in

Sarawak throughout. Also in Brunei and Kalimantan.
Ecology. In lowland and freshwater swamp forests, up to 500 m altitude. Flowering
throughout the year; fruiting heaviest in JulyOctober.
5. Dialium procerum (Steenis) Steyaert
Fig. 6.
(Latin, procerum = very tall, high; alluding a great respect to B.H. Danser by van Steenis,
the founder of the genus Dansera)
Reinwardtia 2 (1953) 355; Whitmore l.c. 261; Anderson l.c. 234; Whitmore, Tantra & Sutisna l.c. 201;
Turner l.c. 302; Rojo l.c. 614; Coode et al. (eds.) l.c. 163. Basionym: Dansera procera Steenis, Bull.
Bot. Gard. Buitenz. 3, 17 (1948) 415. Type: NIFS. 117T. 3P. 390, Sumatra, Palembang (holotype L;
isotype BO).
Tree up to 45 m tall, 90 cm diameter. Leaves unifoliolate; petiole and petiolule 13

cm long. Leaflets coriaceous, slightly hairy beneath, glabrescent; ovate or ovate-oblong,
or elliptic, 6.516 3.57 cm, base rounded or cuneate, apex acuminate; lateral veins
68 pairs; intercostal venation reticulate, distinct. Inflorescences axillary or terminal,
sometimes fasciculate; rachis 915 cm long; pedicel 24 mm long. Flowers: calyx lobes
3, elliptic, 34 mm long, minutely hairy inside; stamens 6, anthers linear, c. 2.5 mm
long, glabrous or hairy on both sides of the connective; ovary 1.52 mm long. Fruits
c. 4(5) 2.5 cm, slightly compressed, blackish, hairs easily fallen. Seeds reniform,
c. 1.5 1 cm, brown, shining.
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sarawak known from Kapit
district; not yet found in Sabah. Also recorded for Brunei.
Ecology. In primary and old secondary forests, up to 600 m altitude. Flowering in June
March; fruiting in JanuaryApril.
7. INTSIA Thouars
(a Malagasy plant name)
merbau (Malay, preferred name)
Gen. Nov. Madag. (1806) 22; Backer & Bakhuizen f., FJ 1 (1964) 529; Whitmore, TFM 1 (1972) 262;
Cockburn, TS 1 (1976) 169; Anderson, CLTS (1980) 235; Cowan & Polhill in Polhill & Raven (eds.),
Adv. Leg. Syst. 1 (1981) 128; Corner, WSTM 3rd. edition 1 (1988) 438; Whitmore, Tantra & Sutisna,
CLK 2, 1 (1990) 202; Ding Hou, Blumea 38 (1994) 322, FM 1, 12 (1996) 622; Coode et al. (eds.),
CLBD (1996) 163; Loo & Tan, Gard. Bull. Sing. 49, 1 (1997) 80; Argent et al. (eds.), MNDT-CK 2
(1997) 357.
Trees, often with buttresses. Stipules intrapetiolar, minute, connate, caducous. Leaves
compound, once pinnate and paripinnate, usually each with 24 pairs of leaflets. Leaflets
usually with 1 or 2 small crateriform glands at the basal part on lower surface; petiolules
149
twisted. Inflorescences terminal or axillary, racemose, often fasciculate, or paniculate;

bracts caducous. Flowers bisexual, bilaterally symmetrical; hypanthium cupular, narrowly
infundibuliform or cylindrical; calyx lobes 4, smooth outside; petals: only 1 fully developed,
flabellate; stamens 3(4) fertile; filaments and staminodes connate at the base, anthers
dorsifixed; ovary puberulous, style slender, stigma small, capitellate. Fruits smooth, oblong,
rarely obcordate, straight, or falcate, flattened, glabrous, dehiscent, often 3(more)-seeded;
valves coriaceous or slightly woody, straight or slightly twisting during dehiscence. Seeds
ovoid, oblong, discoid, or sublenticular, flattened, not arillate, scurfy.
Distribution. A genus comprising 2 species or more, distributed from Madagascar, islands
of the Indian Ocean, tropical Asia, through Malesia to N Australia, Melanesia and
Micronesia. Two species occur in Sabah and Sarawak.
Key to Intsia species

Leaves each with (1)2(3) pairs of leaflets. Hypanthium 610(16) mm long. Fruits
8.515(28) 45(7.5) cm........................1. I. bijuga
Leaves each with (2)4(5) pairs of leaflets. Hypanthium 34 mm long. Fruits 1519
(40) 67(9) cm...............2. I. palembanica
1. Intsia bijuga (Colebr.) Kuntze
Fig. 7E.
(Latin, bis = twice, -jugum = paired; leaf with two pairs of leaflets)
Revis. Gen. Pl. 1 (1891) 192; Merrill l.c. (1921) 296; Masamune l.c. 346; Browne l.c. 229; Backer &
Bakhuizen f. l.c. 529; Whitmore l.c. 262; Cockburn l.c. 170; Anderson l.c. 235; Corner l.c. 438; Keng
l.c. (1990) 36; Whitmore, Tantra & Sutisna l.c. 202; Ding Hou l.c. (1994) 324; Turner l.c. 304; Ding
Hou l.c. (1996) 623; Coode et al. (eds.) l.c. 163; Loo & Tan l.c. 81; Argent et al. (eds.) l.c. 357. Basionym:
Macrolobium bijugum Colebr., Trans. Linn. Soc. 12 (1818) 359, t. 17. Type: Based on Colebrookes
desc. & pl. Synonyms: Afzelia retusa Kurz, J. As. Soc. Beng. 42, 2 (1873) 73, Ridley l.c. (1922) 639; I.
retusa (Kurz) Kuntze, Revis. Gen. Pl. 1 (1891) 192, Meijer Drees, Bull. Jard. Bot. Buitenz. 3, 16 (1938)
88.
Tree up to 40 m tall, 100 cm diameter, but usually smaller; bole straight, sometimes
crooked; buttresses when present up to 2(4) m high and 200 cm wide. Bark dippled
or later small-scaly. Leaves each with (1)2(3) pairs of leaflets; petiole and rachis (1.5
) 2.511.5 cm long, glabrous. Leaflets (broadly)elliptic, ovate, obovate, rarely suborbiculate,
(2.5)510(18) (1.5)46(12) cm, base acute, cuneate, or obtuse, often slightly
asymmetrical, apex obtuse, rounded, sometimes shortly acuminate, rarely slightly notched;
petiolules 26 mm long. Inflorescences up to 10(17) cm long, pubescent, glabrescent,
or more or less glabrous; pedicel 712 mm long. Flowers: hypanthium 610(16) mm
long; calyx lobes elliptic, 610(16) 45(7.5) mm; petals: lamina 610(30) 10
15(35) mm, claw c. 5 mm long; fertile stamens: filaments c. 30 mm long; ovary c.
7.5 mm long, style c. 40 mm long. Fruits brown or black when ripe, 8.515(28)
45(7.5) cm. Seeds 23.5 cm long and wide, c. 0.8 cm thick.
Vernacular names. Sabahepil, merbau paya (Brunei Malay), ipil (Bajan), ipil laut
(preferred name), talolonandim (Beaufort). Sarawakipil (preferred name).
150
vi
Distribution. As under the genus. Widespread in Sabah and Sarawak. Also in Brunei
and Kalimantan.
Ecology. Along (sandy) sea coasts, edges of rivers, tidal or temporarily inundated places
with brackish water, back mangroves, in primary and secondary inland forests, at altitudes
from sea level up to 600 m. Flowering and fruiting all the year round.
2. Intsia palembanica Miq.
Fig. 7AD.
(of Palembang, Sumatra)

Fl. Ned. Ind., Suppl. (1861) 289; Browne l.c. 227; Whitmore l.c. 262; Cockburn l.c. 170; Anderson l.c.
235; Corner l.c. 438; Whitmore, Tantra & Sutisna l.c. 203; Ding Hou l.c. (1994) 324, l.c. (1996) 623;
Turner l.c. 304; Coode et al. (eds.) l.c. 163; Argent et al. (eds.) l.c. 360. Lectotype (Ding Hou, 1994):
Teijsmann 4535, Sumatra, Lampong (holotype U). Synonyms: Afzelia palembanica (Miq.) Baker in
Hooker f., Fl. Brit. Ind. 2 (1878) 275; I. plurijuga Harms, Bot. Jahrb. 55 (1917) 54.
Tree up to 45 m (rarely more) tall, 100 cm (or more) diameter; buttresses up to 7 m

high, extending outwards 600 cm, 30 cm thick. Bark grey to red-brown, often rather
smooth, scaly in big trees; inner bark pale red, fibrous. Sapwood pale white to yellow,
heartwood yellow-brown to dark brown. Leaves each with (2)4(5) pairs of leaflets;
petiole and rachis (3.5)9.517.5(26) cm long, both puberulous, glabrescent. Leaflets
ovate, elliptic, sometimes broadly ovate or broadly elliptic, or suborbicular, rarely obovate
or lanceolate, (3)1013.5(18.5) (3)57.5(10.5) cm, base obtuse, rounded, cuneate,
rarely slightly truncate or subcordate, often somewhat asymmetrical, apex obtuse or
rounded, sometimes slightly notched, (shortly)acuminate; lateral veins 610 per side;
petiolules 25 mm long. Inflorescences up to 10 cm long, pubescent, glabrescent; pedicel
312 mm long. Flowers: hypanthium 34 mm long; calyx lobes (broadly)elliptic, ovate,
or slightly obovate, 68(12) 36.5(8) mm; petals: lamina 36(10) 36.5(8) mm,
claw 25 mm long; fertile stamens: filaments 2040 mm long; ovary 56 mm long, style
2035 mm long. Fruits brown or black when ripe, smooth, 1519(40) 5.57(9) cm.
Seeds 34.5 1.53 cm, 0.51.2 cm thick.
Vernacular names. Sarawakipil (Murut), lemelai (Kayan, Kenyah), merbau ayer, merbau
bukit (Iban).
Distribution. India (Andamans), Burma, Thailand, and Malesia: throughout (except Java?).
Widespread in Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. In coastal regions, edges of seasonal swamps on inundated lands, in lowland
primary or older secondary forests, sometimes on sandstone and limestone hills, at altitude
from sea level up to 900 m. Flowering in JanuaryApril and AugustNovember; fruiting
in FebruaryDecember.
Uses. Timber is rather heavy and is fairly durable. It is used for flooring and fine furnishings.
(cf.: PROSEA 5, 1 (1993) 264270; Vink in Ding Hou, Blumea 38 (1994) 322323).
152
8. KOOMPASSIA Maingay ex Benth.

(from the Malay plant name kempas)
impas, kempas (Murut), menggris (Bidayuh), tapang (Malay, Sadong)
In Hooker, Icon. Pl. 12 (1873) 58, t. 1164; Prain, J. As. Soc. Beng. 66, 2 (1897) 165; Ridley, FMP 1
(1922) 620; de Wit, Bull. Bot. Gard. Buitenz. 3, 17 (1947) 309; Whitmore, TFM 1 (1972) 264; Cockburn,
TS 1 (1976) 172; Anderson, CLTS (1980) 235; Irwin & Barneby in Polhill & Raven (eds.), Adv. Leg.
Syst. 1 (1981) 101; Corner, WSTM 3rd. edition 1 (1988) 438; Whitmore, Tantra & Sutisna, CLK 2, 1
(1990) 203; Ding Hou, FM 1, 12 (1996) 631; Coode et al. (eds.), CLBD (1996) 163; Loo & Tan, Gard.
Bull. Sing. 49, 1 (1997) 83; Argent et al. (eds.), MNDT-CK 2 (1997) 360. Synonym: Abauria Becc.,
Malesia 1 (1877) 169.
Trees. Stipules free, very small and caducous. Leaves compound, once pinnate and
imparipinnate. Leaflets 512(17) per leaf, alternate, petiolulate; midrib furrowed above,
prominent below; lateral veins usually many, very slender, often obscure and hardly
distinguishable especially on the upper surface; intercostal venation loosely finely reticulate,
often obscure or indistinct. Inflorescences axillary or terminal, paniculate; bracts and
bracteoles minute, inconspicuous, usually caducous; pedicel very short or absent. Flowers
radially symmetrical; hypanthium 0; sepals 5, puberulous to thinly pubescent; petals 5,
more or less equal in shape and size; stamens 5, free, more or less equal in length,
filaments very short, glabrous, anthers basifixed, opening with apical and basal pores,
both pores connected by a thin (nearly) dehiscent rim and sometimes appearing as
longitudinally dehiscent; ovary sessile or subsessile, 1-ovuled, style and stigma rather
short and small. Fruits compressed, indehiscent or slightly dehiscent only when germinating,
surrounded by a broad, papery wing. Seeds strongly laterally compressed, without
endosperm.
Distribution. A small genus of 3 species found in Sumatra, Peninsular Malaysia, Borneo,
the Philippines, and New Guinea. Two species in Sabah and Sarawak.
Uses. Timber eminently suitable for heavy permanent structures, flooring material, and
also good for railway sleepers, dock blocks, fence posts, shingles, etc. (cf.: Foxworthy,
Mal. For. Rec. 3 (1927), 8; K. Heyne, Nutt. Pl. Ned.-Ind. 1, 1950, 739; Burkill, EPMP
2 (1966) 1285; Cockburn l.c. 174; PROSEA 5, 1 (1993) 270275).
Key to Koompassia species

Leaflets obtuse or acute and slightly notched at the apex. Petals acute or acuminate at
the apex. Fruits glabrous............1. K. excelsa
Leaflets acuminate at the apex. Petals rounded at the apex. Fruits pubescent.
......2. K. malaccensis
1. Koompassia excelsa (Becc.) Taub.
Fig. 8AB.

In Engler & Prantl, Nat. Pflanzenfam. 3, 3 (1891 156; Merrill, Philip. J. Sci. 10 (1915) Bot. 12, l.c.
153
(1921) 299; Ridley l.c. (1922) 620; Masamune l.c. 347; de Wit l.c. (1947) 311; Browne l.c. 230; Whitmore
l.c. 264; Meijer l.c. 187; Cockburn l.c. 172; Anderson l.c. 235; K. Larsen & S.S. Larsen l.c. (1984) 84;
Corner l.c. 439; Whitmore, Tantra & Sutisna l.c. 203; Ng l.c. 429; Turner l.c. 304; Ding Hou l.c. (1996)
632; Coode et al. (eds.) l.c. 163; Argent et al. (eds.) l.c. 360. Basionym: Abauria excelsa Becc., Malesia
1 (1877) 169. Type: Beccari 1818, Sarawak, Mt. Matang near Kuching (holotype FI). Synonym: K.
parvifolia Prain l.c. (1897) 166 & 486, Ridley, Agric. Bull. Str. Fed. M.S. 1 (1902) 138 (as parviflora).
Giant tree up to 88 m tall, 100(270) cm diameter; bole straight, smooth and cylindrical;
buttresses steep, thick and up to 12 m high. Bark quite smooth, grey, often with a green
tinge; inner bark red-brown, granular outwards, more fibrous inwards. Stipules oblong,
24.5 mm long, pubescent outside. Leaves each with 712(17) leaflets; rachis slender,
67.5 cm long, pubescent, glabrescent; petiole c. 1.5 cm long. Leaflets glabrous except
sparsely hairy above, densely pubescent beneath; elliptic-oblong, sometimes elliptic,
symmetrical, 3.13.7(4.2) 11.4(1.7) cm, base subrotundate to cuneate, apex obtuse
or acute and slightly notched; petiolules short, 12.5 mm long, pubescent. Inflorescences
to 11(12.5) cm long; rachis pubescent; pedicel c. 1.5 mm long, densely puberulous.
Flowers: sepals lanceolate, equally sized, 2.53.25 mm long, thinly puberulous outside;
petals oblong or narrowly elliptic, 2.53 0.5 mm, acute or acuminate at the apex,
glabrous; stamens slightly longer than the petals, filaments stout, about as long as the
anthers, anthers oblong, 23 mm long, glabrous; ovary glabrous except with some hairs
at the basal part, style very short, c. 0.3 mm long, stigma indistinct. Fruits oblong or
elliptic-obovate, 812.5 23.5 cm (including the wing), strongly veined, glabrous. Seeds
very flat, compressed, oblong, 23 1 cm.
Vernacular names. Sabahmangaris (preferred name). Sarawaktapang (Malay, Sadong),
tahid (Kayan, Kenyah, Punan Tutoh), tanid (Murut), tanyit (Berawan).
Distribution. S Thailand (Peninsula), Sumatra, Peninsular Malaysia, Singapore, the
Philippines (Palawan), and Borneo. In Sabah common below 600 m altitude; in Sarawak
throughout. Also in Brunei and Kalimantan.
Ecology. In primary forests, in stream valleys and lower slopes of ridges, up to 400(
600) m altitude. Flowering at irregular intervals, on average once in 5 to 6 (or even
25) years, in AprilMay, JulySeptember, and December; fruiting rather rare, mainly in
JuneAugust, and October.
Uses. Timber used for indoor construction. The trees often remain in areas of secondary
growth because they are rarely felled, and often deliberately left standing to provide nesting
sites for wild honey-bees. (cf.: Burkill, EPMP, 2 (1966) 1305).
2. Koompassia malaccensis Maingay ex Benth.

(of Malacca, Peninsular Malaysia)
Fig. 8C.
In Hooker, Icon Pl. 3, 12 (1876) 58, t. 1164; Prain l.c. (1897) 166; Merrill, Philip. J. Sci. 10 (1915) Bot.
12; Ridley l.c. (1922) 620; Steenis, Bull. Jard. Bot. Buitenz. 3, 12 (1932) 170; de Wit l.c. (1947) 317;
Browne l.c. 232; Whitmore l.c. 265; Meijer l.c. 189; Cockburn l.c. 174; Anderson l.c. 235; K. Larsen &
S.S. Larsen l.c. (1984) 84; Corner l.c. 397; Keng l.c. (1990) 36; Whitmore, Tantra & Sutisna l.c. 203; Ng
154
vi
l.c. 429; Turner l.c. 304; Ding Hou l.c. (1996) 634; Coode et al. (eds.) l.c. 163; Loo & Tan l.c. 84; Argent
et al. (eds.) l.c. 363. Lectotype (de Wit, 1947): Maingay 604, Peninsular Malaysia, Malacca (holotype
K; isotype L). Synonym: K. beccariana Taub., Ber. Deutch. Bot. Ges. 10 (1892) 640, Merrill l.c. (1915)
Bot. 12, l.c. (1921) 299, l.c. (1929) 105, Masamune l.c. 347.
Giant tree up to 60 m tall, 120 cm diameter; buttresses steep, plank-like, thick and up
to 6 m high. Bark grey to brown, minutely fissured and flaking. Stipules broadly ovate,
c. 2 mm long, pubescent outside. Leaves each with 59(14) leaflets; rachis slender,
6.519 cm long, pubescent, glabrescent; petiole 1.54 cm long. Leaflets glabrous except
minutely puberulous on the midrib above, densely to thinly pubescent beneath; ovateelliptic, elliptic-oblong or lanceolate, more or less symmetrical, 5.510.4(12.5) 1.5
5 cm, base usually rounded, apex acuminate; petiolules short, 47.5(9) mm long, densely
minutely puberulous. Inflorescences up to 12 cm long; rachis pubescent, densely flowered;
pedicel 0.752.5 mm long, densely puberulous. Flowers: sepals lanceolate, 2.53 1
mm, densely puberulous outside; petals orbicular to obovate, c. 2.5 1.1 mm, rounded
at the apex, glabrous; stamens nearly half the length of the petals, filaments slender,
0.50.75 mm long, anthers heart-shaped, about as long as the filaments, glabrous; ovary
c. 1 mm long, densely dark brown puberulous, style very short and stigma indistinct.
Fruits oblong, 9.515 34.5 cm (including the wing), pubescent, veined. Seeds very
flat, compressed, irregularly oblong, up to 3.3 1 cm, purplish-black and shallowly rugose.
Vernacular names. Sabahimpas (preferred name), kempas (Malay). Sarawakapas
(Melanau), impas (Murut), kempas (Murut), kumpas (Bidayuh, Sadong), menggris (Bidayuh,
preferred name).
Distribution. Sumatra, Peninsular Malaysia (throughout), Singapore, and Borneo. In Sabah
less common than K. excelsa; in Sarawak throughout. Also in Brunei and Kalimantan.
Ecology. In lowland and hill forests, up to 800 m, often favouring an altitude not exceeding
150 m, also found in peat and freshwater swamp forests. Flowering and fruiting all the
year round.
9. LEUCOSTEGANE Prain
(Greek, leukos = white, steganos = covered; the flower covered with white calyx)
Ann. Bot. Gard. Calc. 9, 1 (1901) 37; Ridley, FMP 1 (1922) 643; Whitmore, TFM 1 (1972) 266; Anderson,
CLTS (1980) 236; Cowan & Polhill in Polhill & Raven (eds.), Adv. Leg. Syst. 1 (1981) 128; Ding Hou,
FM 1, 12 (1996) 635.
Trees. Stipules intrapetiolar, foliaceous, persistent. Leaves compound, once pinnate and
paripinnate, each with (1)25(6) pairs of leaflets. Leaflets petiolulate, stipellate.
Inflorescences on trunks or older branches, or in the axils of fallen leaves, racemose,
fascicled; bracts and bracteoles small, persistent beyond anthesis or deciduous; pedicel
articulated above the 2 bracteoles. Flowers bisexual, bilaterally symmetrical; hypanthium
infundibuliform, tubular, or cylindrical; calyx lobes 4; petals 2(or 1) developed, lateral;
stamens 2 fertile; ovary 25(or more)-ovuled. Fruits recurved, oblong, compressed, smooth,
156
vi
shortly beaked, dehiscent, 35-seeded; valves coriaceous or woody, at length twisted.

Seeds discoid or ellipsoid.
Distribution. In Malesia 2 species. Only 1 species in Sarawak, not yet recorded in Sabah.
Ecology. In lowland forest.
Leucostegane grandis Ridl.
Fig. 9.
(Latin, grandis = great; inference unclear)

Kew Bull. (1929) 257; Anderson l.c. 236; Ding Hou l.c. (1996) 636. Type: Haviland 1799, Borneo,
Sarawak, Kuching (holotype K; isotype L).
Tree up to 12 m tall, 16 cm diameter. Stipules ovate, up to 5 mm long. Leaves each

with (1)2(3) pairs of leaflets, glabrous; petiole and rachis 3.58.5 cm long. Leaflets
elliptic, oblong, 8.518(27.5) 2.58.5(9.5) cm, base obtuse, sometimes slightly unequal,
apex acuminate; lateral veins 712 per side, inconspicuous, sometimes visible beneath;
intercostal venation often obscure or invisible; petiolules c. 6 mm long; stipels very small.
Inflorescences 3.55(8) cm long; pedicel up to 20 mm long. Flowers: calyx lobes obovate
or oblanceolate, 1012 45 mm, almost glabrous except sometimes ciliate on the margins;
petals small, ovate or elliptic, 79 24.5 mm, shortly clawed at base; stamen filaments
c. 12 mm long, dilated at base; ovary pubescent. Fruits hatchet-shaped, c. 26 5 cm.
Seeds (young) elliptic, c. 1.5 0.8 cm.
Distribution. Sumatra and Borneo. In Borneo, only known from Kuching and Nanga
Pelagos, in Sarawak.
Ecology. In lowland forest, also found in swampy places. Flowering in January, April,
July, and September; fruiting in April.
10. PELTOPHORUM (Vogel) Benth., nom. cons.

(Greek, pelte = shield, pherein = to bear; bearing a shield-shaped stigma)
In Hooker, J. Bot. 2 (1840) 75; Prain J. As. Soc. Beng. 66, 2 (1897) 223; Ridley, FMP 1 (1922) 646;
Backer & Bakhuizen f., FJ 1 (1964) 547; Whitmore, TFM 1 (1972) 267; Cockburn, TS 1 (1976) 175;
Anderson, CLTS (1980) 237; Polhill & J.E. Vidal in Polhill & Raven (eds.), Adv. Leg. Syst. (1981) 90;
Corner, WSTM 3rd. edition 1 (1988) 440; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 204; Ding
Hou, FM 1, 12 (1996) 650; Coode et al. (eds.), CLBD (1996) 163; Loo & Tan, Gard. Bull. Sing. 49, 1
(1997) 86. Basionym: Caesalpinia sect. Peltophorum Vogel, Linnaea 11 (1837) 406. Synonym:
Baryxylum Lour., Pl. Cochinch. (1790) 266, nom. rejic.
Trees, unarmed. Stipules lobed or branched, often persistent, or small and caducous.
Leaves compound, bipinnate, deciduous. Leaflets small, numerous (427 pairs per pinna).
Inflorescences terminal and/or axillary, racemose or paniculate; bracts present, sometimes
caducous; bracteoles lacking. Flowers bisexual; hypanthium very short or obscure; calyx
lobes 5; petals 5, 1225 mm long; stamens 10, free, filaments thickened and pilose at
158
the basal part; ovary 38-ovuled, style filiform, incurved, stigma prominent, peltate or
capitate. Fruits oblong, hard, compressed, indehiscent, 16-seeded; with a firm, winglike extension on each suture, usually longitudinally striate. Seeds lenticular, compressed,
without endosperm.
Distribution. A pantropical genus of about 15 species with 3 of them occurring in Sabah
and Sarawak.
Key to Peltophorum species

1. Inflorescences or infructescences terminal and axillary, paniculate. Pedicel 57 mm
long............2. P. pterocarpum
Inflorescences or infructescences axillary, racemose. Pedicel 1040 mm long..........2
2. Leaves with 35 pairs of pinnae; stipules lanceolate to linear, caducous. Pedicel 10
12 mm long. Seeds longitudinally positioned in the pod..................3. P. racemosum
Leaves with 59 pairs of pinnae; stipules lobed and usually relobed, often persistent.
Pedicel 2040 mm long. Seeds more or less transversely positioned in the pod.........
.....1. P. dasyrhachis
1. Peltophorum dasyrhachis (Miq.) Kurz
Fig. 10AC
(Greek, dasus = densely hairy, rachis = axis; axis of a compound leaf or an inflorescence
densely hairy)
J. As. Soc. Beng. 45, 2 (1876) 128, l.c. (1877) 293; Prain l.c. (1897) 224; Ridley l.c. (1922) 647; Backer
& Bakhuizen f. l.c. 547; Whitmore l.c. 267; K. Larsen & S.S. Larsen l.c. (1980) 60, l.c. (1984) 54;
Corner l.c. 440; Ng l.c. 430; Turner l.c. 307; Ding Hou l.c. (1996) 650. Basionym: Caesalpinia
dasyrhachis Miq., Fl. Ned. Ind., Suppl. (1861) 292. Type: Teijsmann s.n., Sumatra, Lampong (holotype
L; isotype K).
Tree up to 30 m tall; young branchlets brownish red tomentose, glabrescent. Stipules

lobed and usually relobed, often persistent. Leaves: petiole up to 5.5(7) cm and rachis
up to 32 cm long, both tomentose; pinnae 59 pairs. Leaflets 616 pairs per pinna,
sessile, finely pubescent on both surfaces, glabrescent, sometimes almost glabrous above;
oblong or elliptic-oblong, 12.5 0.41 cm, base acute, obtuse or rounded, apex obtuse
or slightly emarginate. Inflorescences axillary, racemose, 1530 cm long; rachis pubescent;
bracts linear, 1012 mm long, more or less persistent; pedicel 2040 mm long. Flowers:
calyx lobes 1015 56 mm, tomentose outside; petals obovate, 1525 1012 mm,
hairy towards the base of the inner side; stamens with filaments 1015 mm long; ovary
velutinous, 48-ovuled, style c. 12 mm long. Fruits reddish brown, elliptic, 1015
24 cm (including wing-like extension of 45 mm wide), tapering towards both ends,
brown-pubescent, glabrescent, not or indistinctly striate. Seeds 48, flat, 0.11.2 0.5
cm, more or less transversely positioned in the pod.
Distribution. Thailand, Cambodia, Laos, Vietnam, Sumatra, Peninsular Malaysia, and
Borneo. In Sabah known from Sandakan and Keningau districts; in Sarawak recorded
for Kuching district. Also in Kalimantan.
159
Ecology. In lowland forest. Flowering in FebruaryJuly and NovemberDecember; fruiting

in January, March, MayJune, and December.
2. Peltophorum pterocarpum (DC.) Backer ex K. Heyne

(Greek, pteron = wing, karpos = fruit; the winged fruit)
Nutt. Pl. Ned.-Ind., ed. 2, 2 (1927) 755; Backer & Bakhuizen f. l.c. 547; Burkill l.c. 1715; Whitmore l.c.
268; Hattink, Reinwardtia 9 (1974) 59; Cockburn l.c. 175; Verdcourt l.c. 16; Anderson l.c. 237; K.
Larsen & S.S. Larsen l.c. (1980) 62, l.c. (1984) 54; Corner l.c. 440; Keng l.c. (1990) 37; Whitmore,
Tantra & Sutisna l.c. 205; Rudd l.c. 57; Ng l.c. 430; Turner l.c. 308; Ding Hou l.c. (1996) 651; Coode et
al. (eds.) l.c. 163; Loo & Tan l.c. 86. Basionym: Inga pterocarpa DC., Prodr. 2 (1825) 441. Type: S.
coll., s.n.. in ins. Timor. Fl. ign. (v.s. comm. Mus. Par.) (holotype P; isotype K). Synonyms:
Caesalpinia inermis Roxb., Fl. Ind. ed. Carey 2 (1832) 367; P. inerme (Roxb.) Llanos in Blanco, Fl.
Filip. Ed. 3 (187783) t. 335, Merrill l.c. (1921) 301, Masamune l.c. 350.
Tree up to 35 m tall, 100 cm diameter. Young shoots pubescent or tomentose, glabrescent.

Stipules small, linear, not lobed, caducous. Leaves: petiole up to 10 cm and rachis up
to 26 cm long; pinnae 414 pairs. Leaflets (8)1015(22) pairs per pinna, sessile, nearly
glabrous above, finely pubescent, glabrescent beneath; oblong, 12 0.31 cm, base
unequally acute or attenuate, or rounded, apex obtuse, rounded or slightly emarginate.
Inflorescences terminal and axillary, paniculate, up to 40 cm long, densely pubescent;
bracts minute, deltoid, 58 mm long, caducous; pedicel usually 57 mm long. Flowers:
calyx lobes slightly triangular, 58 56 mm, pubescent outside; petals obovate, 17
25 1013 mm, densely hairy towards the basal part; stamen filaments 1215 mm long;
ovary hairy, 34-ovuled. Fruits oblong-elliptic, up to 14 2.5 cm (including wing-like
extension of 45 mm wide), pubescent, glabrescent or almost glabrous, 14-seeded. Seeds
oblong, flat, 11.2 0.5 cm, light brown, longitudinally positioned in the pod.
Vernacular name. Sabah and Sarawakkupang (Malay).
Distribution. Sri Lanka, Thailand, Cambodia, Vietnam, throughout Malesia to northern
Australia. In Sabah recorded from Kota Kinabalu (Pulau Gaya), Kudat, Sandakan, and
Tawau districts; in Sarawak uncommon but widespread. Also in Brunei.
Ecology. Frequently growing along beaches and behind the mangroves, often in lowland,
open forests, occasionally found on limestone plateau. Flowering and fruiting all the year
round.
Uses. Widely introduced and cultivated in the tropics. Planted as shade trees in plantations
and as an very attractive ornamental in parks and along roadsides. Bark is used for dyes
and for various traditional medicinal purposes. (cf.: K. Heyne, Nutt. Pl. Ned.-Ind. 1 (1950)
755; Burkill, EPMP 2 (1966) 1715).
3. Peltophorum racemosum Merr.
Fig. 10DE.
(Latin, racemus = raceme; the inflorescence)

Philip. J. Sci. 11 (1916) Bot. 84, l.c. (1921) 301; Masamune l.c. 350; Meijer l.c. 191; Cockburn l.c. 175;
160
vi
Whitmore, Tantra & Sutisna l.c. 205; Ding Hou l.c. (1996) 653. Type: Hubert Winkler 2424, Borneo,
Kalimantan (isotype L).
Tree up to 60 m tall, 80 cm diameter; buttresses flat, c. 1 m high, 100 cm extending

outward, 4 cm thick. Young shoots prominently ferruginous-pubescent. Stipules lanceolate
to linear, c. 5 mm long, caducous. Leaves: petiole up to 10 cm and rachis up to 20
cm long; pinnae 35 pairs, up to 10 cm long. Leaflets 411 pairs per pinna, sessile
or subsessile, upper surface glabrous except sometimes slightly hairy on the midrib, lower
surface covered with short, appressed hairs; oblong, oblong-elliptic, 2030(60) 8
12(25) mm, base subacute or obtuse, apex broadly rounded and minutely apiculate; lateral
veins up to 14 per side, visible or distinct on both surfaces. Inflorescences axillary, solitary,
racemose, up to 27 cm long; rachis pubescent; bracts triangular, c. 3 mm long, caducous;
pedicel 1012 mm long. Flowers: hypanthium shortly discoid, c. 5 mm diameter; calyx
densely pubescent outside, lobes oblong, 710 45 mm, pubescent at the lower half
inside; petals suborbicular, subreniform, or broadly obovate, 1216 mm long and wide,
pubescent at the lower half outside and at the basal part inside; stamen filaments 7
10 mm long; ovary elliptic or lanceolate, 46 mm long, densely pubescent, 4-ovuled.
Fruits elliptic, narrow-elliptic, or obovate-elliptic, flat, 1215.5(19) 34 cm (including
marginal extension of c. 5 mm wide), loosely pubescent, glabrescent, or almost glabrous.
Seeds elliptic, flat, c. 2 1 cm, longitudinally positioned in the pod.
Vernacular name. Sabahtimbarayong (preferred name).
Distribution. Endemic to Borneo. In Sabah and Sarawak scattered and widespread. Also
in Kalimantan.
Ecology. In lowland primary forest. Flowering in April and June; fruiting in February
October.
11. SARACA L.
(possibly relates to the Javanese plant name soka,
derived from the Indian Sanskrit name asoka)
babai (Iban), biyau (Bidayuh), gaparang (Murut), gapis (Malay), kelangan (Kenyah)
Mant. Pl. 1 (1767) 98; Prain, J. As. Soc. Beng. 66, 2 (1897) 210; Ridley, FMP 1 (1922) 640; Backer &
Bakhuizen f., FJ 1 (1964) 527; Zuijderhoudt, Blumea 15 (1968) 414; Whitmore, TFM 1 (1972) 268;
Cockburn, TS 1 (1976) 175; Anderson, CLTS (1980) 238; Cowan & Polhill in Polhill & Raven (eds.),
Adv. Leg. Syst. 1 (1981) 128; Corner, WSTM 3rd. edition 1 (1988) 441; Ding Hou, FM 1, 12 (1996)
660; Coode et al. (eds.), CLBD (1996) 163; Argent et al. (eds.), MNDT-CK 2 (1997) 366. Synonym:
Jonesia Roxb., Asiat. Res. 4 (1799) 355, cum pl., Wight, Icon. Pl. Ind. Orient. 1 (1839) t. 206.
Shrubs or trees. Stipules axillary, enveloping the bud, minute and usually caducous. Leaves
spiral, compound, once pinnate and paripinnate, each with 17 pairs of leaflets, subsessile
to shortly petioled. Leaflets sometimes with a pair of wart-like glands at the base and/
or at the apex; lateral veins more or less straight or slightly curved and obliquely upward
162
towards the margins, usually elevated and distinct beneath; petiolules very short.
Inflorescences terminal or axillary, or on older branches, often rather dense, corymbose
or paniculate, usually many-flowered; bracts caducous or persistent; bracteoles (sub)opposite,
often coloured and showy; pedicel very short. Flowers bisexual, or functionally male
(with a short, small, sterile or rudimentary pistil); hypanthium cylindrical or tubular;
calyx lobes 4(6), subequal, petaloid; petals 0; stamens usually 4 fertile, in one whorl,
filaments exserted, anthers often shortly hairy at apex and/or base; ovary oblong or
lanceolate, usually 28-ovuled; style filiform, stigma terminal, minute, obtuse. Fruits
smooth outside, suborbicular, broadly ovoid, ellipsoid, or obovoid, flattened, straight or
distinctly curved, coriaceous or woody, dehiscent, 18-seeded; valves twisting and enrolling
during dehiscence. Seeds ovoid, ellipsoid, or subglobose, smooth, compressed or subterete,
without aril.
Distribution. A genus of about 11 closely allied species distributed in India, Sri Lanka,
SE Asia, and Malesia. Two species in Sabah and Sarawak.
Ecology. In lowland forests especially growing along streams.
Uses. Trees of some species are cultivated in gardens for ornamental purpose. The trees
are rather small, so the wood is used for making small utensils.
Key to Saraca species

Hypanthium 715(32) mm long, 0.52 times longer than the calyx lobes. Ovary with
more ovules (often 68). Fruits oblong or lanceolate, usually 731 37 cm, 68seeded. ....1. S. declinata
Hypanthium (3)5.57 mm long, about the same length as the calyx lobes. Ovary with
only 2 ovules. Fruits suborbicular, broadly ovoid or obovoid, or oblong, 48 34 cm,
12-seeded......2. S. hullettii
1. Saraca declinata (Jack) Miq.
Fig. 11AE.
(Latin, declinatus = bent or curved downwards, or drooping; the branches)

Fl. Ned. Ind. 1, l (1855) 84 & 1080; Prain l.c. (1897) 211; Ridley l.c. (1922) 641; Merrill, J. Arn. Arbor.
33 (1952) 228; Zuijderhoudt l.c. 419; Whitmore l.c. 269; Cockburn l.c. 178; Verdcourt l.c. 89; Anderson
l.c. 238; J.E. Vidal, Fl. Cambodia, Laos & Vietnam 18 (1980) 133; Fl. Thail. 4 (1984) 95; Corner l.c.
444; Turner l.c. 309; Ding Hou l.c. (1996) 665; Coode et al. (eds.) l.c. 163.; Argent et al. (eds.) l.c. 369.
Basionym: Jonesia declinata Jack, Mal. Misc. 2(7) (1822) 74; reimpr. in Hooker, Comp. Bot. Mag. 1
(1836) 223. Type: Jack s.n., Sumatra (isotype G?). Synonyms: S. macroptera Miq., Fl. Ned. Ind. 1, 1
(1855) 1080, Merrill l.c. (1921) 296, Ridley l.c. (1922) 642; J. palembanica Miq., Fl. Ned. Ind., Suppl.
(1861) 291; S. palembanica (Miq.) Baker in Hooker f., Fl. Brit. Ind. 2 (1878) 272, Ridley l.c. (1922)
642; S. triandra auct. non (Roxb.) Baker, Ridley l.c. (1922) 642; S. lanceolata Merr. l.c. (1929) 100; S.
crassifolia Ridl., Kew Bull. (1933) 491; S. elmeri Ridl., Kew Bull. (1938) 492; S. elegans Ridl. l.c.
(1938) 278; S. longistyla Ridl. l.c. (1938) 279.
Tree up to 30 m tall, 60 cm diameter, sometimes treelet up to 6 m tall; buttresses very

small. Bark smooth, grey, greenish black, lenticellate, longitudinally fissured, flaking in
163
vi
small pieces; inner bark brown, reddish brown, firmly fibrous. Sapwood white, soft. Leaves
each with (1)35(7) pairs of leaflets; petiole and rachis (4.5)8.556 cm long. Leaflets
glabrous; elliptic-lanceolate, lanceolate, or oblanceolate, 430(50) 1.511(20) cm,
base cuneate, obtuse, or rounded, rarely subcordate for the lowest pair, symmetrical, apex
acuminate, acute, or obtuse; lateral veins 612 per side; petiolules often very short,
sometimes up to 10 mm long. Inflorescences slightly pubescent or subglabrous, main
branches (1)710(30) cm long; pedicel 1015(35) mm long. Flowers puberulous
outside; hypanthium 715( 32) mm long; calyx lobes ovate, elliptic, or obovate, 615
37 mm; stamens (3)4(5), filaments 1638 mm long, glabrous; ovary puberulous,
68(10)-ovuled, style 720 mm long. Fruits oblong or lanceolate, 731 37 1.2
cm, woody, 68-seeded; valves coiling during dehiscence. Seeds ovoid-oblong, c. 4
2 cm.
Vernacular names. Sabahbakis, bokis, balanut, dulanut, jambakia (Dusun); gapis
(preferred name), kachang kachang, langsat monyet (Malay), marang parang, pasoi,
selungapid, tagandop, tambakis (Dusun); tululs hobui (Kadazan). Sarawakbabai (Iban).
Distribution. India, Burma, Indo-China, Thailand, and Malesia. Widespread in Sabah and
Sarawak. Also in Brunei and Kalimantan.
Ecology. In primary mixed dipterocarp forest, along rivers and streams, often drooping
over the water, sometimes in swamp forest or secondary vegetation, on hill slopes, basalt
hillside, clay or clay loam soils, rich soils, and occasionally on slopes of limestone hills,
up to 800 m altitude. Flowering and fruiting all the year round.
2. Saraca hullettii Prain
Fig. 11F.
(R. Hullett, 18431914, England, a schoolmaster in Singapore, a member of the Gardens

Committee)
J. As. Soc. Beng. 65, 2 (1897) 492; Merrill l.c. (1921) 296; Ridley l.c. (1938) 278; Masamune l.c. 352;
Zuijderhoudt l.c. 418; Anderson l.c. 238; Ding Hou l.c. (1996) 667. Syntypes: Hullett 312 and Haviland
s.n. (CAL), and Beccari PB 916 (K), all from Sarawak.
Tree up to 20 m tall, 30 cm diameter. Bark smooth, greyish, light or pink-brown. Sapwood

white. Leaves each with 2 or 3 pairs of leaflets; petiole and rachis 316 cm long. Leaflets
glabrous; lanceolate, 5.535 211 cm, base cuneate to attenuate, sometimes obtuse,
symmetrical, apex acuminate rarely caudate; lateral veins 911 per side; petiolules 2
10 mm long. Inflorescences puberulent, main branches 714 cm long; pedicel 2230
mm long. Flowers glabrous outside; hypanthium (3)5.57 mm long, hairy at the upper
half inside; calyx lobes ovate, elliptic, oblong, or obovate, (3)510 (2)45.5 mm;
stamens 4, rarely 3, filaments 1525 mm long, glabrous; ovary 2-ovuled, style 812 mm
long. Fruits suborbicular, broadly ovoid or obovoid, or oblong, 48 34 cm, thinly
woody, 12-seeded. Seeds (young) ovoid, c. 1.5 l cm.
Vernacular name. Sarawakbi-n (Iban).
Distribution. Endemic to Borneo. In Sabah recorded for Tawau district; in Sarawak known
from Bintulu, Kuching, Marudi, Miri, and Serian districts. Also in Brunei and Kalimantan.
165
Ecology. In lowland and riverine forests, often immediately beyond tidal limits, on clayey
soils, sandstone or limestone, once found in peatswamp forest, up to 300 m altitude.
Flowering in January, MarchApril, and JulyNovember; fruiting in July and October.
12. SINDORA Miq.

(Latinised local tree name sindur)
borak (Bidayuh, Padawan), petir (Iban), sepetir (Malay)
Fl. Ned. Ind. 1 (1860) 287; de Wit, Bull. Bot. Gard. Buitenz. 3, 18 (1949) 5; Backer & Bakhuizen f., FJ
1 (1964) 526; Whitmore, TFM 1 (1972) 270; Cockburn, TS 1 (1976) 178; Anderson, CLTS (1980) 238;
Cowan & Polhill in Polhill & Raven (eds.), Adv. Leg. Syst. 1 (1981) 132; Corner, WSTM 3rd. edition 1
(1988) 445; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 207; Ding Hou, FM 1, 12 (1996) 691; Coode
et al. (eds.), CLBD (1996) 164; Loo & Tan, Gard. Bull. Sing. 49, 1 (1997) 98; Argent et al. (eds.),
MNDT-CK 2 (1997) 369. Synonym: Echinocalyx Benth. in Bentham & Hooker f., Gen. Pl. 1 (1865)
584 & (1867) 1003.
Trees. Stipules foliaceous, free, rarely connate at the lower half. Leaves compound, once
pinnate and paripinnate, each with 210 pairs of leaflets, petioled. Leaflets: midrib slightly
grooved above, prominent below; lateral veins many, usually more than 20 per side, obscure
or faint above, visible or distinct beneath, obliquely ascending towards and close to the
margins and forming a thickened vein; intercostal venation fine, very closely reticulate
on both surfaces; petiolules short. Inflorescences solitary or fascicled, paniculate or
racemose; bracts and bracteoles small, often caducous or sometimes present at anthesis.
Flowers bisexual, bilaterally symmetrical, shortly pedicelled or subsessile; hypanthium
very short; calyx lobes 4, narrowly imbricate, outer surface often with spiny outgrowths;
petals 1, usually fleshy, rudimentary ones often wanting; stamens 10 (9 + 1), uppermost
one free and reduced to a short staminode, remaining 9 shortly and obliquely united
at base and sheath-like, the united part often hairy on both surfaces, 2 upper ones fertile
with elongate filaments and ovoid or oblong, longitudinally dehiscing, dorsifixed anthers,
7 lower ones with shorter filaments and with or without small, imperfect anthers; ovary
25(or more)-ovuled, style filiform, often recurved, stigma small. Fruits flat, rounded
ovate or elliptic, often spiny, rarely unarmed, dehiscent, woody, 2-valved. Seeds few, black,
shining, with a large fleshy aril at the base.
Distribution. A genus comprising 1820 species, distributed in W Africa (Gabon, 1 sp.),
SE Asia, Peninsular Malaysia, Singapore, Indonesia, Borneo, and the Philippines. Eight
Ecology. In lowland forests, favouring well-drained areas, sometimes occurring near the
banks of brooks or small rivers, often scattered, growing singly or in small groups, up
to 150 m, sometimes to 350 m, very rarely at higher altitudes.
Uses. Plants of some species produce strong and durable timber of excellent quality, resin
or wood-oil, medicine, tannin, etc. (cf.: K. Heyne, Nutt. Pl. Ned.-Ind. 1 (1950) 727
730; Burkill, EPMP 2 (1966) 20672070; PROSEA 5, 1 (1993) 434442).
166
Key to Sindora species

(for flowering material)
1.
Leaflets usually glabrous on both surfaces, except sometimes sparsely hairy on the
midrib beneath......................2
Leaflets usually only minutely puberulous, pubescent, or hirsute on the lower surface,
rarely sparsely puberulous above......................4
2.
Apex of leaflets usually obtuse, acute, sometimes slightly emarginate. Calyx lobes
4.55(6) mm long.............6. S. leiocarpa
Apex of leaflets usually shortly acuminate to acuminate. Calyx lobes 6.510 mm
long..................3
3.
Apex of leaflets with a gland at the tip on the lower surface. Calyx lobes without
spiny outgrowths on the outer surface...................3. S. coriacea
Apex of leaflets without a gland as above. Calyx lobes with spiny outgrowths on
the outer surface....................1. S. affinis
4.
Leaves usually with 3 or 4 pairs of leaflets.....................5

Leaves usually with 5 or 6 pairs of leaflets.................7
5.
Calyx lobes without spiny outgrowths on the outer surface...........2. S. beccariana

Calyx lobes bearing spiny outgrowths on the outer surface....................6
6.
Apex of leaflets obtuse. Calyx lobes broadly ovate or lanceolate, 45.5 1.52.5
mm.....4. S. echinocalyx
Apex of leaflets usually acute or subacuminate. Calyx lobes lanceolate, up to 9.5
3 mm............8. S. wallichii
7.
Under surface of leaflet usually densely hirsute, sometimes slightly glabrescent.....

.......7. S. velutina
Under surface of leaflet sparsely and minutely puberulous, glabrescent, sometimes
almost glabrous except on the midrib...................5. S. irpicina
Key to Sindora species

(for fruiting material)
1. Fruits armed or bearing spines................2
Fruits unarmed or smooth without spines.................7
2. Leaflets usually glabrous on both surfaces..........1. S. affinis
Leaflets puberulous, pubescent, or hirsute on the lower surface..............3
3. Leaves usually with 3 or 4 pairs of leaflets...............4
Leaves usually with 5 or 6 pairs of leaflets...........6
4. Leaflets obtuse, slightly notched at the apex...........4. S. echinocalyx
Leaflets usually acute, subacuminate to attenuate.............5
167
5. Fruits broadly ellipsoid or suborbicular, 78 cm wide, covered with scattered spine..

....2. S. beccariana
Fruits suborbicular, broadly obovoid or ellipsoid, 46 cm wide, covered with dense
spines......8. S. wallichii
.
6. Leaflets densely hirsute on the lower surface. Fruits covered with small, scattered
spines and with fine warts among them...............7. S. velutina
Leaflets often minutely puberulous on the lower surface. Fruits covered with slender,
numerous spines and without fine warts among them..........5. S. irpicina
7. Leaflets shortly to distinctly acuminate, apex with a gland at the tip on the lower
surface...............3. S. coriacea
Leaflets usually obtuse, acute, or shortly acuminate, sometimes slightly emarginate,
apex without a gland as above...............6. S. leiocarpa
1. Sindora affinis de Wit

(Latin, affinis = allied to; inference unknown)
694. Type: bb 19774, Kalimantan, Tidung lands near Nunukan (holotype BO; isotype L).
Tree up to 35 m tall, 60 cm diameter. Bark rather smooth, hardly cracked, light brown;
inner bark pale brown to purplish black. Sapwood white, heartwood light brown, streaked
with darker bands. Leaves each with (3)45(6) pairs of leaflets; petiole 1.53(5) cm
and rachis 414 cm long, slightly puberulous, glabrescent, or almost glabrous. Leaflets
often almost glabrous on both surfaces, except sparsely hairy on the midrib of lower
surface; oblong or oblong-elliptic, elliptic, or ovate 2.58(9.5) 24(5) cm, base
rotundate, often symmetrical, apex shortly acuminate or acuminate, without a gland at
the tip on the lower surface. Inflorescences paniculate, 920 cm long; both rachis and
branches densely tawny puberulous; bracts and bracteoles lanceolate, 34 mm long, densely
puberulous; pedicel 13(5) mm long, densely puberulous. Flowers: calyx lobes with
spiny outgrowths on the outer surface, elliptic or lanceolate, 710 35 mm, densely
puberulous outside and appressed hairy inside; petals elliptic, 8 23 mm, outside densely
puberulous and inside glabrous; stamens: free filaments and staminodes to c. 12 mm
long, sparsely hairy often at the lower part or at base, or glabrous; ovary c. 2 mm long,
densely woolly, 56-ovuled, style up to 15 mm long. Fruits (young) ellipsoid, slightly
asymmetrical, 58.5 34.5 cm, flattened, slightly minutely hairy, not veined, with
scattered, very short spines; beak very short. Seeds only very young ones seen.
Vernacular name. Sabahsepitir (Tawau).
Distribution. Endemic to Borneo. In Sabah found in Tawau district; not reported for
Ecology. In lowland forest, growing in well-drained areas in sandy or clayey soils, once
found in loam soils on sandstone, up to 60 m altitude. Flowering in April and July;
fruiting in May.
168
2. Sindora beccariana Backer ex de Wit

Bull. Bot. Gard. Buitenz. 3, 18 (1949) 18; Cockburn l.c. 181; Anderson l.c. 239; Whitmore, Tantra &
Sutisna l.c. 207; Ding Hou l.c. (1996) 694; Coode et al. (eds.) l.c. 164; Argent et al. (eds.) l.c. 370.
Type: Beccari 529, Borneo (no precise locality) (holotype BO; cf. de Wit, 1949, f. 2g, based on the
type).
Tree up to 40 m tall, 80 cm diameter. Bark smooth, greenish or dark grey, brown-mottled;

inner bark whitish or reddish. Sapwood whitish to brownish, heartwood streaky. Leaves
each with 34 pairs of leaflets; petiole 2.53.5 cm and rachis 814 cm long, slightly
puberulous, glabrescent, or almost glabrous. Leaflets upper surface often almost glabrous,
lower surface densely minutely rusty puberulous; elliptic, ovate-elliptic, or ovate, 3.5
8.5 1.74 cm, base rotundate, obtuse, or slightly cuneate, often symmetrical, apex
acuminate to attenuate. Inflorescences paniculate, up to 16 cm long; both rachis and
branches densely puberulous; bracts and bracteoles ovate to lanceolate, 12.5 mm long;
pedicel 11.5 mm long. Flowers: calyx lobes narrowly imbricate, lanceolate or elliptic,
5.57 22.5 mm, unarmed, densely puberulous on both surfaces; petals elliptic or slightly
obovate-oblong, 5 2.53.5 mm, outside densely puberulous in the lower half, inside
glabrous except with appressed hairs on the vertical, central part; stamens: free filaments
and staminodes up to 10 mm long, hairy often on the lower half; ovary densely woolly,
23-ovuled, style up to 12 mm long. Fruits purple, broadly ellipsoid or suborbicular,
79(11) 78 cm, flattened, hard, evenly covered with scattered, sharp spines (c. 5
mm long). Seeds broadly ovate or suborbicular, c. 3.5 3 cm.
Vernacular names. Sabahsampa rantu, sinampar, sindur (Malay), sepetir (preferred
name).
Distribution. Endemic to Borneo. In Sabah known from Beaufort, Labuk Sugut, Lahad
Datu, Sandakan, Sipitang, Tenom, and Tawau districts; in Sarawak recorded for the 1st
and 4th Divisions, more common near the coast. Also in Brunei and Kalimantan.
Ecology. In lowland forests in flat or sloping country, sometimes near the banks of brooks,
scattered, rare or sometimes locally common, growing on well-drained often sandy loam
or clay soils, often below 100 m, once recorded from c. 850 m altitude (Sabah: Weston,
Beaufort district). Flowering in JuneJuly and November; fruiting in AprilJune and
SeptemberDecember.
Use. The timber is cut and exported as a fine wood for joinery (Cockburn, 1976).
3. Sindora coriacea (Baker) Prain

J. As. Soc. Beng. 66, 2 (1897) 206 & 482; Ridley l.c. (1922) 639; Merrill l.c. (1929) 99; Masamune l.c.
353; de Wit l.c. (1949) 30; Whitmore l.c. 271; Cockburn l.c. 182; Anderson l.c. 239; K. Larsen & S.S.
Larsen l.c. (1984) 98; Whitmore, Tantra & Sutisna l.c. 207; Turner l.c. 310; Ding Hou l.c. (1996) 697;
Loo & Tan l.c. 99. Basionym: Afzelia ?coriacea Baker in Hooker f., Fl. Brit. Ind. 2 (1878) 275. Type:
Maingay 566, Peninsular Malaysia, Malacca (holotype K).
169
Tree up to 33 m tall, 95 cm diameter; buttresses up to 0.6 m high and spread. Bark

dark brown, smooth, surface faintly hooped or cracked; inner bark fawn or brown, granular.
Sapwood white or pale yellow. Stipules caducous. Leaves each with 3 or 4 pairs of
leaflets; petiole (1.5)2.54 cm and rachis 5.510(14) cm long, glabrous. Leaflets often
almost glabrous on both surfaces; elliptic, ovate, or slightly obovate, (3.5)510(15)
(2)35(7.5) cm, base acute, cuneate, obtuse, or rounded, sometimes unequally sided,
apex shortly acuminate or acuminate, rarely rounded or retuse, with a gland at the tip
on the undersurface. Inflorescences paniculate, up to 20(30) cm long; both rachis and
branches densely puberulous; bracts and bracteoles ovate or lanceolate, 1.53 mm long,
densely puberulous; pedicel 12.5 mm long, densely puberulous. Flowers: calyx lobes
elliptic or lanceolate, 6.57.5 2.53 mm, outside densely puberulous and without spiny
outgrowths, densely appressed hairy insides; petals slightly obovate-oblong, 6.57.5
33.5 mm, outside densely puberulous, inside glabrous except the villose margin; stamens:
free filaments and staminodes up to 12 mm long, sparsely hairy often at the lower part
or at base, or glabrous; ovary c. 4 mm long, densely woolly along the suture, 45ovuled, style up to 10 mm long. Fruits (young) ellipsoid or slightly broadly ellipsoid,
710 46 cm, flattened, often unequally sided, slightly minutely hairy, faintly veined,
unarmed. Seeds 2 or 3, arillate.
Vernacular names. Sabahsepetir or sapetir (Malay, commonly used).
Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, Singapore, and Borneo.
In Sabah scattered, widespread; not reported in Sarawak. Also in Brunei and Kalimantan.
Ecology. In forests at low altitudes, favouring well-drained sandy or leached soils. Flowering
in MarchSeptember; fruiting throughout the year.
4. Sindora echinocalyx (Benth.) Prain

(Greek, echinus = hedge-hog, sea-urchin, calyx = calyx; calyx with some spiny outgrowths)
J. As. Soc. Beng. 66, 2 (1897) 204 & 481 (quoad Griffith K distr. no. 1848, excl. syn.); Symington, Kew
Bull. (1938) 75 & 77; de Wit l.c. (1949) 34; Whitmore l.c. 272; K. Larsen & S.S. Larsen l.c. (1984) 99;
Turner l.c. 310; Ding Hou l.c. (1996) 698. Basionym: Echinocalyx Benth. in Bentham & Hooker f.,
Gen. Pl. 1 (1865) 584 & 1003, pro gen. Lectotype (Symington, 1938): Griffith Kew Distr. no. 1848,
Peninsular Malaysia, Malacca (holotype K; isotype L ). Synonym: S. wallichii auct. non Benth. in
Benth. & Hook. (quoad Griffith K distr. 1848): Benth. in Hooker, Ic. Pl. (1867) t. 1017, Ridley l.c.
(1922) 637, p.p.
Tree up to 40 m tall, 70 cm diameter; buttresses absent. Bark smooth, brown with grey
patches, rough. Stipules foliaceous, up to 8 mm long, early caducous. Leaves each with
3 or 4 pairs of leaflets; petiole 22.5 cm and rachis c. 7 cm long, minutely puberulous,
glabrescent. Leaflets upper surface glabrous, lower surface puberulous especially on the
midrib; elliptic or elliptic-oblong, rarely obovate, 47.5 23 cm, lower ones usually
smaller than the upper ones, base obtuse or rotund, often symmetrical, apex obtuse, the
very tip slightly notched. Inflorescences paniculate, up to 15 cm long; both rachis and
branches densely puberulous; bracts and bracteoles triangular, ovate or lanceolate, 47
mm long; pedicel very short. Flowers: calyx lobes broadly ovate or lanceolate, 45.5
1.52.5 mm, puberulous and bearing some spiny outgrowths outside, puberulous insides;
170
petals slightly obovate or elliptic, c. 5 2.5 mm, puberulous outside and glabrous inside;
stamens: free filaments up to 10 mm long, slightly hairy at the lower part; ovary c.
5 mm long, densely hairy, with numerous minute, spiny outgrowths, 24-ovuled, style
up to 10 mm long. Fruits ovoid or broadly ellipsoid, or suborbiculate, 46.5 3.5
6 cm, flattened, covered with close, hard, straight spines (up to 4 mm long), 12(3)seeded. Seeds arillate.
Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, and Borneo. In Borneo
known from Sandakan district, in Sabah.
Ecology. In forests on hillsides and ridges, from lowland up to 600 m altitude.
5. Sindora irpicina de Wit

(Latin, irpicinus = like a rake or harrow; the pod bearing evenly distributed and equally sized
prickles)
Bull. Bot. Gard. Buitenz. 3, 18 (1949) 50; Cockburn l.c. 182; Whitmore, Tantra & Sutisna l.c. 207; Ding
Hou l.c. (1996) 700; Argent et al. (eds.) l.c. 374. Type: Nieuwenhuis & Jaheri 908 (24-4-1896), cult. in
Hort. Bog. sub no. I-E-4a, coll. 13-5-1941, Borneo, Kalimantan (holotype BO).
Tree up to 40 m tall, 80 cm diameter; buttresses very short. Bark smooth, light brown,
thin, with transverse elevated ridges and scattered large, protruding lenticels; inner bark
light brown, very hard. Sapwood pale or whitish brown, heartwood chocolate with darker
concentric rings. Leaves each with (3)5 pairs of leaflets; petiole 2.33.8 cm and rachis
7.519.5 cm long, slightly puberulous, glabrescent, or almost glabrous. Leaflets upper
surface glossy and glabrous, lower surface glossy, sparsely and minutely puberulous,
glabrescent, sometimes almost glabrous except on the midrib; elliptic, oblong-elliptic or
rarely obovate, 5.410.5 2.14.8 cm, base obtuse, rounded, acute or slightly cuneate,
often symmetrical, apex acuminate to attenuate. Inflorescences paniculate, up to 16 cm
long; both rachis and branches pubescent; pedicel 35 mm long. Flowers: calyx lobes
lanceolate, 3.56 c. 3 mm, unarmed, puberulous outside, densely appressed hairy insides;
petals ovate, c. 5 3 mm, deeply concave to involute, outside densely puberulous in
the lateral parts, inside glabrous; stamens: free filaments and staminodes up to 10 mm
long, sparsely hairy in the lower half, anthers 2 (perfect) largest, ellipsoid, c. 0.35 mm
long; ovary c. 4 mm long, densely woolly, with many minute outgrowths, 35-ovuled,
style up to 14 mm long. Fruits black, ellipsoid or oblong-ellipsoid, rarely slightly reniform,
1114 5.58 cm, flattened, hard, covered with evenly spaced, numerous, slender, sharp
spines (c. 3 mm long), without fine warts among them, 13-seeded. Seeds broadly
suborbicular, c. 1.8 cm long and wide, smooth; aril reniform, c. 3 1.5 cm.
Vernacular name. Sabahsepetir (preferred name).
Distribution. Endemic to Borneo. In Sabah widespread, common; in Sarawak recorded
for Kapit district. Also in Brunei and Kalimantan.
Ecology. In lowland forests, often near the banks of brooks, on sandstone, sandy limestone,
shales, sandy and volcanic soils, up to 700 m altitude. Flowering in May; fruiting in
MarchApril, JuneJuly, September, and November.
171
6. Sindora leiocarpa Backer ex de Wit
(Greek, leio- = smooth, karpos = fruit; the smooth pods)

Bull. Bot. Gard. Buitenz. 3, 18 (1949) 58; Cockburn l.c. 181; Anderson l.c. 239; Whitmore, Tantra &
Sutisna l.c. 207; Kessler & Sidiyasa l.c. 74; Ding Hou l.c. (1996) 702; Coode et al. (eds.) l.c. 164;
Argent et al. (eds.) l.c. 374. Type: Grashoff 667, Sumatra, Kubulands (holotype L).
Tree up to 45 m tall, 80 cm diameter. Bark smooth, finely cracked or fissured, redbrown, grey-mottled, or black; inner bark yellowish brown, brown, red, fibrous and brittle.
Sapwood white or yellow. Stipules lanceolate, 35 mm long, caducous. Leaves each
with (2)3(4) pairs of leaflets; petiole 1.52.8 cm and rachis 411 cm long, both minutely
puberulous, glabrescent, or almost glabrous. Leaflets glabrous on both surfaces, except
sometimes sparsely hairy on the midrib of lower surface; elliptic, obovate-elliptic or broadly
elliptic, 2.55(9.5) 23(5) cm, base obtuse or cuneate, slightly asymmetrical, apex
obtuse, acute, or shortly acuminate, sometimes slightly emarginate, without a gland at
the tip on the undersurface. Inflorescences paniculate, up to 16(25) cm long; both rachis
and branches densely pubescent; bracts and bracteoles ovate, up to 4 mm long, densely
puberulous; pedicel 11.5 mm long, densely puberulous. Flowers yellow, yellowish brown;
calyx lobes elliptic or elliptic-oblong, obovate, or ovate, 4.55(6) 23(3.5) mm, without
spiny outgrowths on the outer surface, densely puberulous outside, appressed hairy inside;
petals slightly oblong, 56 mm long, slightly longitudinally folded and somewhat bent
inward from the middle, the lower half (claw) densely puberulous outside and glabrous
inside, the upper half (limb) glabrous except for the villose margins; stamens: free filaments
and staminodes up to 7 mm long, ciliate; ovary c. 2 mm long, densely woolly along
the margins, style 810 mm long. Fruits ellipsoid, ovoid, broadly ovoid, or suborbicular,
59 4.55.8 cm, flattened, smooth, almost glabrous, 13-seeded. Seeds broadly ovoid
or suborbicular, c. 1.3(2.1) cm long, smooth, black; aril broader than high, often larger
than the seed, smooth or rugose.
Vernacular names. Sabahsepetir, sepetir paya (Malay). Sarawakensurut, tampar
hantu, tampar hantu paya (Iban).
Distribution. Sumatra and Borneo. In Sabah recorded for Beaufort, Kinabatangan, Lahad
Datu, Sandakan, and Tawau districts; in Sarawak throughout. Also in Brunei and Kalimantan.
Ecology. In lowland, well-drained or seasonal freshwater swamp forests, on sandy loam
soils, usually below 100 m altitude, rarely up to 400 m. Flowering in January, March
June, and AugustDecember; fruiting in MarchMay and AugustDecember.
Uses. The wood is used for making boards, partition walls and household utensils.
7. Sindora velutina Baker

(Latin, velutinus = velvety; the densely hairy branchlets and leaf under surface)
In Hooker f., Fl. Brit. Ind. 2 (1878) 269; Prain l.c. (1897) 205 & 482; Ridley l.c. (1922) 638; Merrill
l.c. (1929) 99; Masamune l.c. 353; de Wit l.c. (1949) 72; Whitmore l.c. 273; Meijer l.c. 195; Cockburn
l.c. 181; Anderson l.c. 239; Whitmore, Tantra & Sutisna l.c. 207; Turner l.c. 310; Ding Hou l.c.
(1996) 707; Argent et al. (eds.) l.c. 374. Type: Maingay 607, Peninsular Malaysia, Malacca (holotype
172
K). Synonyms: S. mucronata Pierre, Fl. For. Coch. (1899) sub t. 385; S. parvifoliola Symington,
Kew Bull. (1938) 77.
Tree up to 54 m tall, 95 cm diameter; buttresses very short. Bark smooth to slightly

cracking with horizontal rings, greyish white, brown or dark purplish brown; inner bark
white, brown or red-brown, firmly fibrous or laminated, granular at the edge. Sapwood
white or cream colour, fairly hard, heartwood dark brown. Branchlets glabrous to densely
tomentose. Stipules lanceolate, c. 8 mm long, caducous. Leaves each with (3)56
(7) pairs of leaflets; petiole 23(5) cm long and rachis 516(22.5) cm long, both
tawny woolly or densely tomentose when young and later thinner. Leaflets upper surface
glabrous, rarely sparsely hairy near the midrib, lower surface usually densely tawny hirsute,
sometimes slightly glabrescent; ovate-elliptic or elliptic, sometimes ovate or broadly ovate,
39(14.5) 1.54.5(7.5) cm, base rotund or obtuse, usually symmetrical, apex shortly
or normally acuminate, sometimes acute. Inflorescences paniculate, up to 11(20) cm
long; both rachis and branches densely brown-hirsute; bracts lanceolate, c. 3 1.5 mm,
densely appressed hirsute outside; bracteoles resembling bracts but about half the length;
pedicel very short. Flowers: calyx lobes oblong-lanceolate, 1113 mm long, unarmed,
densely hirsute often on both surfaces; petals narrowly lanceolate, 1012 mm long, densely
woolly outside, glabrous inside; stamens: free filaments and staminodes up to 1.5 cm
long, hirsute towards the lower 1/21/3; ovary c. 3 mm long, woolly, with many minute
outgrowths, style up to 15 mm long. Fruits elliptic-ovate or broadly ovate, slightly irregular
in shape, 611.7(15) 56.5(8) cm, with rather small, scattered, moderately developed
sharp spines (c. 4 mm long), with fine warts among them, 14-seeded. Seeds irregularly
ovoid, 1.62.2 cm across, flattened, smooth, purple-black, almost glossy; aril longer than
wide (c. 2.6 cm wide).
Vernacular name. Sabahsepetir (Malay).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah known from Sandakan,
Lahad Datu and Tawau districts; in Sarawak recorded for Bintulu district. Also in Brunei.
Ecology. In forest on dryland, hillsides, rarely along riverbanks and in swamps, growing
usually scattered, in low altitude usually below 100 m, sometimes up to 360 m. Flowering
in MarchMay, July, and OctoberNovember; fruiting in February, MayJune, September,
and NovemberDecember.
Uses. The wood is said to be soft and subject to attack by termites and other insects.
It splits when drying. It is used as beams and rafters for house-building and for household
utensils and boards. (cf.: Foxworthy, Mal. For. Rec. 3 (1927) 92; K. Heyne, Nutt. Pl.
Ned.-Ind. 1 (1950), 729; Burkill, EPMP 2 (1966) 2070; de Wit, Bull. Bot. Gard. Buitenz.
3,18 (1949) 76).
8. Sindora wallichii Benth.
Fig. 12.
(N. Wallich, 17861854, former Superintendent of the Calcutta Botanic Garden, India)
In Hooker, Icon. Pl. 11 (1867) sub t. 1018, excl. t. 1017; Prain l.c. (1897) 203 & 481, as S. wallichiana;
Ridley l.c. (1922) 637; Symington l.c. (1938) 75 & 77; de Wit l.c. (1949) 76; Whitmore l.c. 273; Cockburn
l.c. 181; Keng l.c. (1990) 39; Whitmore, Tantra & Sutisna l.c. 207; Kessler & Sidiyasa l.c. 74; Turner l.c.
173
vi
310; Ding Hou l.c. (1996) 708; Loo & Tan l.c. 99; Argent et al. (eds.) l.c. 374. Lectotype (de Wit,
1949): Wallich 5805, Singapore (holotype K; isotype BM). Synonyms: S. wallichii Benth. var. intermedia
Baker in Hooker f., Fl. Brit. Ind. 2 (1878) 268; S. intermedia (Baker) Prain l.c. (1897) 204 & 482, Ridley
l.c. (1922) 638, p.p.
Tree up to 35 m tall, 80 cm diameter; bole straight; buttresses steep. Bark smooth, grey
or dark grey; inner bark yellow, mottled. Stipules foliaceous, up to 15 mm long, tardily
caducous. Leaves each with 3 or 4 pairs of leaflets; petiole 1.22.5 cm and rachis 4.5
8.5 cm long, minutely puberulous, glabrescent, or sometimes glabrous in the upper half.
Leaflets upper surface glabrous, sometimes puberulous in the lower half or very minutely
puberulous all over, lower surface puberulous to densely pubescent; elliptic-ovate or ellipticoblong, ovate-oblong or obovate-oblong, 410.5 2.55 cm, upper pair usually the largest,
base obtuse or rotund, often more or less unequally sided, apex acute, subacuminate,
obtuse, the very tip slightly notched. Inflorescences paniculate, up to 24 cm long, both
rachis and branches densely puberulous; pedicel very short. Flowers: calyx lobes lanceolate,
to 9.5 3 mm, outside puberulous and bearing some distinctly spiny outgrowths often
in the upper half, insides glabrous; petals elliptic or oblong-elliptic, c. 8 22.5 mm,
outside woolly, inside glabrous; stamens: free filaments up to 10 mm long, slightly hairy
at the lower part; ovary c. 5 mm long, densely hairy, with many minute, spiny outgrowths,
24-ovuled, style up to 10 mm long. Fruits suborbicular, broadly obovoid or ellipsoid,
610 46 cm, flattened, covered with close, hard, straight spines (up to 5 mm long),
12(3)-seeded. Seeds suborbicular, c. 2 cm in diameter; aril often broader than long.
Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo. In Sabah known from
Keningau, Lahad Datu, Sandakan, and Tawau districts; not recorded in Sarawak. Also
in Kalimantan.
Ecology. In lowland forests, on well-drained sandy or sometimes clayey soils, sometimes
near mangrove swamp, often below 100 m, sometimes up to 300 m altitude. Flowering
in March, May, July, and OctoberDecember; fruiting in January, March, August, October,
and December.
13. SYMPETALANDRA Stapf

(Greek, sum- or sun- = joined, petalon = petal, andros = male; flower with petals and
stamens joined at the base)
In Hooker, Icon. Pl. 4, 28 (1905) t. 2721; Whitmore, TFM 1 (1972) 273; Steenis, Blumea 22 (1975) 162;
Cockburn, TS 1 (1976) 183; Anderson, CLTS (1980) 239; Polhill & J.E. Vidal in Polhill & Raven (eds.),
Adv. Leg. Syst. 1 (1981) 95; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 208; Ding Hou, FM 1, 12
(1996) 709; Argent et al. (eds.), MNDT-CK 2 (1997) 379.
Trees unarmed; buttresses present. Stipules minute, caducous. Leaves compound, once
pinnate and paripinnate with (1)2(3) pairs of leaflets, or bipinnate with 13 pairs
of pinnae, each pinna with 16 pairs of leaflets. Leaflets pellucid-punctate. Inflorescences
axillary and/or terminal, racemose, simple or fascicled, or paniculate; bracts persistent
175
beyond anthesis; bracteoles absent at anthesis. Flowers bisexual, calyx and corolla pellucidglandular; hypanthium obscure or absent; calyx 5-lobed; petals 5, 2.55 mm long; stamens
10, alternately long and short, joined with petals at the base, filaments glabrous, anthers
with a large, dorsal, dark brown connective, and a deciduous gland at apex, glabrous
or slightly hairy on the dorsal connective; ovary hairy, 26-ovuled. Fruits woody, smooth
outside, dehiscent and 2-valved, compressed, prominently veined, 14-seeded; valves
straight or only slightly twisting during dehiscence. Seeds orbicular to broadly ellipsoid,
compressed but thick, 33.5 cm diameter, not arillate.
Distribution. A genus comprising 5 species, distributed in Sumatra, Peninsular Malaysia,
Borneo, Lesser Sunda Islands (Flores), and the Philippines. Two species in Sabah and
Sarawak.
Key to Sympetalandra species

Leaves once pinnate. Inflorescence rachis glabrous. Anthers slightly hairy on the dorsal
connective. Ovary 2-ovuled......................1. S. borneensis
Leaves bipinnate (some leaves once-pinnate in juvenile state). Inflorescence rachis pubescent.
Anthers not hairy. Ovary 36-ovuled........................................2. S. unijuga
1. Sympetalandra borneensis Stapf

(of Borneo)
In Hooker, Icon. Pl. 4, 28 (1905) t. 2721; Merrill l.c. (1921) 295, l.c. (1929) 98; Masamune l.c. 354;
Whitmore, l.c. 274; Steenis l.c. (1975) 163; Anderson l.c. 239; Whitmore, Tantra & Sutisna l.c. 208;
Ding Hou l.c. (1996) 710; Argent et al. (eds.) l.c. 379. Type: Haviland & Hose 1628, Borneo, Sarawak,
Kuching (holotype K; isotypes L, SAR).
Tree up to 25 m high, 25 cm diameter; buttresses up to 0.8 m high, 90 cm wide, 15

cm thick. Leaves once pinnate, each with (1)2(3) pairs of leaflets; petiole 4.57 cm
long and rachis 2.53.5 cm long, both glabrous. Leaflets elliptic, 617 37 cm.
Inflorescences axillary and extra-axillary, densely racemose, up to 14 cm long, both
peduncle and rachis glabrous; bracts ovate, c. 1 mm long; pedicel very short. Flowers:
calyx c. 1.5 mm long, glabrous outside; lobes triangular or crescent, 0.30.5 mm long;
petals oblong or slightly obovate-oblong, 2.53.5 1.5 mm; stamen filaments 34 mm
long, glabrous, anthers c. 0.6 mm long, slightly hairy at the upper half of dorsal connective;
ovary 2-ovuled. Fruits unknown.
Vernacular name. Sabahmerbau lalat (preferred name).
Distribution. Endemic to Borneo. In Sarawak confined to coastal areas, throughout; so
far erroneously recorded for Sabah (see treatment under next species).
Ecology. In forests, including kerangas forest, at low and medium altitude. Flowering
in MarchApril, and SeptemberOctober.
176
vi
2. Sympetalandra unijuga (Airy Shaw) Steenis
Fig. 13.
(Latin, uni- = one, -jugum = paired; leaf with one pair of leaflets)
Blumea 22 (1975) 163; Whitmore, Tantra & Sutisna l.c. 208; Turner l.c. 311; Ding Hou l.c. (1996) 713.
Basionym: Erythrophleum unijugum Airy Shaw, Kew Bull. (1939) 180. Type: Agullana 1780, Borneo,
Sabah (holotype K; isotype L). Synonyms: Erythrophleum densiflorum auct. non (Elm.) Merr.: Merrill
l.c. (1921) 295, l.c. (1929) 98; S. borneensis auct. non Stapf: Kostermans, Gard. Bull. Sing. 17 (1958) 5,
p.p., Cockburn l.c. 183; Serianthes gigalobium Kosterm., Reinwardtia 2 (1953) 357.
Tree up to 30 m tall, 70 cm diameter; buttresses up to 4 m high, extending up to 100

cm from the trunk. Leaves bipinnate (some leaves once pinnate in juvenile state), with
1 (2 or 3) pair(s) of pinnae, each pinna with (2)34(5) pairs of leaflets; petiole 2
5(8) cm long; main rachis 610 cm long; stalk of pinnae 25 cm long; rachis of pinnae
615(18) cm long. Leaflets ovate-oblong to oblong-elliptic, 610(17) 24(8) cm,
base more or less rounded, apex acuminate; petiolules 35 mm long. Inflorescences
racemose and paniculate, up to 10 cm long, rachis loosely to densely hairy; pedicel
very short. Flowers: calyx lobes not ciliate; petals yellowish white; anthers glabrous;
ovary 36-ovuled. Fruits falcate, 1536 3.54(5) cm, coarsely and prominently
reticulate-veined; pericarp woody, c. 2 mm thick. Seeds 15 per fruit, c. 4 2.5 2
cm, even larger in 1-seeded pods.
Vernacular name. Sabahmerbau lalat (preferred name).
Distribution. Sumatra, Peninsular Malaysia, Borneo, and the Philippines. In Sabah
widespread; in Sarawak rare.
Ecology. In well-drained forests, from lowland up to 600 m altitude. Flowering in April
May, June and September; fruiting in July and SeptemberNovember.
14. UITTIENIA Steenis

(H.Uittien, 18981944, botanist,
formerly curator of the Utrecht Herbarium, the Netherlands)
Bull. Bot. Gard. Buitenz. 3, 17 (1948) 416; Hutchinson, Gen. Fl. Pl. (1964) 237; Irwin & Barneby in
Polhill & Raven (eds.), Adv. Leg. Syst. 1 (1981) 101 (as syn. of Dialium); Ding Hou, FM 1, 12 (1996)
714. Synonym: Dialium subgen. Uittienia (Steenis) Steyaert, Reinwardtia 2 (1953) 355.
Trees. Stipules minute, linear, caducous. Leaves unifoliolate; petiole thickened at basal
end; petiolules very short, wrinkled. Inflorescences axillary, fascicled short-thyrses.
Flowers: hypanthium obscure or 0; calyx lobes 5; petals 5; disk distinct, cushion-like,
hairy; stamens 5(7), all fertile, rarely 13 smaller; ovary slightly hairy, 1-ovuled. Fruits
not winged, drupaceous, indehiscent, globose, woody, with a crustaceous, glabrous exocarp
united with a thick, hard mesocarp around the single subglobular seed; endosperm almost
filling the seed; embryo minute, conical.
178
vi
Distribution. Only 1 species known in Sumatra and Borneo. In Sabah only known from
Tawau district; not yet found in Sarawak. Also in Kalimantan.
Ecology. In primary forest, up to 400 m altitude.
Uittienia modesta Steenis
Fig. 14.
(Latin, modestus = modest; alluding the modest personality of Uittien)

714. Type: bb 19274, Borneo, E Kalimantan (holotype BO; isotype L). Synonym: Dialium modestum
(Steenis) Steyaert, Reinwardtia 2 (1953) 355.
Tree up to 40 m tall, 100 cm diameter; buttresses up to 5 m high, extending outward

100200 cm. Leaves glabrous; petiole 0.82 cm long. Leaflets ovate to ovate-oblong,
elliptic or lanceolate, 720 3.57.5 cm; base obtuse or rounded, apex acuminate; lateral
veins 47 per side; intercostal venation loosely reticulate, obscure or visible on both
surfaces; petiolules very short. Inflorescences with rachis or branches up to 7 cm long,
sparsely puberulous; bracts and bracteoles minute, deciduous. Flowers: calyx lobes oblong
or ovate, 3.5 1.52 mm; petals subequally obovate, 46 23 mm, glabrous, distinctly
pinnately nerved with a strong midrib; disk c. 2 mm diameter; stamen filaments glabrous.
Fruits often only one developed from an inflorescence, on a thickened woody stalk (c.
1 cm diameter), c. 6 cm diameter, with a roughish, somewhat scurfy-granular brown
surface, glabrous; pericarp rather hard and thick (45 mm). Seed one in each fruit,
subglobose, c. 4 cm diameter.
Distribution. Sumatra and Borneo. In Sabah known for Tawau district; not recorded in
Ecology. In primary forest, often occurring below 100 m altitude, sometimes up to 400(
600) m. Flowering in April, JuneJuly, and November; fruiting in JuneJuly and September
October.
180
MORACEAE
K.M. Kochummen1 & Rusea Go2
(Edited with the assistance of Dr. C.C. Berg)
1
Formerly Forest Research Institute Malaysia, Kepong, Malaysia
Lindley, Veg. Kingd. (1847) 266; Merrill, EB (1921) 217; Ridley, FMP 3 (1924) 317 (under Urticaceae); Masamune, EPB (1942)
238; Browne, FTSB (1955) 348 (under Urticaceae); Backer & Bakhuizen f., FJ 2 (1965) 12; Corner, Gard. Bull. Sing. 19 (1962)
187, WSTM 3rd. edition 2 (1988) 509; Burgess, TBS (1966) 399; Kochummen, TFM 3 (1978) 119; Anderson, CLTS (1980) 254;
Primack, Foresters Guide to the Moraceae of Sarawak (1983) 2; Rohwer in Kubitzki et al. (eds.), Fam. & Gen. Vasc. Pl. 2
(1993) 438; Kessler & Sidiyasa, Trees of the Balik Papan-Samarinda Area, East Kalimantan (1994) 172; Turner, Gard. Bull.
Sing. 47 (1995) 347; Coode et al. (eds.), CLBD (1996) 210; Argent et al. (eds.), MNDT-CK 2 (1997) 429.
Dioecious or monoecious trees, shrubs, climbers, stranglers, epiphytes or herbs with white or
rarely pale yellow-brown latex (except some species of Streblus); bole (if present) with or
without buttresses. Bark smooth, rough, fissured or scaly, often lenticellate, sometimes spiny.
Buds small to very large. Stipules small or large, sometimes connate to form a cap over the bud.
Twigs glabrous or hairy, slender or stout, often with ring-like scars of fallen stipules. Leaves
alternate (distichous or spiral), very rarely subopposite or subverticillate; simple, entire, lobed,
or deeply incised down to the midrib, or rarely compound-imparipinnate; pinnately veined or
with distinct basal pair(s) of lateral veins; margin plane, revolute or faintly toothed.
Inflorescences axillary, or on older branches or trunk, or borne on runners/stolons arising from
the base of stem, unisexual or bisexual, paniculate, racemose, spicate, capitate or urceolate.
Flowers small, unisexual or very rarely bisexual; perianth usually with (0)4(10) free or fused,
valvate or imbricate, persistent tepals (= perianth lobes). Male (staminate) flowers with
(1)4(8) stamens opposite the tepals, filaments free or united, straight or inflexed in bud,
anthers small, mucronate or bilobed, varying in shape from crescentic to turbinate (with
transverse equatorial dehiscence); pistillode (rudimentary pistil) present or absent. Female
(pistillate) flowers without staminodes (rudimentary stamens); ovary unilocular, very rarely 2locular, superior or inferior or immersed in sockets of the inflorescence axes, styles 2 or 1,
stigmas 2 or 1, not capitate or peltate; ovule 1, anatropous or campylotropous, apical or
subapical. Fruits drupaceous or in Ficus very often dry and achene-like, simple and often
enclosed in an urn-shaped receptacle (syconium, in Ficus) or united into a large fleshy
compound structure called syncarp (= multiple fruit), with the persistent perianth becoming
enlarged, fleshy (and edible in several Artocarpus species). Seeds large or small, testa
membranous or disintegrated; endosperm present or absent; embryo curved or straight, radicle
long or short, cotyledons plicate, conduplicate or plane, foliaceous or thickened, equal or
unequal.
Distribution. About 37 genera with c. 1050 species, mostly in the tropics. Eight genera with 179
species are recorded for Sabah and Sarawak.
Current address: 2Universiti Putra Malaysia, Serdang, Malaysia
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Ecology. Occurs in a wide variety of habitats and forest types from lowland to upper montane
forests up to 3200 m altitude. The flowers are either wind- or insect-pollinated (agaonid wasps,
stingless bees, flies, and a few others). Fruits are dispersed by birds, bats and a variety of
terrestrial and arboreal mammals.
Uses. Several members of the family, e.g., species of Antiaris, Artocarpus, and Parartocarpus,
produce valuable timber. Fruits of a number of species of Artocarpus (e.g., A. altilis, A.
heterophyllus, A. integer, and A. odoratissimus) are edible, and the trees are commonly
cultivated in village orchards or occasionally in large scale commercial plantation. The leaves of
mulberry tree (Morus alba), a native of China, are a source of food for silkworms. Many species
of Ficus have potential for use in landscaping while some others are used in traditional medicine.
The latex of Antiaris and Parartocarpus species is poisonous and used by the aborigines for
making arrow poison. Bark of Antiaris and certain species of Artocarpus are used in making
cloth.
Taxonomy. The Moraceae are closely related to the Ulmaceae and Urticaceae but differ from
both in having latex, stigma not capitate or peltate, ovule anatropous or campylotropous, and in
the more or less connate large fleshy multiple fruits. In the more recent system of classification
(Rohwer, 1993), the family is classified into 5 tribes, viz. the Artocarpeae, Castilleae,
Dorstenieae, Ficeae and Moreae. The disposition of the eight genera occurring in Sabah and
Sarawak is as follows:
Tribe Artocarpeae
Tribe Castilleae
Tribe Dorstenieae
Tribe Ficeae
Tribe Moreae
:
:
:
:
:
Artocarpus, Parartocarpus, Prainea

Antiaris
No representative
Ficus
Maclura, Streblus, Trophis
Key to the genera

(for fertile specimens)
1. Flowers enclosed inside urceolate receptacles or syconia..............3. Ficus
Flowers not enclosed inside syconia..............2
2. Male flowers in spicate inflorescences..........3
Male flowers usually in capitate or sometimes paniculate inflorescences................4
3. Shrubs or trees...................6. Streblus
Climbers...................
Trophis P. Br., nom. conserv.
Nat. Hist. Jamaica (1756) 357; Corner, Gard. Bull. Sing. 19 (1962) 230; Berg, Proc. Kon. Ned. Ak. C 91 (1988) 352;
Rohwer in Kubitzki et al. (eds.), Fam. & Gen. Vasc. Pl. 2 (1993) 446. Synonyms: Olmedia Ruiz & Pav., Prod. (1798);
Malaisia Blanco, Fl. Filip. (1837) 789, Corner l.c. (1962) 240, Backer & Bakhuizen f. l.c. 15; Cephalotrophis Blume,
Mus. Bot. Lugd.-Bat. 2 (1856) 75.
About 9 species, in tropical America, Madagascar, Runion, and Asia.
Dioecious, unarmed twining climbers (in Sabah), or trees or shrubs (outside Sabah and
Sarawak) with white latex. Leaves simple, distichous, pinnately veined; stipules free, lateral. Male
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MORACEAE (KOCHUMMEN & GO)
inflorescences spicate with sterile abaxial grooves; female ones almost capitate; bracts
linear to spathulate, incurved. Male flowers 34-merous, tepals valvate, joined at the
base; filaments inflexed in bud; pistillode minute. Female flowers: perianth utricular,
obscurely 4-lobed; style with 2(1) long stigmatic arms. Fruits (syncarps) fleshy, set with
few projecting drupes, enclosed in a perianth.
In Sabah and Sarawak, represented by one species, T. scandens (Lour.) Hook. & Arn.
(basionym: Malaisia scandens Lour.), which is rare and known so far by one collection,
SAN 8393, from Keningau district in Sabah.
4. Climbing plants with axillary spines....
Maclura Nutt., nom. conserv.
Gen. N. Amer. Pl. 2 (1818) 233; Corner l.c. (1962) 235; Backer & Bakhuizen f. l.c. 17; Berg l. c. (1988) 352; Rohwer
l.c. 446.
About eleven species, mainly found in the tropics, with one species in N America.
Dioecious climbers (in Sabah), or scrambling shrubs or trees (outside Sabah and
Sarawak), usually with axillary spines and white latex. Leaves simple, spirally arranged
or distichous, pinnately veined; stipules free, lateral. Male inflorescences paniculate, with
a sterile groove, or capitate; female ones capitate or shortly spicate; bracts spathulate to
clavate or absent. Male flowers 4-merous; tepals free; anthers extrorse to laterally
dehiscent, filaments inflexed or straight in bud; pistillode subulate or absent. Female
flowers sessile, free or connate proximally; tepals 4, decussate, free or fused to the
middle; ovary free or sunken in sockets of the receptacle; stigmatic arms 12, short or
long. Fruits simple, drupaceous, very thinly pulpy, enclosed in the fleshy perianth, mostly
not projecting, often concrescent below into a fleshy syncarp.
In Sabah one species, M. cochinchinensis (Lour.) Corner, very rare, known only by one
collection, J.H. Beaman 7932, from Mt. Kinabalu.
Shrubs or trees................5
5. Styles 2. Fruits simple, one-seeded................1. Antiaris
Style 1. Fruits many-seeded syncarps (= multiple fruits)..............6
6. Stamens 23. Capitate inflorescences with distinct and persistent involucre of bracts at the
base. Bark with abundant large lenticels.........4. Parartocarpus
Stamen 1. Capitate inflorescences without distinct involucre of bracts at the base. Bark
without large lenticels............................7
7. Dioecious trees. Male and female inflorescences similar in shape. Drupes projecting out from
the syncarps................5. Prainea
Monoecious trees. Male and female inflorescences different in shape. Drupes immersed in
the syncarps......................2. Artocarpus
Forest key to the genera

(based mainly on field and vegetative characters)
1. Tall forest trees..................2
Shrubs, small trees, climbers, creepers, epiphytes, or stranglers...................7
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2. Flowers enclosed in a syconium...........3. Ficus (in part)

Flowers not enclosed in a syconium..................3
3. Fruits many-seeded syncarps, often prickly...................4
Fruits simple, one-seeded, with smooth surface................6
4. Bark with abundant large lenticels. Capitate inflorescences with persistent involucre of
bracts.......4. Parartocarpus
Bark
without
large
lenticels.
Capitate
inflorescences
without
persistent
involucre
of
bracts............................5
5. Dioecious trees. Drupes projecting out of the syncarp surface. Leaves with prominently
looping lateral veins............5. Prainea
Monoecious trees. Drupes immersed in the syncarp surface. Leaf lateral veins not
prominently looping.........2. Artocarpus
6. Leaves hairy. Inflorescences discoid-capitate................1. Antiaris
Leaves glabrous. Inflorescences spicate..............6. Streblus (in part)
7. Shrubs or small trees......................8
Climbers, creepers, epiphytes or stranglers...............9
8. Flowers enclosed in a syconium...........3. Ficus (in part)
Flowers not enclosed in a syconium................6. Streblus (in part)
9. Stem spiny.................Maclura
Stem not spiny.................10
10. Flowers enclosed in a syconium...3. Ficus (in part)
Flowers not enclosed in a syconium...............Trophis
1. ANTIARIS Lesch., nom. conserv.

(from the Indonesian plant name antjar)
ipoh (Iban, Malay)
Rusea Go
Ann. Mus. Hist. Nat. Paris 16 (1810) 476; Merrill, EB (1921) 220; Ridley, FMP 3 (1924) 350; Masamune, EPB (1942) 238;
Corner, Gard. Bull. Sing. 19 (1962) 244, WSTM 3rd. edition 2 (1988) 510; Backer & Bakhuizen f., FJ 2 (1965) 19; Berg, Bull.
Jard. Bot. Etat. 47 (1977) 308; Kochummen, TFM 3 (1978) 120; Anderson, CLTS (1980) 254; Primack, Foresters Guide to the
Moraceae of Sarawak (1983) 13; Whitmore, Tantra & Sutisna, CLK 2,1 (1990) 236; Rohwer in Kubitzki et al. (eds.), Fam. &
Gen. Vasc. Pl. 2 (1993) 450; Coode et al. (eds.), CLBD (1996) 210; Argent et al. (eds.), MNDT-CK 2 (1997) 429; PROSEA 5, 3
(1998) 73. Synonyms: Ipo Pers., Syn. Pl. 2 (1807) 566; Antcha Horsfield, Verh. Bat. Gen. 7 (1814) 8; Toxicaria Aepnel ex
Steud., Nomencl. Bot. (1821) 434; Lepurandra Graham, Cat. Pl. Bombay (1839) 193.
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Monoecious or deciduous trees, without spines. Bark smooth, grey; sap creamy white and
watery, turning dirty brownish and granular on exposure. Stipules free, subamplexicaul (slightly
clasping the twig). Leaves distichous, pinnately veined, base rounded or cordate. Inflorescences
unisexual, axillary, the females solitary, the males in groups of 24, or the males below a single
female in the leaf axil. Male inflorescences discoid-capitate, pedunculate, many-flowered,
involucrate with small bracts arranged in a single series; interfloral bracts absent. Male flowers
sessile; tepals 45, minute, free, imbricate; stamens (3)4(8), filaments very short or almost
none, anthers introrse; pistillode absent. Female inflorescences 1(2)-flowered, sessile or
pedunculate; bracts small, few, scattered. Female flower with 45 minute tepals; ovary
(sub)inferior, concrescent with the receptacle, style very short, with 2 long stigmatic arms; ovule
immersed, apical, transversely anatropous. Fruits simple, drupaceous, globose to ellipsoid or
pear-shaped, surface velvety, fleshy; endocarp thinly crustaceous. Seeds rather large; embryo
straight, radicle very short, cotyledons thick, equal, not folded.
Distribution. One variable species, found throughout the Old World tropics, ranging from
tropical Africa to Madagascar, Yemen, Sri Lanka, India, Burma, Thailand, Indo-China, S China,
throughout the Malesian region, Australia, and Tonga.
Antiaris toxicaria Lesch.

(Latin, toxicarius = poisonous; the latex)
Fig. 1.
Ann. Mus. Hist. Nat. Paris 16 (1810) 478, t. 22; Merrill l.c. (1921) 220; Ridley l.c. (1924) 350; Masamune l.c. 238; Corner l.c.
(1962) 244, l.c (1988) 510; Backer & Bakhuizen f. l.c. 20; Kochummen l.c. 120; Anderson l.c. 254; Primack l.c. 13; Whitmore,
Tantra & Sutisna l.c. 236; Turner, Gard. Bull. Sing. 47 (1995) 347; Coode et al. (eds.) l.c. 210; Argent et al. (eds.) l.c. 429.
Tree up to 50 m tall, 100 cm diameter, often deciduous; buttresses 13 m high. Bark greyish
white, smooth to slightly fissured and lenticellate-pustulate; inner bark whitish, turning redbrown on exposure, fibrous; sap creamy white to yellowish brown, gummy or sticky, turning
dirty brown on exposure. Sapwood creamy white. Young twigs, petioles and veins on the lower
leaf surface hispid with brown hairs 12 mm long. Terminal buds ovoid, often oblique. Stipules
paired, somewhat interpetiolar, linear to lanceolate, 11.2 0.2 cm, hispid with brown hairs,
caducous, scars distinct. Leaves membranous to thin-coriaceous, appressed hairy to somewhat
scabrid on both surfaces; elliptic, subovate to obovate, (10)1215(18) 610 cm, base
cordate to cuneate, often somewhat asymmetric, margin denticulate to subentire, apex acuminate;
lateral veins 712 pairs, curving towards the leaf margin; intercostal venation reticulate, distinct
beneath; petiole up to 1.5 cm long. Inflorescences borne on the new shoots together with new
leaves, minutely appressed hairy. Male inflorescences discoid to reflexed pulvinate, 12 cm
across; peduncle slender, c. 1.5 0.1 cm; involucral bracts linear, short. Male flowers up to 2
mm long; tepals spathulate. Female inflorescences 1-flowered, subsessile, with several small
lanceolate bracts. Female flowers: tepals minute, lanceolate, free; styles soft-pubescent,
stigmatic arms 1.31.7 cm long. Fruits borne on the twigs behind the leaves, ellipsoid to
pyriform, 1.24.5 12 cm, densely brown velutinous, ripening red to crimson, purplish and
finally black; peduncle short, less than 1 cm long; pulpy wall c. 1.5 mm thick; mesocarp fleshy;
endocarp crustaceous, thin. Seeds 11.5 1 cm, smooth.
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Vernacular names. Sabahipoh (Malay). Sarawakipoh (Iban, Malay).

Distribution. As given under the genus. Of scattered occurrence in Sabah and Sarawak. Also in
Ecology. From lowland to lower montane forests up to 1500 m altitude, often near streams.
Flowering in March to August; fruiting in May to December.
Uses. The poisonous components of the latex are cardiac glycosides (cardenolides), among
which the antiarin in two forms is predominant (Bisset, Ann. Bog. 2 (1957) 219; Planta Medica
10 (1962) 143; Lloydia 29 (1966) 1 & 172). It arrests heart beat, like digitalin, but more potently.
In Sarawak, the sap is tapped by making diagonal slashes on the bark, and is used by the
aborigines to prepare arrow and dart poison. The untreated latex is used without alteration or
after mixing it with sap of other plants, such as that of Strychnos species (Loganiaceae). The dart
tips are dipped into the latex, turning the dart tips black. The poison remains virulent for many
years. The bark can be made into bark cloth after thorough soaking in water to remove the
poison. The trees yield a lightweight hardwood timber and sold under the trade name terap. It is
locally used for light construction, interior finishing, furniture, moulding, panelling, flooring, and
for making tool-handles. The timber is also suitable for manufacturing plywood, veneer and
blockboard.
2. ARTOCARPUS J.R. & G. Forster, nom. conserv.

(Greek, artos = bread, karpos = fruit; the bread-fruit tree)
beruni (Dusun), pudau, selangking (Iban), terap (Malay)
K.M. Kochummen
Char. Gen. (1786) 101; Blume, Bijdr. Fl. Ned. Ind. (1825) 479; Miquel, Fl. Ind. Bat. 1, 2 (1859) 284; King in Hooker f., Fl. Br.
Ind. 5 (1888) 539, Ann. Roy. Bot. Gard. Cal. 2 (1889) 1; Beccari, For. Born. (1902) 623; Merrill, En. Philip. Pl. 2 (1923) 40;
Ridley, FMP 3 (1924) 351; Burkill, EPMP 1 (1935) 247; Browne, FTSB (1955) 349; Jarrett, J. Arn. Arb. 40 (1959) 113, ibid. 41
(1960) 73; Backer & Bakhuizen f., FJ 2 (1965) 18; Burgess, TBS (1966) 399; Kochummen, TFM 3 (1978) 120; Anderson, CLTS
(1980) 254; Wong, DMT (1982) 9; Primack, Foresters Guide to the Moraceae of Sarawak (1983) 16; Corner, WSTM 3rd.
edition 2 (1988) 511; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 236; PROSEA 2 (1991) 79; Rohwer in Kubitzki et al. (eds.),
Fam. & Gen. Vasc. Pl. (1993) 447; Kessler & Sidiyasa, Trees of Balikpapan-Samarinda Area, East Kalimantan (1994) 173;
PROSEA 5, 2 (1995) 59; Coode et al. (eds.), CLBD (1996) 216; Argent et al. (eds.), MNDT-CK 2 (1997) 430.
Monoecious, small to medium-sized or large trees with thick white latex in all living parts,
without spines; bole with or without buttresses. Bark without large lenticels, grey white, orangebrown or dark brown, smooth to fissured, scaly or dippled; inner bark yellowish brown. Stipules
either intrapetiolar, large, amplexicaul (clasping the twigs) and on falling leaving annular scars
on the twigs (subgen. Artocarpus), or small, extrapetiolar, non-amplexicaul and on falling
leaving lateral scars on the twig (subgen. Pseudojaca). Leaves spirally arranged (subgen.
Artocarpus) or distichous (subgen. N 134550, CI from SAN 42439.) Pseudojaca), simple, entire or
pinnately lobed or deeply pinnately incised, or compound-imparipinnate (A. anisophyllus).
Inflorescences unisexual, capitate, with the heads cylindrical, club-shaped, ellipsoid or globose,
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short or long-peduncled or sessile, arising from the trunk and older branches or axillary; males
generally smaller than the females; involucral bracts absent; interfloral bracts many, stalked,
peltate, clavate or spathulate. Flowers numerous, compactly covering the surface of the
receptacle; each perianth enclosing a single ovary or stamen, intermixed with interfloral bracts.
Male flowers free; perianth tubular, entire or 24-lobed; stamen 1, exserted, anthers globose to
oblong; pistillode absent. Female flowers at least distally connate with each other and/or with
interfloral bracts; perianth well-developed and tubular (subgen. Artocarpus) or highly reduced or
absent (subgen. Pseudojaca), lower part thin-walled, enclosing the ovary, upper part thickwalled with a narrow opening through which the style projects out, with the apices either
completely fused to form an areolate or smooth surface or free and forming variously shaped
processes on the surface of the female heads; ovary one-loculed, superior (subgen. Artocarpus)
or inferior (subgen. Pseudojaca), style 1, apical to lateral, simple or bifid (the stigmatic arms
equal or unequal), ovule subapical to lateral. Syncarp (multiple fruit) fleshy, formed by the
enlargement of the entire female head (subgen. Artocarpus) or by the swelling of the receptacle
of the female head (subgen. Pseudojaca), small to gigantic in size; in subgen. Artocarpus, at
least the proximal parts of the perianth fleshy, and perianth of unfertilized ovaries elongate and
become ribbon-like. Fruits drupaceous, not projecting from the syncarp surface. Seeds large;
embryo straight or oblique, cotyledons equal or unequal; germination hypogeal, first two leaves
opposite and fully developed or scale-like.
Distribution. About 55 species; native to the Indian subcontinent south of the Himalayas, Sri
Lanka, Burma, Thailand, Indo-China, Southern China, Taiwan, Hainan, Malesia, and Melanesia.
Twenty three (and one incompletely known) species are known in Sabah and Sarawak.
Ecology. Primary and secondary forests, mostly below 100 m, a few species occurring up to
2000 m altitude. Animals are the main seed dispersers. The fleshy perianth or the entire syncarp
is eaten by arboreal mammals or birds. The very large fruit of A. integer arising from the stem is
probably consumed by wild cattles, pigs and elephants. Some species with male heads emit a
sweet scent of honey and burnt sugar which are insect-pollinated. Others have scentless male
heads producing clouds of pollen at anthesis and hence appear to be wind-pollinated.
Uses. Species of Artocarpus produce light and medium hardwood. The light hardwood timber is
known by the trade names terap (Peninsular Malaysia and Sabah) or pudau (Sarawak), while the
medium hardwood timber is known as keledang (Peninsular Malaysia), beruni (Sabah), or
selangking (Sarawak). In terap, the heartwood is yellow to pale brown and usually not
differentiated from the sapwood; the wood is non-durable under tropical conditions. It is easy to
work with, can be peeled satisfactorily and is suitable for plywood production. Locally, terap
timber is used for light construction, for making boxes, crates, wooden pallets and veneer. The
wood is sometimes nicely figured and therefore is suitable for decorative purposes like furniture,
joinery and panelling. In keledang, the heartwood is orange to yellowish brown and well
differentiated from the paler sapwood; the wood is non-durable to moderately durable, and is
comparatively resistant to termite attack but susceptible to powder post beetle attack. It is difficult
to saw, blunting saw tooth, probably due to the presence of tension wood or silica. Keledang
timber is used for telegraph poles, wharves, large canoes, boat building, and light construction,
or when under cover for house and bridge building, beams, poles, flooring, furniture, joinery,
cabinet work, household utensils, musical instruments, turnery, veneer and plywood. It is
favoured as hewn coffin among the Chinese. The roots of old Artocarpus heterophyllus are
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highly prized for carving and picture framing. Many species produce fruits which are edible.
Examples are A. altilis, the breadfruit, A. heterophyllus, the Jackfruit and A. integer, the
chempedak, with the former two species are widely cultivated throughout the tropics. A.
odoratissimus is widely cultivated in Sabah and Sarawak for its edible fruits. Seeds are eaten
after roasting. The bark of some species is used for cordage and formerly widely utilized by the
aborigines to make loin cloths and jackets. The latex, bark, leaves and roots of some species have
medicinal properties, and the latex is used as bird-lime and when mixed with wax is used in batik
printing. Large leaves of a few species such as A. elasticus and A. odoratissimus are used as
thatch, to make partitions in long houses, and for temporary field huts.
Key to Artocarpus species

1. Leaves compound-imparipinnate; leaflets of unequal sizes......1. A. anisophyllus
Leaves simple, entire, lobed, faintly toothed towards apex, shallowly crenate, or deeply
pinnately incised down to the midrib with equal segments...........2
2. Leaves of adult and juvenile trees deeply, pinnately lobed or incised down to the
midrib...................
A. altilis (Parkinson) Fosberg
(Latin, altus = tall; the tree)
J. Acad. Sci. Wash. 31 (1941) 95. Synonyms: A. communis J.R. & G. Forst., Char. Gen. (1776) 101; A. incisus Linn. f.,
Suppl. Pl. (1781) 411.
Pulon, kukor, sukun, (breadfruit). Native of New Guinea and Micronesia; cultivated
throughout the tropics for the edible fruits.
Leaves of adult trees entire, lobed, faintly toothed towards apex, or shallowly crenate; those
of juvenile trees entire, or lobed, or deeply pinnately incised.......3
3. Lateral veins sunken above............4
Lateral veins raised or flat above...............8
4. Leaf margin lobed..............5
Leaf margin entire, plane or faintly toothed towards apex....6
5. Twig and petiole densely hispid-pubescent with tawny subpatent hairs 47 mm long. Leaves
1825 cm broad; petiole 12 cm long........20. A. sarawakensis
Twig and petiole sparsely pubescent with patent reddish brown hairs up to 5 mm long.
Leaves 1118 cm broad; petiole 3.54.5 cm long..........21. A. tamaran
6. Petiole up to 0.7 cm long. Leaves soft-hairy below................12. A. longifolius
Petiole at least 1 cm long. Leaves rough-hairy below...........7
7. Petiole 25.5 cm long. Lower leaf surface often glaucous; leaf margin plane. Syncarp
warty........18. A. primackiana
Petiole 13 cm long. Lower leaf surface not glaucous; leaf margin faintly toothed towards
apex. Syncarp densely covered with sharp rigid processes.........
.....................19. A. rigidus (in part)
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8. Leaves often glaucous below; midrib sunken above..........7. A. glaucus

Leaves not glaucous below; midrib raised or flat above............9
9. Leaf apex cuspidate or rounded...........10
Leaf apex acute, acuminate, or obtuse.................11
10. Leaf apex cuspidate; intercostal venation scalariform (ladder-like)..............2. A. annulatus
Leaf apex rounded, intercostal venation reticulate (net-like)..........15. A. obtusus
11. Leaves glabrous (to unaided eye) on both surfaces or sparsely pubescent only on the veins
below................12
Leaves sparsely or densely hairy all over at least on the lower surface..........18
12. Leaf margin lobed, undulate, or shallowly crenate..............13
Leaf margin entire and plane...............15
13. Leaves thick-coriaceous, drying grey-green..........11. A. lanceifolius
Leaves thin-coriaceous, drying reddish brown........14
14. Leaves scabrid (sandpapery to the touch) above; petiole 35 cm long; lateral veins 1215
pairs............................3. A. corneri
Leaves glabrous or nearly so above; petiole 13.5 cm long; lateral veins 612
pairs........22. A. teysmannii
15. Lateral veins and intercostal venation distinctly visible above...............14. A. nitidus (in part)
Lateral veins and intercostal venation invisible or very faintly visible above.................16
16. Lateral veins 1220 pairs..............................................6. A. excelsus
Lateral veins 913 pairs...............17
17. Syncarp c. 4.5 3.5 cm, borne on twigs behind the leaves, surface areolate
.................10. A. kemando (in part)
Syncarp 30100 cm long, borne on stem and older branches, surface short-spiny.....
A. heterophyllus Lamarck
(Greek, heteros = different, phullon = leaves; with different types of leaves)
Encycl. Meth. 3 (1789) 210.
Nangka, Jackfruit. Possibly indigenous to S India. Cultivated throughout the tropics for
the edible fruits.
18. Leaves sparsely hairy below....................................19
Leaves densely hairy below.................21
19. Hairs of midrib and lateral veins on lower leaf surface long and wavy. Inflorescences and
syncarps arising from trunk and older branches...............8. A. integer
Hairs of midrib and lateral veins on lower leaf surface short and appressed. Inflorescences
and syncarps axillary or on twigs behind the leaves....................................20
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20. Lateral veins and intercostal venation distinctly visible above...14. A. nitidus (in part)
Lateral veins and intercostal venation faintly visible above...............10. A. kemando (in part)
21. Leaves soft-hairy below...............22
Leaves rough-hairy below................24
22. Indumentum yellowish. Syncarp with peduncle up to 3 cm long........4. A. dadah
Indumentum reddish brown. Syncarp with peduncle (3)45(7) cm long........23
23. Twig up to 3 mm thick. Syncarp irregularly lobed........23. A. tomentosulus
Twig 38 mm thick. Syncarp smooth..........17. A. peltatus
24. Leaves 1035 cm broad...............25
Leaves usually less than 10 cm broad..............26
25. Petiole 410 cm long. Syncarp cylindrical, covered with fleshy hispid processes of different
sizes..........5. A. elasticus
Petiole 13 cm long. Syncarp subglobose, covered with rigid cylindrical processes of similar
size.................16. A. odoratissimus
26. Petiole 35 cm long................9. A. jarrettiae
Petiole up to 3 cm long....................27
27. Leaf margin faintly toothed towards apex. Syncarp densely covered with sharp, rigid
processes.................19. A. rigidus (in part)
Leaf margin entire. Syncarp almost smooth.....13. A. melinoxylus
1. Artocarpus anisophyllus Miq.

(Greek, aniso- = unequal, phullon = leaves; the unequally sized leaflets)
Fig. 2.
Fl. Ind. Bat., Suppl. (1861) 422; Merrill, PEB (1929) 45; Masamune, EPB (1942) 238; Brown l.c. 352; Jarrett l.c. (1959) 138;
Burgess l.c. 399; Kochummen l.c. 123; Anderson l.c. 254; Primack l.c. 26; Corner l.c. (1988) 515; Whitmore, Tantra & Sutisna
l.c. 236; Kessler & Sidiyasa l.c. 174; PROSEA l.c. (1995) 64; Turner, Gard. Bull. Sing. 47 (1995) 347; Coode et al. (eds.) l.c.
210; Argent et al. (eds.) l.c. 432. Type: Teijsmann HB 3698, Sumatra, Palembang, Batu Raja (holotype U; isotypes BO, K, L).
Synonyms: A. klidang Boerl., Handl. Fl. Ned. Ind. 3 (1900) 333; A. superba Becc. l.c. 625, Merrill l.c. (1921) 219, Ridley l.c.
(1924) 356.
Tree up to 45 m tall, 90 cm diameter; buttresses spreading, up to 2.5 m tall. Bark dark grey,
smooth to dippled; inner bark yellowish brown. Twigs stout, c. 1 cm thick, smooth or irregularly
ridged, set with ring-like stipular scars. Stipules amplexicaul, 417 cm long, appressed
pubescent. Leaves spirally arranged, compound-imparipinnate, rachis up to 90 cm long. Leaflets
oblong, ovate or lanceolate, 3.540 213 cm, subopposite, unequal in size, smaller ones
alternating with larger ones, terminal leaflets almost always bifid; midrib raised above; lateral
veins 720 pairs, sunken above, raised below; venation scalariform, faint to distinct below, faint
above. Inflorescences axillary, males and females usually paired; bracts slenderly stalked, apical
parts funnel-shaped; peduncle 56.5 cm long. Male heads ellipsoid-oblong, 37.5 1.52 cm;
perianth tubular, c. 1.8 mm long, shortly bilobed, minutely hairy; stamens 22.5 mm long,
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filament slender, cylindrical, anthers subglobose, c. 0.2 mm long. Syncarps subglobose, up to 11

8 cm, covered with cylindrical, rigid, minutely punctate, obtuse processes (perianth-apices) 6
8 11.5 mm; pulp (fleshy perianth) orange; peduncle 613 cm long. Seeds ellipsoid, c. 1.7 1
cm.
Vernacular names. Sabahterap ikal (Dusun, preferred name). Sarawakbintau (Bidayuh,
Sadong), bintawak (Bidayuh, Iban), denging (Kayan), entawa, mentawa (Iban), kayo bakek
(Kayan), kayo bibungan (Berawan), kelidang (Murut), tawak (Kenyah, Punan Tutoh).
Distribution. Sumatra, Peninsular Malaysia and Borneo. Widely distributed in Sabah,
uncommon in Sarawak. The typical variety also occurs in Brunei and Kalimantan.
Ecology. In lowland and hill mixed dipterocarp forests, up to 600 m altitude.
Uses. In Sarawak, the timber is traded under the name bintawak and locally used for indoor
constructions and for making household utensils and tool-handles. The fruits are edible.
Key to varieties
Twig ridged. Leaflets distinctly stalked..............var. anisophyllus
Twig smooth. Leaflets sessile................
var. sessilifolius Kochummen
(Latin, sessile = stalkless, folium = leaf)
Gard. Bull. Sing. 50 (1998) 200. Type: Wood SAN 16549, Borneo, Sabah, Sandakan district, Sepilok FR (holotype SAN;
isotypes A, BO, K, KEP, L, SING).
Endemic to Borneo. Widely distributed in Sabah, but in Sarawak known only by a single
collection, Hewitt 177, from Kuching district. Not known in Brunei and Kalimantan.
2. Artocarpus annulatus Jarrett

(Latin, annulatus = marked with rings; the ring-like constrictions on male inflorescences)
Blumea 22 (1975) 409; Anderson l.c. 254; Whitmore, Tantra & Sutisna l.c. 236; Primack l.c. 26. Type: Chew & Anderson CWL
665, Borneo, Sarawak, Kuching district, Mt. Mentawa (holotype K; isotypes L, SAR).
Tree up to 10 m tall, 20 cm diameter. Twigs brown, 35 mm thick, sparsely pubescent with

subrecurved, apically barbed hairs, or glabrous. Stipules amplexicaul, c. 4 cm long, sparsely
hairy. Leaves simple, spirally arranged, both surfaces glabrous or the midrib and veins hairy
above; broadly elliptic or obovate, 1030 516.5 cm, base cuneate, margin entire, plane, apex
cuspidate; midrib raised above; lateral veins 47 pairs, raised on both surfaces, distinctly
curving along the leaf margin; intercostal venation scalariform,
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distinct below, faint above; petiole 2.55 cm long. Inflorescences solitary, axillary or borne on
the stem; peduncle 1.52.5 cm long, hairy; interfloral bracts with stout stalks and peltate apical
parts. Male heads ovoid, c. 3.5 3.5 cm, surface wrinkled by ring-like constrictions; perianth
deeply 2(4)-lobed, c. 0.8 mm long; stamens one or two, filaments sometimes basally connate, c.
1 mm long, anthers ellipsoid, c. 0.1 mm long. Female heads with simple, exserted, filiform
styles. Syncarps ellipsoid, c. 8 6 cm, covered with fleshy, cylindrical, obtuse, short-hairy
processes intermixed with numerous intrafloral bracts; stalks of the bracts thread-like and the
apices funnel-shaped; peduncle up to 3 cm long.
Vernacular names. Sarawakbukoh (Iban), patat (Bidayuh).
Distribution. Endemic to Borneo; very rare, known only from Mt. Mentawa near Kuching in
Sarawak.
Ecology. In forest on limestone hills, at 150300 m altitude.
3. Artocarpus corneri Kochummen
Fig. 3.
(E.J.H. Corner, 19061996, formerly prominent Professor of Tropical Botany, University of Cambridge, U.K., who
undertook extensive studies of the Malesian Moraceae)
Gard. Bull. Sing. 50 (1998) 187. Type: Dayang Awa & Yii S. 46878, Borneo, Sarawak, Dulit Range, Belaga (holotype KEP;
isotypes CGE, K, L, SAN, SAR).
Tree up to 22 m tall, 50 cm diameter. Twigs 79 mm thick, dark brown, closely ridged, very
sparsely short-hairy, with distinct ring-like stipular scars. Stipules amplexicaul, lanceolate, up to
3 cm long, densely covered with long hairs. Leaves simple, spirally arranged, thin-coriaceous,
upper surface scabrid, lower surface glabrous to unaided eye but with short rough hairs on the
midrib and lateral veins, drying reddish brown to chocolate-brown; obovate or oblong, (12)17
28 (5.5)712 cm, base cuneate, margin faintly lobed, apex acuminate or acute; midrib raised
above; lateral veins 1215 pairs, raised below, flat above; intercostal venation scalariform,
faintly visible below, invisible above; petiole 35 cm long, sparsely pubescent with short hairs.
Inflorescences and flowers unknown. Syncarps (immature) axillary, solitary, cylindrical, 57
3.54 cm; peduncle 5.58 cm long; perianth densely hairy, styles distinctly bifid.
Distribution. Endemic to Borneo; known only by two collections, S. 38879 from the Maligan
Range, Lawas district and the type, from Belaga district, in Sarawak.
Ecology. In hill mixed dipterocarp forest, at 8001000 m altitude.
4. Artocarpus dadah Miq.

(after the vernacular name dadah from Lampong, Sumatra)
Fl. Ind. Bat., Suppl. (1861) 420; Corner, Gard. Bull. Str. Settl. 10 (1939) 282, l.c. (1988) 516; Browne l.c. 352; Jarrett l.c. (1960)
91; Kochummen l.c. 124; Anderson l.c. 254; Primack l.c. 28; Whitmore, Tantra & Sutisna l.c. 236; PROSEA l.c. 347; Coode et
al. eds.) l.c. 210; Argent et al. (eds.)l.c. 432. Type: Teijsmann HB 4391, Sumatra (holotype U; isotypes BO, K, L). Synonyms:
A. mollis Miq. l.c. (1861) 420; A. rufescens Miq. l.c. (1861) 420; A. tampang Miq. l.c. (1861) 421; Ficus tampang (Miq.) Miq. l.c.
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(1861) 425; Ficus inconstatissima Miq. l.c. (1861) 431; A. inconstantissima (Miq.) Miq., Ann. Mus. Bot. Lugd.-Bat. 3 (1867)
211; A. dadah Miq. Var. pubescens Miq. L.c. (1867) 213; A. lakoocha Roxb. var. malayana King in Hooker f., Fl. Brit. Ind. 5
(1888) 544; A. reniformis Becc. l.c. 631; A. dasyphylla auct., non Miq.: Merr PEB (1929) 46.
Deciduous tree up to 35 m tall, 60 cm diameter. Bark grey-brown, distanly fissured. Twigs 2.5-5
mm thick, densely pubescent with short yellowish to reddish brown hairs. Stipules nonamplexicaul. Leaves simple, distichous, upper surface glabrous except for the sparse short hairs
on the midrib and lateral veins, lower surface densely pubescent with yellowish to reddish brown
soft hairs; obovate-oblong, elliptic-oblong, or ovate-elliptic, 10-25 (-30) x 5-11 cm, base
rounded to broadly cuneate or shallowly cordate, margin entire, plane, apex obtuse or acute;
midrib raised above; lateral veins 12-20 pairs, looping near the leaf margin, raised below, flat
above; intercostals venation scalariform or reticulate, distinct below, faint above; petiole 0.5-2
cm long, hairy. Leaves of juvenile trees pinnately lobed. Inflorescences solitary, axillary, bracts
stoutly stalked, their apical parts narrowly peltate; peduncle 0.8-3 cm long, yellowish hairy.
Male heads globose, 0.8-1.5 cm across; perianth 2-3-lobed, c. 0.3 mm long, stamens c. 0.5 mm
long. Female heads with similar bracts; female flowers with exserted styles. Syncarps
subglobose or obovoid, 5-7 cm across, green with deep pink flesh, surface smooth; peduncle up
to 3 cm long.
Vernacular names. Sabah-beruni (Dusun, preferred name). Sarawak-dadah (Iban, Malay),
dadak, merubi, selangking (Iban).
Distribution. Burma, Thailand, Sumatra, Peninsular Malaysia, and Borneo. Common in Sabah;
rare in Sarawak, known by one collection each from the 1st and 7th Divisions. Also in Brunei
and Kalimantan.
Ecology. In lowland forest on hilly sites, up to 300 m altitude.
Uses. A. dadah is fairly important source of beruni and selangking timber, and the wood is used
for house poles and flooring. The latex is reported to have disinfectant property and is applied on
wounds. The fruit is edible but sour.
5. Artocarpus elasticus Reinw. ex Blume

(Latin, elasticus=elastic; the latex)
Bijdr. FI. Ned. Ind. (1825) 481; Merrill l.c. (1921) 218; Burkill, EPMP (1935) 251; Masamune l.c. 234; Jarret l.c. (1959) 346;
Backer & Bakhuizen f. l.c. 18 Browne l.c. 403; Kochummen l.c. 237; Ardeson l.c. 254; Primack l.c. 29; Corner l.c. (1988) 516;
Whitmore, Tantra & Sutisna l.c.237; PROSEA l.c. (1995) 65; Turner l.c. 347; Coode et al. (eds) l.c. 210: Argent et al. (eds) l.c
433. Type: Reinwardt s.n. ( =RHL Sheet No. 9081851027 ), Java (holotype L). Synonyms: A. blumei Trec., Ann. Sci. Nat. Bot.
3, 8 (1847) 111, Merrill l.c (1921) 218, Masamune l.c 238; A. kunstleri King in Hooker f., F1. Brit. Ind. 5 (1880) 540, Riddley l.c
(1924) 353; A. blumei Trec.var. kunstleri Boerl., Handl. F1. Ned. Ind. 3 (1900) 370; A. pubescens auct., non Willd.: Blume l.c
(1825) 481; A. sericarpus Jarrett l.c. (1959) 350, Anderson l.c. 256.
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Tree up to 45 m tall, 90 cm diameter, with prominent buttresses up to 3 m tall. Bark grey-brown,

smooth to slightly scaly; inner bark pale brown. Twigs 1020 mm thick, rough-hairy. Stipules
amplexicaul, 620 cm long, hispid-pubescent with yellowish or reddish brown, patent or
appressed hairs. Leaves simple, spirally arranged, upper surface sparsely rough-hairy, lower
surface densely rough-hairy; ovate or elliptic to oblong, 1560 1035 cm, base usually
rounded, rarely cuneate, margin entire, plane or shallowly crenate, apex acute; midrib raised
above; lateral veins 1214 pairs, prominently raised below, flat and distinct above; intercostal
venation scalariform or reticulate, prominent below, faint above; petiole 410 cm long. Leaves of
juvenile trees 23-times pinnatifid, up to 200 cm long. Inflorescences solitary, axillary; peduncle
47.5 cm long, hairy. Male heads cylindrical, 615 cm long, with distinct deep grooves; perianth
bilobed, c. 0.8 mm long, appressed pubescent, stamens c. 0.9 mm long, anthers oblong, c. 0.4 m
long. Female heads with simple or bifid, exserted styles. Syncarps cylindrical, up to 12 6 cm,
yellow-brown, covered by fleshy, whitish, short-hispid processes of two lengths, the longer
flexuous and slightly tapering, solid, 1018 11.5 mm, and the shorter ones narrowly conical,
perforate, c. 4 1 mm; bracts with funnel-shaped upper parts, scattered; peduncle 6.512 cm
long, hairy.
Vernacular names. Sabahterap togop (Dusun, preferred name). Sarawakkian (Berawan,
Kayan, Kenyah), pedalai, tekalong (Iban), pien (Bidayuh), terap (Malay, preferred name), tian
(Bidayuh, Sadong).
Distribution. Burma, Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, the Philippines,
Sulawesi, Maluku, and Lesser Sunda Islands. Common in Sabah and Sarawak. Also in Brunei
and Kalimantan.
Ecology. In primary and secondary lowland to lower montane forests, up to 1500 m altitude.
Uses. The fleshy fruiting perianth is edible, and the seeds are also eaten after roasting. The bark
is used by the aborigines in Sarawak as cordage for making ropes and cloths. The latex is used
for preparing bird-lime. The timber is known and sold as terap and is used mainly for light
construction.
6. Artocarpus excelsus Jarrett

Blumea 22 (1975) 409; Coode et al. (eds.) l.c. 210. Type: Chew & Corner RSNB 7046, Borneo, Sabah, Mt. Kinabalu, Mesilau R.
(holotype K; isotypes L, SING).
Tree up to 35 m tall, 50 cm diameter, buttresses sometimes present. Bark grey, scaly. Twigs c. 5
mm thick, dark brown, rugose. Stipules amplexicaul, appressed hairy, oblong, up to 5.5 cm long.
Leaves simple, spirally arranged, upper surface glabrous, lower surface appressed hairy on the
midrib and veins only; narrowly oblong or elliptic, 1022 26.5 cm, base cuneate, margin
entire, plane, apex acute; midrib black on drying, flat above; lateral veins 1220 pairs, flat and
faintly visible above, raised below; intercostal venation reticulate, visible below only; petiole
1.53 cm long. Juvenile leaves shallowly irregularly pinnatifid. Male heads cylindrical, 1.72.2
0.5 cm; perianth bilobed, c. 0.8 mm long, stamens c. 1 mm long, filaments slender, cylindrical,
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anthers ellipsoid, c. 0.2 mm long. Female heads with simple exserted styles. Syncarps
irregularly cylindrical, c. 7 4 cm, dull ochre-yellow, covered by fleshy, conical blunt processes
projecting up to c. 1 mm high and 1.5 mm broad; fruiting perianth numerous, proximal free parts
thinly fleshy, whitish; peduncle 12.5 cm long, appressed hairy.
Disribution. Endemic to Borneo. Recorded only for Mt. Kinabalu where it is common, and from
G. Pagon Periok in BruneiSarawak border.
Ecology. In lower montane forest, at 14001800 m altitude.
7. Artocarpus glaucus Blume

(Latin, glaucus = sea-green; the colour of the leaf undersurface)
Bijdr. Fl. Ned. Ind. (1825) 483; Browne l.c. 353; Jarrett l.c. (1960) 118; Backer & Bakhuizen f. l.c. 19; Kochummen l.c. 126;
Anderson l.c. 254; Primack l.c. 30; Whitmore, Tantra & Sutisna l.c. 237; PROSEA l.c. (1995) 66; Turner l.c. 347; Coode et al.
(eds.) l.c. 210; Argent et al. (eds.) l.c. 433. Type: Blume s.n. (= RHL Sheet No. 908185719), Java (holotype L). Synonyms: A.
glaucescens Trc., Ann. Sc. Nat. Bot. 3, 8 (1847) 120; A. zollingeriana Miq. in Zollinger, Syst. Verz. 2 (1854) 90 & 95; A.
biformis Miq., Fl. Ind. Bat., Suppl. (1861) 419; A. tephrophylla Miq. l.c. (1861) 422; A. glaucescens Trc. var. tephrophylla Miq.,
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 212; A. denisoniana King in Hooker f., Fl. Brit. Ind. 5 (1888) 544, Ridley l.c. (1924) 355.
Tree up to 40 m tall, 80 cm diameter, with or without buttresses. Bark reddish or greyish brown,
smooth or somewhat fissured, flaky; inner bark fibrous, yellowish; sap yellowish white. Twigs
slender, 1.54 mm thick, finely rugose, appressed hairy, early glabrescent. Stipules nonamplexicaul. Leaves simple, distichous, upper surface glabrous, lower surface glaucous and
appressed hairy; elliptic-oblong, obovate, or ovate-lanceolate, 620 38 cm, base asymmetric,
broadly cuneate, margin entire, plane, apex acute; midrib sunken above; lateral veins 815 pairs,
raised below, flat above; intercostal venation scalariform or reticulate, distinct below, flat above;
petiole 0.74.5 cm long. Inflorescences solitary or paired, axillary; bracts with peltate upper
parts; peduncle up to 0.3 cm long. Male heads oblong or club-shaped; perianth 23-lobed,
divided almost to the base. Female flowers with exserted styles. Syncarps knobbly, subglobose,
up to 3 cm across, yellow to orange-brown with light orange flesh, surface papillate or smooth,
short-pubescent; peduncle 0.20.5 cm long, velvety hairy.
Vernacular names. Sarawakmerubi (Malay Baram), pudau putih (preferred name),
selangking (Iban).
Distribution. Sumatra, Peninsular Malaysia, Java, Lesser Sunda Islands, and Borneo.
Uncommon in Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. In lowland to hill forests, up to 1000 m altitude.
Use. The timber is classified as terap, and is used for house-building.
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8. Artocarpus integer (Thunb.) Merr.

(Latin, integer = entire; the leaf margin)
Interpr. Rump. Herb. Amb. (1917) 190; Merrill l.c. (1921) 219 (integra); Ridley l.c. (1924) 354; Corner, Gard. Bull. Str. Settl. 10
(1939) 56, l.c. (1988) 518; Masamune l.c. 234 (integra); Jarrett l.c. (1959) 329; Backer & Bakhuizen f. l.c. 19; Burgess l.c. 399;
Kochummen l.c. 127; Anderson l.c. 255; Primack l.c. 32; Whitmore, Tantra & Sutisna l.c. 237; PROSEA 2 (1991) 91; Turner l.c.
347; Coode et al. (eds.) l.c. 211; Argent et al. (eds.) l.c. 434. Basionym: Radermachia integra Thunb., Vet. Akad. Handl.
Stockholm 37 (1776) 254. Type: Thunberg s.n., Java (holotype UPS; isotype L). Synonyms: A. jaca Lam., Encycl. 3 (1789) 209;
Polyphema champeden Lour., Fl. Cochinch. (1790) 546, p.p.; A. champeden (Lour.) Stokes, Bot. Mat. Med. 4 (1812) 330; A.
hirsutissima Kurz, Natuurk. Tijdschr. Ned. Ind. 27 (1864) 182.
Tree up to 24 m tall, 60 cm diameter; bole without buttresses, with bumps where syncarps are
produced. Bark dark grey-green, dippled to somewhat scaly; inner bark yellowish. Twigs 2.54
mm thick, smooth or shallowly rugose, sparsely to densely pubescent with patent, rufous hairs up
to 3 mm long. Stipules amplexicaul, ovate, acute, 1.59 cm long, hairy. Leaves simple, spirally
arranged, upper surface glabrous except for the sparsely hairy midrib and lateral veins, lower
surface often sparsely appressed pubescent with reddish brown, long, wavy hairs; elliptic to
oblong or obovate, 525 2.512 cm, base cuneate or rounded, margin entire, plane, apex
acute; midrib raised above; lateral veins 610 pairs, flat above, distinct below; intercostal
venation scalariform, faint; petiole 0.83 cm long, hairy. Inflorescences solitary, axillary or
borne on short leafy shoots on older branches or from the bumps on the main trunk; peduncle
2.56 cm long, hairy. Male heads cylindrical to club-shaped, 35.5 11.2 cm; perianth 0.71
mm long, shortly bilobed, stamens 11.3 mm long, anthers ellipsoid, c. 0.3 mm long. Female
flowers with simple thread-like styles exserted to c. 1.5 mm long. Syncarps arising from trunk
and older branches, cylindrical or oblong, 2035 1015 cm, yellow to yellowish green with
strong penetrating odour when ripe; surface covered by firm, obtuse, minutely hispid processes
24 3 mm, fleshy perianth surrounding fruit yellow, shinning, sweet; peduncle 5.56.5(12)
cm long, hairy.
Vernacular names. Sabahpulutan (Dusun, preferred name). Sarawakbeluno (Murut),
bukoh, temedak (Iban), chempedak (Malay, preferred name), nakan (Berawan, Kayan, Kenyah,
Punan), tibadak (Bidayuh, Sadong), tibudak (Bidayuh).
Distribution. Thailand, Peninsular Malaysia, Sumatra, Borneo, Sulawesi, Maluku, and New
Guinea. Common in Sabah and Sarawak. Also in Brunei and Kalimantan. Two varieties, viz. var.
integer and var. sylvestris, are recognised. Only the former occurs in Sabah and Sarawak.
Ecology. In primary and secondary lowland forests, up to 500 m altitude.
Uses. Planted extensively for the edible fruits. The young fruits are cooked as vegetable, while
the ripe fruits are consumed fresh or coated with flour and deep-fried. Seeds are eaten after
boiling or roasting.
9. Artocarpus jarrettiae Kochummen

(J. Francis Jarrett, former botanist at the Royal Botanic Gardens, Kew, U.K., who revised the
genus Artocarpus for the Malesian region)
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Gard. Bull. Sing. 50 (1998) 198. Type: Amin & Francis SAN 120933, Borneo, Sabah, Ranau district (holotype SAN; isotypes K,
KEP, L).
Tree up to 15 m tall, 30 cm diameter. Twigs 67 mm thick, dark brown, closely ridged. Stipules
amplexicaul, lanceolate, up to 5.5 cm long, with recurved edges, densely long-hairy outside.
Leaves simple, spirally arranged, upper surface sparsely rough-hairy, lower surface densely
rough-hairy; elliptic or oblong, 16.523 8.510(12) cm, base cuneate, margin entire, plane,
apex obtuse or acute; midrib raised above; lateral veins 1113 pairs, raised below, flat above;
intercostal venation scalariform, visible below, faint above; petiole 35 cm long, short-hairy.
Inflorescences not seen. Syncarps (immature) solitary, axillary, yellowish green when fresh,
cylindrical, c. 4.5 3 cm; perianth of female flowers covered with glandular hairs with swollen
bases, style exserted, simple; peduncle 4.55.5 cm long, rough-hairy.
Distribution. Endemic to Borneo; very rare, known only by one collection each from Sabah (the
type) and Sarawak, S. 54906, from Sarikei district.
Ecology. In lowland forest by streams.
10. Artocarpus kemando Miq.

(from the Sumatran vernacular namekemando)
Fl. Ind. Bat., Suppl. (1861) 418; Beccari l.c. 628; Merrill l.c. (1921) 219, l.c. (1929) 46; Ridley FMP 5 (1925) 335; Burkill l.c.
256; Corner l.c. (1939) 282, l.c. (1988) 519; Masamune l.c. 240; Jarrett l.c. (1959) 354; Kochummen l.c. 129; Anderson l.c. 255;
Primack l.c. 33; Whitmore, Tantra & Sutisna l.c. 237; PROSEA l.c. (1995) 67; Turner l.c. 347; Coode et al. (eds.) l.c. 211;
Argent et al. (eds.) l.c. 434. Type: Teijsmann HB 4515, Sumatra, Lampong, Kebong (holotype U; isotypes BO, L). Synonym: A.
brunneifolia S. Moore, J. Bot. 63 (1925) Suppl. 112.
Tree up to 35 m tall, 80 cm diameter, with spreading buttresses up to 2.5 m tall. Bark grey,
smooth; inner bark white to reddish. Twigs 22.5 mm thick, finely pubescent with appressed
reddish hairs. Stipules amplexicaul, lanceolate, acute, 0.72 cm long, appressed pubescent with
reddish hairs. Leaves simple, spirally arranged, upper surface glabrous, lower surface glabrous
or sparsely appressed short-hairy on the midrib and lateral veins; elliptic to oblong, 521 29
cm, base rounded or cuneate, margin entire, plane, apex acute or obtuse; midrib flat above;
lateral veins 913 pairs, looping near the leaf margin, distinct below, flat or faintly visible
above; intercostal venation scalariform or reticulate, visible below, faintly visible above; petiole
12 cm long. Inflorescences solitary or paired, axillary or on twigs behind the leaves; peduncle
0.73.5 cm long, pubescent. Male heads cylindrical, 24 cm long, wrinkled; perianth c. 0.4 mm
long, deeply bilobed; stamens c. 0.5 mm long, filaments cylindrical, anthers ellipsoid, c. 0.2 mm
long. Female flowers with exserted styles. Syncarps axillary or borne on twigs behind the
leaves, ellipsoid to subglobose, c. 4.5 3.5 cm, surface hairy, smooth or areolate, or covered by
contiguous, low, umbonate processes 23 mm across; peduncle 1.55.8 cm long, pubescent.
Vernacular names. Sabahpudu (Dusun, preferred name). Sarawakpudau (Malay, Iban),
pudu (Iban), pupud, selibut (Kayan), puduh (Bidayuh), puroh (Berawan).
Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo. Very common in Sabah
and Sarawak. Also in Brunei and Kalimantan.
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Ecology. In lowland and hill mixed dipterocarp forests, often in swampy habitats, up to 1000 m
altitude.
Uses. The timber is known as terap and is used for making household utensils and door/window
frames. The latex is used for preparing bird-lime, and the fruits are edible.
11. Artocarpus lanceifolius Roxb.

(Latin, lanci- = lance-shaped, folium = leaf; the lance-shaped leaves)
Fl. Ind., ed. Carey 3 (1832) 527; King in Hooker f., Fl. Br. Ind. 5 (1888) 543, Ann. Roy. Bot.Calc. 2 (1889) 3; Ridley l.c. (1924)
354; Foxworthy, Mal. For. Rec. 3 (1927) 128; Burkill l.c. 256; Jarrett l.c. (1959) 140; Kochummen l.c. 129; Anderson l.c. 255;
Primack l.c. 34; Corner l.c. (1988) 519; Whitmore, Tantra & Sutisna l.c. 237; PROSEA l.c. (1995) 67; Turner l.c. 347; Coode et
al. (eds.) l.c. 211; Argent et al. (eds.) l.c. 436. Type: Drawing by Wight (1843). Synonyms: A. clementis Merr., J. Str. Br. Roy.
As. Soc. 85 (1922) 164; A. lanceifolius Roxb. subsp. clementis (Merr.) Jarrett l.c. (1959) 142, syn. nov.
Tree up to 35 m tall, 90 cm diameter, occasionally with low buttresses. Bark dark grey, smooth
to scaly; inner bark yellowish white. Twigs 68 mm thick, rugose, appressed hairy, minutely
punctate. Stipules amplexicaul, ovate or lanceolate, acute, 1.54.5 cm long, appressed pubescent
with short hairs. Leaves simple, spirally arranged, thick-coriaceous, entire in adult trees, deeply
pinnately incised with equal segments in juvenile trees, glabrous on both surfaces, drying greygreen; elliptic to ovate, obovate or oblong, rarely lanceolate to elliptic, 1033 517 cm, base
cuneate, asymmetric, margin undulate, apex acute or obtuse; midrib raised above; lateral veins
714 pairs, raised on both surfaces; intercostal venation scalariform, very faint below, invisible
above; petiole 13 cm long. Inflorescences solitary or paired, axillary; bracts with funnel-shaped
upper parts; peduncle 2.57 cm, short-hairy. Male head ellipsoid to cylindrical, 36 1.21.8
cm. Female heads with peltate bracts; female flowers with exserted bifid styles. Syncarps
subglobose, 812 7 cm, olive- or chestnut-brown, tessellate by hardened truncate processes;
peduncle 510 cm long; fleshy perianth whitish.
Vernacular names. Sarawakkaliput, pala tupai (Iban).
Distribution. Sumatra, Peninsular Malaysia and Borneo. Widely distributed in Sabah; rare in
Sarawak, recorded only for Belaga (Dulit Ranges) and Kapit districts. Also in Brunei and
Kalimantan.
Ecology. In lowland and hill mixed dipterocarp forests, up to 300 m altitude, very rarely up to
1800 m.
Uses. The pulp surrounding the seeds is edible. An important source of keledang timber which is
used in heavy construction, furniture manufacture, boat-building, and for making household
utensils and coffins.
12. Artocarpus longifolius Becc.

(Latin, longus = long, folium = leaf; long-leaved)
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For. Born. (1902) 629; Merrill l.c. (1921) 219; Masamune l.c. 240; Jarrett (1960) l.c. 83; Anderson l.c. 255; Primack l.c. 35;
Whitmore, Tantra & Sutisna l.c. 237; Turner l.c. 347; Coode et al. (eds.) l.c. 211. Type: Teijsmann HB 11312, Borneo (holotype
FI).
Tree up to 12 m tall, 30 cm diameter. Bark reddish brown, smooth. Twigs 46 mm thick,

densely pubescent with reddish brown hairs. Stipules non-amplexicaul, lanceolate, 0.51.5 cm
long, persistent. Leaves simple, distichous, upper surface glabrous except for the hairy veins,
lower surface soft-hairy; oblanceolate, oblong or narrowly obovate, 1136.5 4.516 cm, base
attenuate, margin entire, plane, apex acuminate, acumen up to 2 cm long; midrib flat above;
lateral veins 1115 pairs, looping near the leaf margin, raised below, sunken above; intercostal
venation scalariform or reticulate, raised below, sunken above; petiole 0.30.7 cm long, hairy.
Inflorescences solitary, axillary; bracts slenderly stalked, with peltate upper parts; peduncle 1.2
3 cm long, soft-hairy. Male heads obovoid, 0.82 cm long; perianth comprises 3 free tepals,
stamens c. 0.5 mm long, filaments broad, constricted at the upper parts, anthers globose, c. 0.2
mm across. Female flowers with exserted styles. Syncarps ellipsoid, c. 2 cm diameter
(immature), surface almost smooth, soft-hairy; peduncle 36 cm long, soft-hairy.
Distribution. Endemic to Borneo; rare in Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. In lowland mixed dipterocarp and kerangas forests, up to 100 m altitude.
13. Artocarpus melinoxylus Gagnep.

(Greek, melinos = yellowish, xylon = wood; the yellowish wood)
Bull. Soc. Bot. France 73 (1926) 87; Jarrett l.c. (1959) 143; Anderson l.c. 355; Whitmore, Tantra & Sutisna l.c. 238; Coode et al.
(eds.) l.c. 211. Lectotype (Jarrett, 1959): Poilane 7079, Indo-China, Ba Na (holotype P; isotype A).
Distribution. Indo-China and Borneo.

Two subspecies, subsp. melinoxylus and subsp. brevipedunculatus are recognised, of which only
the latter is known in Sabah and Sarawak.
subsp. brevipedunculatus Jarrett
(Latin, brevi- = short, pedunculatus = furnished with stalk; the short-stalked inflorescences)
J. Arn. Arb. 40 (1959) 144; Anderson l.c. 255; Primack l.c. 35; Whitmore, Tantra & Sutisna l.c. 238; Coode et al. (eds.) l.c. 211.
Type: Wood SAN A 1733, Borneo, Sabah, Beaufort district (holotype A; isotypes BO, BRI, K, KEP, L, SING).
Tree up to 25 m tall, 40 cm diameter. Twigs (3)5(7) mm thick, rugose, hispid with appressed,
yellow hairs. Stipules amplexicaul, ovate, acute, 12.5 cm long, appresssed yellowish hairy.
Leaves simple, spirally arranged, upper surface glabrous or with scattered hairs on the veins,
lower surface densely rough short-hairy, especially on the veins; elliptic to narrowly obovate,
(7.5)1420 (3.5)610 cm, base cuneate, margin entire, plane, apex obtuse to acute; midrib
flat above; lateral veins 79 pairs, flat above; invenation reticulate, visible on both surfaces;
petiole 11.5 cm long. Inflorescences solitary, axillary; bracts stoutly stalked, upper parts
peltate; peduncle 820 cm long, hairy. Male heads subglobose, 1.52.5 cm across; perianth 23lobed, c. 0.8 mm long, stamens c. 1.5 mm long, anthers globose, c. 0.1 mm across. Female
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heads with peltate bracts; female flowers with exserted styles. Syncarps subglobose, c. 4 cm
across, surface almost smooth or covered by low, hardened, cylindrical, obtuse, hispid-hairy
processes c. 3 2 mm; peduncle 24 cm long, hairy. Seeds ellipsoid c. 1 0.6 cm.
Vernacular names. Sarawakpala munsoh, pala tupai, temponek (Iban).
Distribution. Endemic to Borneo; common in Sabah, less frequent in Sarawak. Also in Brunei.
14. Artocarpus nitidus Trc.

(Latin, nitidus = shinning; the leaf upper surface)
Ann. Sci. Nat. Bot. 3, 8 (1847) 119; Elmer l.c. 624; Jarrett l.c. (1960) 121; Kochummen l.c. 131; Anderson l.c. 255; Primack l.c.
37; Corner l.c. (1988) 520; Whitmore, Tantra & Sutisna l.c. 238; PROSEA l.c. (1995) 68; Turner l.c. 347; Coode et al. (eds.) l.c.
210; Argent et al. (eds.) l.c. 436. Lectotype (Jarrett, 1960): Cuming 1078, the Philippines, Luzon, Albay (holotype P; isotypes
BM, CGE, K, L, SING). Synonyms: A. lanceolata Trc. l.c. 121; A. humilis Becc. l.c. 629, Merrill l.c. (1921) 214, Masamune
l.c. 239; A. lamellosa auct., non Blanco: Merr., Publ. Govt. Lab. Manila 27 (1905) 80; A. sampor Gagnep., Bull. Soc. Bot. Fr. 73
(1926) 90; A. lingnanensis Merr., Lignan Sci. J. 7 (1931) 302, type: Balansa 4112, Tonkin (P); A. nitidus Trc. subsp.
lingnanensis (Merr.) Jarrett l.c. (1960) 124, syn. nov.; A. nitidus Trc. subsp. humilis (Becc.) Jarrett l.c. (1960) 126, Primack l.c.
38, Whitmore, Tantra & Sutisna l.c. 238; A. borneensis Merr., J. Str. Br. Roy. As. Soc. 85 (1922) 165, type: Ramos 1592, Borneo,
Sabah (A, BM, BO, K, L, P, SING); A. nitidus Trc. subsp. borneensis (Merr.) Jarrett l.c. (1960) 127, Primack l.c. 38, Whitmore,
Tantra & Sutisna l.c. 238, syn. nov.; A. gomeziana Wall. var. griffithii King in Hooker f., Fl. Brit. Ind. 5 (1888) 544, syntypes:
Griffith 4665, Kings Collectors 6651, 7533, Scortechini 683 (K, SING); A. griffithii (King) Merr., Pap. Mich. Acad. 1, 24 (1939)
64, A. nitidus Trc. subsp. griffithii (King) Jarrett l.c. (1960) 128, syn. nov.
Tree up to 35 m tall, 70 cm diameter, with buttresses up to 1 m tall. Bark reddish brown, smooth
to shallowly fissured. Twigs 14 mm thick, smooth or finely rugose, appressed hairy, early
glabrescent. Stipules non-amplexicaul. Leaves simple, distichous, thin to thick-coriaceous, upper
surface glabrous, shining, lower surface glabrous or sparsely hairy; elliptic, obovate or oblong,
3.523 1.59 cm, base cuneate or rounded, margin entire, plane, apex obtuse or acute; midrib
appressed short-hairy below, raised above; lateral veins 515 pairs, appressed short-hairy
below, raised and distinct on both surfaces; intercostal venation reticulate, including the areoles
distinctly visible on both surfaces; petiole 0.52.5 cm long. Inflorescences solitary or paired,
axillary; bracts with slender stalk, upper parts peltate, ciliate; peduncle up to 0.3 cm long. Male
heads oblong or club-shaped, 0.31.2 cm long; perianth with 24 free tepals or fused at the base,
0.50.7 mm long; stamens 0.81 mm long, filaments broad, tapering or contracted at the upper
parts, anthers ellipsoid, c. 0.2 mm long. Female heads with scattered bracts; femaleflowers with
exserted styles, up to 0.5 mm long. Syncarps axillary, subglobose, 1.56 cm across, surface
smooth or often irregularly swollen, hairy or glabrous; peduncle 0.152 cm long. Seeds
subglobose to ovoid, 0.81 0.70.8 cm.
Vernacular names. Sabahberuni (Dusun). Sarawakdadak, empaka, karon (Bidayuh),
ngidunuk (Bidayuh, Bau), selangkin (Iban), sinojoh (Bidayuh, Padawan), taburakin (Bidayuh,
Sadong).
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Distribution. A very variable species occurring in Assam, Burma, Thailand, Indo-China,

Southern China, Sumatra, Peninsular Malaysia, Borneo, and the Philippines. Very common in
Ecology. In lowland to lower montane forests, up to 1500 m altitude.
Uses. The timber is categorized as terap and is used in house-building and light contruction. The
fruit is edible but not very tasty.
Taxonomy. Jarrett (l.c. 121) recognised 4 subspecies, viz. subsp. borneensis, subsp. griffithii,
subsp. lingnanensis, and subsp. nitidus, based on the presence or absence of indumentum on the
syncarp and its colour. However, after going through all the collections from Sabah and
Sarawak, I have concluded that the differences are not consistent, and that there are no vegetative
characters which can be used to distinguish the subspecies. Therefore, A. nitidus is here treated
as a widely distributed, vegetatively variable species.
15. Artocarpus obtusus Jarrett

(Latin, obtusus = blunt; the blunt leaf apex)
Blumea 22 (1975) 410; Anderson l.c. 255; Primack l.c. 39; Whitmore, Tantra & Sutisna l.c. 238. Type: Anderson S. 15740,
Borneo, Sarawak, Semengoh Arboretum (holotype K; isotypes CGE, L, SAR).
Tree up to 35 m tall, 60 cm diameter. Twigs up to 5 mm thick, rugose, subglabrous. Stipules

amplexicaul, 14 cm long, appressed hairy. Leaves simple, spirally arranged, glabrous, rarely
appressed hairy below; elliptic or obovate, 823 48 cm, base broadly cuneate, margin entire,
plane, apex rounded; midrib raised above; lateral veins 46 pairs, flat above; intercostal
venation finely reticulate, faint below, almost invisible above; petiole 22.5 cm long. Male
heads ovoid to subconical, c. 2.5 1 cm, irregularly wrinkled with short narrow grooves;
peduncle 1.22.5 cm long; perianth c. 0.7 mm long, bilobed, finely hairy. Female flowers with
exserted, unequally bifid styles. Syncarps ellipsoid, c. 8 6 cm, covered by fleshy, hairy
processes of two lengths, the longer ones flexuous, c. 10 3 mm, and the shorter ones conical
and rigid, 24 2 mm; peduncle 37.5 cm long, hairy. Seeds reniform to subglobose, c. 1.5
1.2 cm.
Distribution. Endemic to Borneo; very rare, known only by 4 collections from Sarawak (Sibu,
Mukah, Bintulu and Kuching districts). A sterile collection, SAN 56056, from Beaufort district in
Sabah probably belongs here.
Ecology. In lowland mixed dipterocarp, peatswamp and freshwater swamp forests.
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16. Artocarpus odoratissimus Blanco

(Latin, odoratus = strongly sweet-smelling; the fragrant ripe fruits)
Fl. Filip. (1837) 671; Merrill, Spec. Blanc. (1918) 124, l.c. (1921) 219; Masamune l.c. 240; Jarrett l.c. (1959) 147; Anderson l.c.
255; Primack l.c. 40; Whitmore, Tantra & Sutisna l.c. 238; PROSEA 2 (1991) 94; Coode et al. (eds.) l.c. 211; Argent et al. (eds.)
l.c. 437. Neotype (Merrill, 1918): Merrill Spec. Blanc. 1019, the Philippines, Mindoro (BM, L). Synonym: A. tarap Becc. l.c.
626; A. mutabilis Becc. l.c. 627.
Tree up to 25 m tall, 50 cm diameter. Bark dark greyish brown, smooth; inner bark yellowish
brown. Twigs 415 mm thick, smooth or ridged, sparsely to densely pubescent with yellow to
reddish brown patent hairs. Stipules amplexicaul, ovate, acute, 18 cm long, hispid-pubescent
with yellow to reddish brown, appressed or patent hairs. Leaves simple, spirally arranged,
densely rough-hairy on both surfaces; broadly elliptic to obovate, 1650 1128 cm, base
cuneate, margin entire, plane or undulate, apex acuminate; midrib raised above; lateral veins 9
15 pairs, looping near the leaf margin, raised on both surfaces; intercostal venation scalariform,
raised below, faint above; petiole 13 cm long, hairy. Leaves of juvenile trees pinnately lobed.
Inflorescences solitary, axillary; bracts with peltate upper parts; peduncle 2.57 cm long, hairy.
Male heads ellipsoid to club-shaped, 49 2.53.5 cm; perianth tubular, c. 2 mm long, shortly
bilobed, minutely pubescent; stamens c. 2 mm long, filaments cylindrical, slender, anthers
subglobose, c. 0.2 mm long. Female heads with pubescent peltate bracts; female flowers with
exserted styles. Syncarps subglobose, to 16 13 cm, greenish yellow, drying pale reddish
brown, covered by closely set, rigid, cylindrical, hispid-pubescent processes 813 1 mm,
fleshy part of perianth white, juicy; peduncle 5.516 cm long, hairy. Seeds ellipsoid, c. 1.2 0.8
cm.
Vernacular names. Sabahtimadang (Dusun, preferred name). Sarawakkeiran (Kelabit),
pingan, tekalong (Iban), terap (Malay, preferred name).
Distribution. Endemic to Borneo. Common in Sabah and Sarawak. Also in Brunei and
Kalimantan. Introduced and commonly cultivated in the Philippines.
Ecology. In primary and secondary lowland and hill mixed dipterocarp forests on sandy clay
soils, up to 1000 m altitude.
Uses. Commonly cultivated in villages for the sweet edible aromatic pulp surrounding the seeds.
Young fruits are cooked and eaten as vegetable. Seeds may be eaten after roasting. In Sarawak,
the latex is used to treat inflammation from wounds.
17. Artocarpus peltatus Merr.

(Latin, peltatus = shield-shaped; the peltate leaves)
Fig. 4.
J. Str. Br. Roy. As. Soc. 85 (1922) 166; Masamune l.c. 240. Type: Villamil 168, Borneo, Sabah, Sandakan district (holotype
PNH; isotype BO).
Tree up to 35 m tall, 60 cm diameter. Bark reddish brown, scaly. Twigs 38 mm thick, ridged,
densely reddish brown hairy. Stipules non-amplexicaul. Leaves simple, distichous,
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upper surface glabrous except for sparse hairs on the midrib and veins, lower surface densely
reddish brown soft-hairy; obovate to elliptic or oblong, 1535 718 cm, base rounded,
asymmetric, sometimes subcordate, often narrowly peltate, margin entire, plane, apex acute;
midrib flat above; lateral veins 1119 pairs, looping and joining near the leaf margin, raised
below, flat above; intercostal venation scalariform or reticulate, distinct below, faint above;
petiole 1.33 cm long, hairy. Inflorescences solitary, axillary; bracts with peltate apical parts, up
to 0.4 mm across; peduncle 3.55 cm long, reddish brown hairy. Male heads obovoid, 1.53.5
12.5 cm; perianth comprises 2 free tepals c. 0.4 mm long; stamens c. 0.8 mm long. Female
flowers with exserted styles. Syncarps obovoid or subglobose, up to 7 cm diameter, yellowgreen, surface smooth, pubescent; peduncle 47.5 cm long, reddish brown hairy. Seeds ellipsoid,
c. 1 0.8 cm.
Distribution. Endemic to Borneo. Common in Sarawak; very rare in Sabah, known by single
collection, SAN 76283, from Kota Belud district. Not known in Brunei and Kalimantan.
Ecology. In lowland forest, at about 200 m altitude.
18. Artocarpus primackiana Kochummen

(Richard B. Primack, Boston University, U.S.A., who wrote the Foresters Guide to the
Moraceae of Sarawak)
Gard. Bull. Sing. 50 (1998) 199. Type: Aban SAN 99596, Borneo, Sabah, Kota Merudu district (holotype SAN; isotype KEP).
Small to medium-sized tree, rarely up to 33 m tall, 50 cm diameter; buttresses up to 3.3 m high.

Bark grey-brown or orange-brown, smooth to cracking. Twigs 57 mm thick, covered with short
rough hairs. Stipules non-amplexicaul, ovate, c. 0.4 cm long, hairy outside. Leaves simple,
distichous, upper surface glabrous except the midrib, lower surface rough-hairy, often glaucous;
oblong or obovate, 1233 7.519 cm, base cuneate or rounded, margin entire, plane, apex
acuminate, acumen up to 1 cm long; midrib flat above; lateral veins 1016 pairs, prominently
raised below, sunken above; intercostal venation scalariform, raised below, faint above; petiole
25.5 cm long, short-hairy, often glaucous. Inflorescences solitary, axillary; bracts numerous
with peltate upper parts. Male heads globose, c. 1.8 cm diameter, sessile or with c. 0.5 cm long
hairy peduncle; perianth 3-lobed, covered with short hairs; stamen one, exserted with stout
filament. Female flowers with exserted, 3-lobed styles. Syncarps sessile or short-peduncled,
subglobose, c. 6 cm across, surface warty, velvety hairy, pale yellow with pink flesh when ripe.
Vernacular names. Sabahberuni (Dusun). Sarawakdadah (Iban).
Distribution. Endemic to Sabah and Sarawak. Most collections are from Sabah, where it is
widely distributed. In Sarawak, only known by one collection, S. 43409, from the G. Mulu NP,
Miri district.
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Notes. Differs from A. tomentosulus in the rough-hairy large leaves with sunken lateral veins,
small ovate stipules and in the sessile or shortly stalked syncarp. Jarrett (l.c. 1960) cited a
specimen, Wood SAN A 3684, along with the type from Sabah. In view of the large leaves and
glabrous nature of the intercostal venation, she suggested that the specimen was taken from a
juvenile shoot. However, this specimen was collected from a 13 m tall tree.
19. Artocarpus rigidus Blume

(Latin, rigidus = rigid; processes of the syncarp)
Fig. 5.
Bijdr. Fl. Ned. Ind. (1825) 482; King in Hooker f. l.c. (1888) 540, l.c. (1889) 8; Merrill l.c. (1921) 220; Ridley l.c. 352;
Masamune l.c. 240; Jarrett l.c. (1959) 150; Backer & Bakhuizen f. l.c. 18; Browne l.c. 353; Kochummen l.c. 131; Anderson l.c.
256; Primack l.c. 41; Corner l.c. (1988) 521; Whitmore, Tantra & Sutisna l.c. 238; Kessler & Sidiyasa l.c. 175; Turner l.c. 348;
PROSEA l.c. (1995) 69; Argent et al. (eds.) l.c. 437. Type: Blume s.n. (= RHL Sheet No. 908185863), Java (holotype L).
Synonyms: Radermachia rotunda Houtt., Nat. Hist. Pl. 11 (1779) 455; A. rotunda (Houtt.) Panz. in Panzer & Christmann,
Planzensyst. 10 (1783) 380, Backer & Bakhuizen f. l.c. 18; A. cuspidatus Griff., Not. Pl. Asiat. 4 (1854) 400; A. kertau Zollinger
ex Miq. in Zollinger, Syst. Verz. 2 (1854) 89, 95; A. dimorphophylla Miq. l.c. (1861) 417; A. varians Miq. l.c. (1861) 417.
Tree up to 45 m tall, 90 cm diameter, with buttresses up to 3 m tall. Bark dark grey, smooth,
fissured to scaly. Twigs 26 mm thick, smooth or rugose, densely or sparsely hispid-pubescent
with reddish brown hairs. Stipules amplexicaul, ovate or lanceolate, acute, 0.53 cm long,
hispid-pubescent with yellow to reddish brown hairs. Leaves simple, spirally arranged, upper
surface sparsely hairy, lower surface densely rough-hairy; elliptic to ovate, obovate or oblong,
932 410 (15) cm, base cuneate to rounded, margin faintly toothed towards apex, apex
acute; midrib raised above; lateral veins 920 pairs, raised below, flat or sunken above;
intercostal venation scalariform or reticulate, raised below, very faint above; petiole 13 cm long.
Leaves of juvenile trees pinnately lobed. Inflorescences solitary, axillary; bracts with peltate
upper parts c. 0.3 mm across, shortly ciliate; peduncle 0.20.6 cm long, densely appressed hairy.
Male heads obovoid to subglobose, 1.32 cm across; perianth tubular, c. 1 mm long, shortly
bilobed, minutely hairy; filaments slender, cylindric, anthers globose, c. 0.1 mm long. Female
heads with caducous, pubescent, peltate bracts; female flowers with simple exserted styles.
Syncarps globose, up to 13 cm diameter, dull orange, surface strongly spiny by the closely set,
rigid, tapering, acute, hispid-pubescent processes 79 2 mm; pulp around the fruit orange,
fleshy, sweet; peduncle up to 4 cm long.
Vernacular names. Sarawakpala munsoh, pudau, temponek (Iban).
Distribution. S Burma, S Thailand, Sumatra, Peninsular Malaysia, Borneo, Java, Kangean
Archipelago, and Bali. Common and widely distributed in Sabah and Sarawak. Also in
Kalimantan.
Ecology. In peatswamp, freshwater swamp, limestone hill, lowland and hill mixed dipterocarp
forests on diverse soils, up to 1000 m altitude.
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Uses. Cultivated for the edible sweet pulp surrounding the seeds. The timber is classified as
beruni or selangking and is used for making beams, furniture and boats.
20. Artocarpus sarawakensis Jarrett

(of Sarawak)
Blumea 22 (1975) 410; Primack l.c. 42. Type: Ilias S. 15109, Borneo, Sarawak, Bintulu district, Segar FR (holotype K; isotypes
C, L, SAN, SAR).
Tree up to 25 m tall. Twigs up to 10 mm thick, rugose, hispid-pubescent with tawny subpatent

hairs 47 mm long. Stipules amplexicaul, up to 10 cm long, densely hispid-pubescent. Leaves
simple, spirally arranged, upper surface glabrous except for scattered long hairs along the lateral
veins, lower surface pubescent with short, rough, appressed hairs; elliptic, 3050 1825 cm,
base cuneate, margin lobed, apex cuspidate; midrib flat above; lateral veins 1217 pairs, raised
below, sunken above; intercostal venation scalariform or reticulate, visible below, very faint
above; petiole 12 cm long, hispid-pubescent. Male heads obovoid to ellipsoid, 36 1.52.5
cm; peduncle 48 cm long, scabrid with scattered long hairs; bracts slenderly stalked, upper parts
peltate; perianth c. 0.8 mm long, stamens c. 1 mm long, anthers globose, c. 0.1 mm long. Female
flowers with simple exserted styles c. 2 mm long. Syncarps subglobose, c. 5 cm across, surface
covered with hardened, cylindrical, obtuse hairy processes c. 5 2 mm; peduncle up to 5.5 cm
long, pubescent with scattered long hairs. Seeds ellipsoid, c. 1.3 0.8 cm.
Vernacular names. Sarawakpedalai, pingan (Iban).
Distribution. Endemic to Borneo. Uncommon in Sarawak, not yet reported for Sabah. Not
known in Brunei and Kalimantan.
Ecology. In lowland mixed dipterocarp forest on ridges, up to 300 m altitude.
21. Artocarpus tamaran Becc.

(from the Iban name of the treetamaran)
For. Born. (1902) 626; Merrill l.c. (1921) 220; Masamune l.c. 240; Jarrett l.c. (1959) 352; Anderson l.c. 256; Primack l.c. 42;
Whitmore, Tantra & Sutisna l.c. 239; PROSEA l.c. (1995) 71; Coode et al. (eds.) l.c. 211; Argent et al. (eds.) l.c. 437. Type:
Beccari PB 2996, Borneo, Sarawak (holotype FI; isotype L).
Tree up to 40 m tall, 100 cm diameter; buttresses up to 3 m high. Bark grey-brown, smooth.

Twigs 510 mm thick, rugose, sparsely pubescent with scattered, patent, reddish brown hairs up
to 5 mm long. Stipules amplexicaul, lanceolate, acute, 318 cm long, covered with reddish
brown, subappressed hairs. Leaves simple, spirally arranged, upper surface glabrous, lower
surface appressed hairy; ovate or elliptic, 2035 1118 cm, base rounded, margin lobed, apex
acute; midrib raised above; lateral veins 1523 pairs, raised below, sunken above; intercostal
venation scalariform, visible below, faint above; petiole 3.54.5 cm long, reddish brown hairy.
Leaves of juvenile trees up to 100 cmlong, deeply pinnately lobed. Inflorescences solitary,
axillary. Male heads cylindrical, 7 11.5 cm, surface tuberculate with many irregular,
cylindrical, obtuse, solid processes c. 3 2 mm; peduncle 3.55.5 cm long, hairy; perianth
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tubular, c. 0.6 mm long, bilobed, minutely pubescent, stamen filaments flat, abruptly contracted
on the upper portions, anthers oblong, c. 0.2 mm long. Female flowers with exserted styles c.
0.5 mm long. Syncarps cylindrical to subglobose, c. 10 5 cm, dark brown, covered by closely
set, fleshy processes of two lengths; peduncle 5.513 cm long, hairy. Seeds ellipsoid, c. 0.6 0.4
cm.
Vernacular names. Sabahtimbangan (Dusun, preferred name). Sarawakkakang, tamaran,
tembaran (Iban), kalong, kihan (Kayan).
Distribution. Endemic to Borneo. Widely distributed in Sabah; less common in Sarawak and
recorded from Bintulu, Kapit and Lundu districts. Also in Brunei and Kalimantan.
Uses. Cloth is made from the bark. The timber is traded and used as terap.
22. Artocarpus teysmannii Miq.

(J.E. Teijsmann, 18021882, former curator of the Bogor Botanic Gardens)
Fl. Ind. Bat., Suppl. (1861) 418; Jarrett l.c. (1959) 341; Anderson l.c. 256; Primack l.c. 43; Whitmore, Tantra & Sutisna l.c. 239;
PROSEA l.c. (1995) 71; Turner l.c. 348; Argent et al. (eds.) l.c. 438. Type: Teijsmann HB 4387, Sumatra, Lampong, Kebong
(holotype U; isotypes BO, L). Synonym: A. peduncularis Kurz, J. Bot. 13 (1875) 331, Beccari l.c. 632, Merrill l.c. (1921) 219,
Ridley l.c. (1924) 353, Burkill l.c. 258, Corner l.c. (1939) 283, Masamune l.c. 240.
Tree up to 45 m tall, 80 cm diameter; buttresses up to 2 m tall. Bark dark brown, smooth. Twigs
35 mm thick, rugose, appressed hairy. Stipules amplexicaul, lanceolate, acute, 29 cm long,
appressed hairy. Leaves simple, spirally arranged, thin-coriaceous, upper surface glabrous or
nearly so, lower surface sparsely appressed pubescent on the veins, drying reddish brown; ovate
to elliptic, 525 313, base broadly cuneate, margin lobed or shallowly crenate, apex obtuse or
acute; midrib flat above; lateral veins 612 pairs, raised below, flat above; intercostal venation
scalariform, invisible on both surfaces; petiole 13.5 cm long. Inflorescences solitary, axillary.
Male head narrowly cylindrical, 3.57.5 0.51 cm; peduncle 28 cm long, sparsely appressed
hairy; sterile perianth solid, filiform, projecting up to c. 1 mm long from the surface, that of
fertile male flowers tubular, c. 1.4 mm long, bilobed, minutely pubescent; stamens up to 2 mm
long, filaments broad, abruptly contracted at the upper parts, anthers oblong, c. 0.5 mm long.
Female flowers with simple projecting styles up to 1.5 mm long. Syncarps cylindrical, up to 8.5
2.2 cm, yellow, covered by closely set, fleshy, conical, appressed hairy processes c. 1.5 1.5
mm; peduncle 310 cm long, appressed hairy. Seeds ellipsoid, c. 0.7 0.6 cm.
Vernacular name. Sarawakchempedak ayer (Malay).
Distribution. Nicobar Islands, Sumatra, Peninsular Malaysia, Borneo, Sulawesi, Maluku, and W
New Guinea. Very rare in Sabah and Sarawak, known by two collections, SAN 23440 (from
Kinabatangan district) and SAN 40656 (from Sandakan district) from Sabah, and by a few
collections from Marudi district, in Sarawak. Also in Kalimantan.
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Ecology. In lowland forest, including alluvial swampy habitats.

Uses. The timber is classified as terap, and the wood is used for boat-building.
23. Artocarpus tomentosulus Jarrett

(Latin, tomentosulus = covered with minute, soft, short, matted hairs; the indumentum of the
plant parts)
J. Arn. Arb. 41 (1960) 117; Primack l.c. 44; Whitmore, Tantra & Sutisna l.c. 239. Type: Keith 7671, Borneo, Sabah, Sandakan
district, Kabili-Sepilok FR (holotype SING).
Tree up to 15 m tall, 30 cm diameter. Twigs slender, c. 3 mm thick, rugose, pubescent with

reddish brown short hairs. Leaves simple, distichous, upper surface glabrous except for the
midrib and lateral veins, lower surface densely reddish brown soft-hairy; elliptic or oblong, 14
20 8.511 cm, base almost rounded, margin entire, plane, apex acuminate, acumen up to 1 cm
long; midrib flat above; lateral veins 914 pairs, raised below, flat above; intercostal venation
scalariform, raised below, invisible above; petiole 11.8 cm long, reddish brown hairy. Male
heads (not fully developed) globose, c. 0.7 cm across, solitary, axillary; bracts minute, stipitate,
apex peltate, c. 0.4 mm across; peduncle c. 0.2 cm long; perianth tubular, c. 0.6 mm long,
bilobed, minutely hairy; anthers ellipsoid, c. 0.2 mm long. Female heads not seen. Syncarps
subglobose, irregularly lobed, c. 4 cm across, reddish brown velvety hairy, styles simple, slightly
projecting; perianth with glandular, broad-based scales; peduncle 35 cm long.
Distribution. Endemic to Borneo. Only known from Sabah by 3 collections, Keith 7671 from
Sandakan district, and SAN 96814 and SAN 116315 from Ranau district.
Notes. Jarrett (l.c.) had wrongly identified a number of collections from Sabah as belonging to A.
tomentosulus. They do not match with the type. I have described a new species, A. primackiana,
based on these collections.
Incompletely known species

Artocarpus sp. A
Tree up to 24 m tall, 40 cm diameter. Bark dark brown, inner bark pale brown. Female heads in
clusters (immature). Known by one collection, SAN 82081, from Sepilok Arboretum, Sandakan
district, Sabah.
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3. FICUS L.
(Latin, ficus = the fig tree)
ara, kayu ara (Malay)

K.M. Kochummen
Sp. Pl. (1753) 1059, Gen. Pl. ed. 5 (1754) 482; Blume, Bijdr. Fl. Ned. Ind. (1825); Miquel, Ann. Mus. Bot. Lugd.-Bat. 3 (1867)
214; King, Ann. Roy. Bot. Gard. Calc. 1, 1 & 2 (18871888) 1; Merrill, EB (1921) 220; Ridley, FMP 3 (1924) 325, ibid. 5
(1925) 333; Corner, J. Mal. Br. Roy. As. Soc. 11 (1933) 1; Elmer, Leafl. Philip. Bot. 9 (1937) 3427; Corner, Gard. Bull. Str.
Settl. 10 (1939) 82, Gard. Bull. Sing. 17 (1960) 368, ibid. 18 (1960) 1, ibid. 19 (1962) 187, ibid. 21 (1965) 1; Backer &
Bakhuizen f., FJ 2 (1965) 20; Kochummen, TFM 3 (1978) 135; Anderson, CLTS (1980) 256; Wong, DMT (1982) 61; Corner,
WSTM 3rd. edition 2 (1988) 521; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 239; Berg, Mem. NY. Bot. Gard. 55 (1990)
169; Ng, Mal. For. Rec. 34, 2 (1991) 453; Rohwer in Kubitzki et al. (eds.), Fam. & Gen. Vasc. Pl. 2 (1993) 452; Coode et al.
(eds.), CLBD (1996) 211; Argent et al. (eds.), MNDT-CK 2 (1997) 438. Synonyms: Covellia Gasp. in Parlatore, Giorn. Bot. 2
(1844) 217; Sycomorphe Miq., Ann. Sc. Nat. 3, 1 (1844) 35; Pharmacosycea Miq. in Hooker, Lond. J. Bot. 6 (1847) 525;
Pogonotrophe Miq. l.c. (1847) 525; Synoecia Miq. in Hooker, Lond. J. Bot. 7 (1848) 469; Bosscheria Teijsm. & de Vries, Nat.
Tijds. Ned. Ind. 23 (1861) 212; Dammaropsis Warb., Bot. Jahrb. 13 (1891) 296.
Monoecious or dioecious trees, shrubs, woody epiphytes, climbers, stranglers, or creepers; all
parts with white or cream latex. Bark generally greyish, smooth. Twigs often with ring-like
stipular scars. Stipules mostly amplexicaul, free or sometimes connate, caducous or persistent.
Leaves simple, alternate (distichous or in spiral) or opposite; margin entire, toothed or lobed,
plane or revolute; basal lateral veins often distinct by departing from the midrib in a more acute
angle and/or distinctly longer than the other lateral veins. Inflorescences/infructescences
(syconia or figs) with urn-shaped and fleshy receptacles, enclosing the flowers or fruits, solitary,
in pairs or clustered, arising from the leaf axils, or borne on the twigs behind the leaves down to
older branches or stem, or on long runners/stolons arising from the base of stem and trailing
along the ground (in geocarpic figs), sessile or peduncled; base of the receptacle sometimes
tapered and stalk-like; basal bracts usually 3, lateral bracts sometimes present, interfloral bracts
and/or bristles present or absent; apex convex, flat and disc-like or depressed, ostiole closed by
overlapping bracts; surface smooth, warty, or wrinkled on drying. Flowers small, unisexual,
numerous, usually of three kinds, male, female and gall flowers, all three in the same syconium
(subgen. Urostigma, Pharmacosycea, Sycomorus), or male and gall flowers in syconia on one
plant and female flowers in syconia of another plant (subgen. Ficus), pedicelled or sessile; tepals
35(7), whitish or reddish, free or more or less united. Male flowers: stamens 15, usually with
distinct, free or united filaments, rarely anthers sessile; rudimentary ovary mostly absent. Gall
flowers resembling female flowers but with swollen and baloon-like ovaries, shorter style, and
not setting seed. Female flowers with a single ovary; style relatively long, terminal, subterminal
or lateral, stigma entire or divided; ovule one, pendulous. Fruits achene-like or rarely
drupaceous, tiny, with smooth or tuberculate surface.
Distribution. A large genus with about 800 species, mostly in tropical Africa, Asia, Malesia,
Australia, C and S America; very common in the Malesian region. A total of 143 species are
recorded for Sabah and Sarawak.
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Ecology. Occur in a wide variety of habitats from sea level up to 3200 m altitude, including
swamp, limestone, riverine, primary and secondary mixed dipterocarp and montane forests.
Epiphytic species infest old trees and crevices of walls of buildings and roof tops, and by their
long tough roots, even hold together crumbling rocks, limestone outcrops and other rockformations. Flowers are pollinated by fig-wasps which are species-specific. The fruits and seeds
are dispersed by frugivorous arboreal and terrestrial vertebrates, such as birds, bats, squirrels,
primates and wild boars, which consume the brightly coloured or dull-green, fleshy fruitingsyconia, and defecate the hard-coated, tiny seeds at a new place away from the parent trees.
Many strangling figs begin their life as epiphytes. Birds, squirrels and monkeys which eat the
figs drop the seeds on branches of trees where they germinate and grow into epiphytic bushes.
The roots of these new generation of fig plants encircle the branches of the host trees and
eventually the whole trunk, forming an anastomosing and interlacing basket. Some aerial roots
grow straight down to the ground from various heights. As the fig roots and the supporting trunk
of the host tree increase in thickness, they press upon each other, but the fig roots being the
stronger, slowly crush the bark of the host tree with the effect that the supporting tree is
gradually killed and the fig plant finally stands in its place on a massive basket of roots. Some
Ficus species are deciduous, shedding their leaves and remaining bare for a few days, usually
after pronounced dry weather. Ficus variegata sheds its leaves twice a year. Due to their year
round fruiting and large crops, fig plants provide resource for a variety of vertebrate frugivores
(fruit eaters) and consequently they are regarded as keystone mutualists in tropical forests. (P.
Stott (ed.), Fig trees and their associated animals, J. Biogeogr. 23, 4 (1996) 405607).
Uses. Ara or kayu ara is the common Malaysian name for the timber of Ficus species. The
timber is soft and light with density ranging from 350640 kg/m air dry. The sapwood is not
differentiated from the heartwood which is light yellow-brown or in various shades of yellow to
pink-grey. Texture is coarse and uneven, with the grain interlocked. The timber is easy to
moderately easy to resaw and easy to cross-cut. Planed surface is smooth and seasoned rapidly. It
is prone to sapstain fungal attack. The wood is suitable for low grade plywood, and for making
disposable chopsticks, packing cases and wooden sandals. The latex is used in the preparation of
bird-lime. Many Ficus species are cultivated as ornamental plants. The most common ones are F.
benjamina and a variety of F. microcarpa. Young plants of F. elastica are used as indoor pot
plants in office buildings because of the shiny dark green leaves and the bright red long stipules.
F. pumila, a creeper originally from China and Japan, is cultivated to grow over walls of
buildings. Many species of Ficus in Sabah and Sarawak are suitable for urban landscape
plantings. For example, F. aurantiacea and F. punctata will provide the same, if not better and
more colourful appearance than F. pumila. Other promising species include F. deltoidea and its
varieties, F. oleifolia and its varieties, and F. ixoroides. F. deltoidea is a well known plant in
traditional medicine in Peninsular Malaysia, Borneo and Indonesia. Decoction of leaves is taken
after child birth. Young leaves of F. grossularioides are cooked and eaten as vegetable. Rough
leaves are sometimes used as sandpaper.
Taxonomy. The genus Ficus is divided into 4 subgenera, viz. subgen. Ficus, subgen.
Pharmacosycea, subgen. Sycomorus and subgen. Urostigma. In Sabah and Sarawak, subgen.
Sycomorus is represented by only a single species, F. racemosa. The species in the other
subgenera are again grouped into series, and subseries, based mainly on floral cIn subgen. Ficus,
the figs (= syconia) containing male and gall flowers are found in different trees from the figs
containing female or seed flowers.
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Key to subgenera
1. Plants dioecious; syconia contain either male and short-styled (gall-)flowers or long-styled
(seed-)flowers only, the two types of syconia are borne on different
plants..............subgen. Ficus
Plants monoecious; syconia contain male, short-style (gall-), and long-styled (seed-)
flowers............2
2. Plants with aerial adventitious roots, often stranglers. Leaves mostly with a single gland at
the base of the midrib beneath. Male flowers with a single stamen. Style with one stigmatic
arm........subgen. Urostigma
Plants without aerial adventitious roots, always terrestrial. Leaves often with two glands, one
in each of the axils of the basal lateral veins. Male flowers with more than one stamen. Style
with 2 stigmatic arms or truncate to elongated stigma......3
3. Ovary white or reddish at the stylar side; style with 2 stigmatic arms; interfloral bracts or
bristles often present....subgen. Pharmacosycea
Ovary dark red; style and stigma simple; interfloral bracts or bristles absent.
.........subgen. Sycomorus
Notes. In the key to Ficus species given below, several categories of growth form are used.
However, in several groups of Ficus the species often develop different growth forms. In the
monoecious subgen. Urostigma, which can often be recognized by the presence of a single gland
(glandular spot) at the base of the midrib beneath, the species produce aerial adventitious roots
that enable them to become stranglers. However, young or small plants can be confused with
epiphytes. Several species in this group are often terrestrial, and then becoming lianescent or, in
open places, developing into trees with short stems and broad crown, and often with aerial roots
hanging from the branches which frequently developing into pillar-roots supporting the branches.
Within the dioecious subgen. Ficus there is a subsection (Palaeomorphe), which can often be
recognized by the not fully amplexicaul stipules and the arrangement of the basal bracts, i.e.
often not neatly arranged in a whorl, but occurring more or less scattered on the peduncle. The
species of this group also produce aerial adventious roots, and many of them exhibit different
habits. They can be terrestrial trees or shrubs or (straggling) climbers, which may become
(partly) epiphytic. Some are stranglers anchored close to the forest floor and others can be
confused with stranglers. A third group comprising some members of the dioecious subgen.
Ficus are also capable of producing aerial adventious roots. These species are rather uniform in
being root-climbers with short adventious roots arising from the stems. The leaves of the sterile
climbing stems are more or less clearly differentiated from those of the spreading fertile branches
which usually do not produce adventitious roots. Finally, a small group of the dioecious subgen.
Ficus comprising F. deltoidea and F. oleifolia can be terrestrial, epiphytic or epilithic.
The users of the key are, therefore, adviced to try more than one possibility as with regard to
habit (at leads 1 and 116) in order to key out material collected from plants with somewhat
unusual habits or growth forms.
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Key to Ficus species

1. Trees or erect shrubs (including some stranglers that develop stems).......................2
Stranglers, climbers, creepers or epiphytes, rarely scandent shrubs................80
2. Leaves glabrous and smooth..................3
Leaves hairy or scabrous, at least on the midrib below...........38
3. Leaf margin entire or rarely lobed (in some varieties of F. deltoidea)......4
Leaf margin toothed or lobed...........32
4. Midrib sunken above..................5
Midrib raised or flat above...........11
5. Leaves thick-coriaceous, cardboard-like, margin revolute................70. F. xylophylla (in part)
Leaves membranous to coriaceous, margin plane or revolute...........................6
6. Lateral veins sunken above.50. F. retusa
Lateral veins flat or raised above...........7
7. Leaves obovate to spathulate, apex rounded....59. F. spathulifolia
Leaves of other shapes, apex acute, acuminate, or rounded..8
8. Petiole up to 1 cm long...............9
Petiole more than 1 cm long.........................10
9. Medium-sized tree. Leaves 920 cm long...............34. F. magnoliifolia
Shrubs or small trees. Leaves 2.510.5 cm long........45. F. oleifolia (in part)
10. Basal pairs of lateral veins not distinct, extending less than 1/3 the length of the
blade...........56. F. septica
Basal pairs of lateral veins distinct, extending more than 1/3 the length of the
blade...........3. F. ashtonii
11. Midrib forked...............15. F. deltoidea (in part)
Midrib not forked.............................12
12. Leaf margin revolute................13
Leaf margin plane.................17
13. Leaf apex acute.....................14
Leaf apex obtuse or rounded....................16
14. Leaf margin lobed................15. F. deltoidea (in part)
Leaf margin entire................15
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15. Twig hollow. Leaves spirally arranged or rarely opposite, base rounded...19. F. fistulosa
Twig solid. Leaves mostly opposite, base cuneate........................47. F. pseudotarennifolia
16. Syconia covered with long bracts all over its surface. Shrubs of lowland riverbanks.
...................33. F. macrostyla (in part)
Syconia without such bracts. Shrubs of montane forest.....................45. F. oleifolia (in part)
17. Lateral veins numerous, closely spaced, more or less straight towards the leaf margin
.................7. F. benjamina (in part)
Lateral veins few, well-spaced, curved towards the leaf margin.....18
18. Lateral veins up to 5 pairs; leaf base asymmetric.........64. F. tinctoria (in part)
Lateral veins more than 5 pairs; leaf base symmetric..........19
19. Basal pairs of lateral veins not distinct.................20
Basal pairs of lateral veins distinct...........21
20. Leaf apex cuspidate; glands in the axils of basal pair of lateral veins present. Syconia up to
0.5 cm across, axillary or borne on twigs behind the leaves...51. F. rubrocuspidata
Leaf apex caudate; glands in the axils of basal pair of lateral veins absent. Syconia up to 3.75
cm across, borne in massive clusters on branched leafless twigs arising from the
stem............54. F. schwarzii
21. Leaves glaucous below. Syconia borne on stem and branches; tree monoecious.
.......49. F. racemosa
Leaves not glaucous below. Syconia axillary, or on twigs behind the leaves, or borne on long
runners/stolons arising from the base of stem; or if in dense clusters on the stem and
branches, then tree dioecious.......22
22. Leaf base auriculate, apex acuminate or caudate, acumen 34.5 cm long...........23
Leaf base not auriculate, apex not so...........25
23. Midrib rounded below; petiole up to 0.3 cm long. Syconia on stolons arising from the base of
stem, with lateral bracts.....................................30. F. leptocalama
Midrib keeled below; petiole 0.31 cm long. Syconia axillary, without lateral bracts....24
24. Stipules caducous................38. F. midotis (in part)
Stipules persistent................52. F. rubromidotis
25. Plants with abundant free hanging aerial roots. Lateral veins steeply ascending, departing
from the midrib at an acute angle.........37. F. microcarpa (in part)
Plants without free hanging aerial roots. Lateral veins not steeply ascending, departing from
the midrib at a broad angle...................................................26
26. Leaves narrowly oblong or lanceolate, 618 1.23.2 cm; lateral veins 816 pairs. Syconia
tapered to the base, solitary, axillary. Riverine shrubs...................................27. F. ixoroides
Not this combination of characters.......................27
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27. Intercostal venation scalariform.......28

Intercostal venation reticulate......30
28. Syconia in clusters on stem and older branches........67. F. variegata
Syconia axillary....................29
29. Midrib keeled below. Syconia without lateral bracts........68. F. virens (in part)
Midrib rounded below. Syconia with lateral bracts...........29. F. lepicarpa
30. Petiole 3.57 cm long. Syconia with 23 cm long peduncle.............10. F. callosa
Petiole up to 3 cm long. Syconia sessile or with peduncle up to 1.5 cm long.....31
31. Leaves with cream and yellow dots on the upper surface; lateral veins 46 pairs. Syconia
usually sessile............13. F. chartacea
Leaves without cream and yellow dots on the upper surface; lateral veins 713 pairs. Syconia
distinctly peduncled..........28. F. lamponga
32. Midrib of leaf sunken above.................32. F. macilenta
Midrib of leaf raised or flat above................33
33. Syconia borne on stem and branches, or on short, leafless twigs arising from the stem, or on
stolons..................................................34
Syconia axillary or on the twigs behind the leaves..............36
34. Leaf apex caudate, acumen up to 6 cm long. Syconia with many lateral bracts
.........31. F. leptogramma
Leaf apex acute, acuminate or obtuse. Syconia without lateral bracts.....................35
35. Syconia surface ridged, apex depressed.55. F. scortechinii
Syconia surface more or less smooth, apex rounded.....63. F. tarennifolia
36. Leaf base with a small auricle on one side......38. F. midotis (in part)
Leaf base without an auricle...............3
37. Leaves narrowly oblong, up to 2.2 cm wide; lateral veins 1321 pairs....12. F. chaii
Leaves elliptic-oblong, 3.59 cm wide; lateral veins 58 pairs....58. F. sinuata
38. Leaf margin crenate, denticulate, serrate, or lobed..39
Leaf margin entire........63
39. Leaves often trilobed, glaucous below...24. F. grossularioides (in part)
Leaves not trilobed, not glaucous below......40
40. Midrib sunken above............41
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41. Twig hollow; petiole 77.5 cm long....16. F. diamantiphylla

Twig solid; petiole up to 2 cm long......42
42. Leaf base strongly asymmetric, rounded on one side, cuneate on the other
.......66. F. uncinata (in part)
Leaf base symmetric, cuneate..............65. F. treubii (in part)
43. Syconia borne on stem or on long, finger-like projections arising from branches and stem...44
Syconia on stolons or runners arising from the base of stem, axillary, or borne on the twigs
behind the leaves..............46
44. Leaves sessile; base asymmetric, auriculate on one side; lateral veins sunken
above......26. F. hemsleyana
Leaves with well-developed petiole 17 cm long; base cuneate, symmetric; lateral veins
raised or flat above.......45
45. Twig hollow. Syconia with faint irregular lines and tiny projections, glabrous..
......43. F. obpyramidata (in part)
Twig solid. Syconia with distinct ridges from base to apex, brown short-hairy
........69. F. virescens
46. Syconia on stolons or runners arising from the base of stem...47
Syconia axillary or borne on twigs behind the leaves..50
47. Stipules persistent. Leaf base auriculate on one side.....22. F. geocharis
Stipules caducous. Leaf base not auriculate on one side......48
48. Petiole up to 0.6 cm long. Leaf base symmetric, cuneate...60. F. stolonifera
Petiole 0.52.5 cm long. Leaf base asymmetric, cordate or cuneate on one side and rounded
on the other.......49
49. Leaf base cordate; lateral veins 1024 pairs...35. F. megaleia
Leaf base cuneate on one side and rounded on the other; lateral veins 313 pairs
.......66. F. uncinata (in part)
50. Leaf base rounded, subcordate or truncate. Syconia at least 1 cm across....51
Leaf base usually cuneate. Syconia usually less than 1 cm across......54
51. Twig glabrous.......8. F. bruneiensis
Twig hairy........52
52. Leaves densely velvety hairy on both surfaces......17. F. endospermifolia
Leaves not velvety hairy......53
53. Leaf margin faintly toothed. Syconia short velvety hairy; basal bracts 0.10.15 cm
long............9. F. brunneoaurata
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Leaf margin distinctly toothed. Syconia covered with long hairs; basal bracts 0.30.4 cm
long.......18. F. eumorpha
54. Leaf margin irregularly lobed.........41. F. montana
Leaf margin variously toothed.........55
55. Leaf apex obtuse.......56
Leaf apex acute or acuminate or caudate.....57
56. Leaf surface smooth.........36. F. melinocarpa
Leaf surface scabrous (sandpapery to the touch)............14. F. cumingii
57. Leaf base asymmetric...............58
Leaf base symmetric.................59
58. Twig angled, soft-hairy. Leaves with petiole up to 1 cm long; apex with an acumen of 23 cm
long. Syconia solitary....5. F. aurita
Twig terete, rough or woolly hairy. Leaves sessile or with petiole up to 0.7 cm long; apex
with an acumen of 36.5 cm long. Syconia in pairs or in fascicles of 38..44. F. obscura
59. Syconia peduncled........60
Syconia sessile..........62
60. Syconia with lateral bracts........25. F. gul
Syconia without lateral bracts......61
61. Leaves densely velvety hairy below; intercostal venation scalariform. Common in lowland
forests............21. F. fulva
Leaves not velvety hairy below; intercostal venation reticulate. In hill and montane
forests.......57. F. setiflora
62. Indumentum dark brown. Leaves with 2 glands in the axils of basal pair of lateral veins
below. Confined to montane forest at Mt. Kinabalu....46. F. paramorpha
Indumentum yellowish. Leaves without glands in the axils of basal pair of lateral veins
below. Widely distributed from lowland to montane forests...............................4. F. aurata
63. Leaves glaucous below..24. F. grossularioides (in part)
Leaves not glaucous below..........64
64. Midrib sunken above........65
Midrib raised or flat above.......66
65. Leaf margin revolute........65. F. treubii (in part)
Leaf margin plane...........6. F. beccarii
66. Syconia on stolons arising from the base of stem or on stout, leafless, branched twigs arising
from the base of stem.......67
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Syconia axillary, or borne on stem, on branches behind the leaves, or on finger-like, leafless
branches arising from the stem.........70
67. Leaves with basal pair of lateral veins short and departing from the midrib at a broad angle;
base cordate, subcordate or rounded....68
Leaves with the basal pair of lateral veins distinct and departing from the midrib at acute
angle; base attenuate or cuneate...69
68. Leaves with one gland in each of the axils of basal lateral veins below. Syconia in small
globular clusters along hanging unbranched leafless twigs arising from stemand main
branches, obconical, 0.50.6 cm across.................................39. F. minahassae
Leaves without gland in the axils of basal lateral veins below. Syconia in dense clusters on
stout leafless branched twigs arising from the base of stem, pear-shaped, 2.53 cm
long..................................11. F. cereicarpa
69. Leaf apex obtuse; margin revolute....33. F. macrostyla (in part)
Leaf apex acuminate, acumen 12.5 cm long; margin plane....62. F. subterranea
70. Syconia borne on stem and branches or on finger-like, leafless twigs arising from the stem.71
Syconia axillary or borne on twigs behind the leaves......74
71. Twig hollow. Syconia obpyramidal.....43. F. obpyramidata (in part)
Twig solid. Syconia of other shapes.........72
72. Leaf apex obtuse.............42. F. nota
Leaf apex acuminate, acumen 13 cm long.73
73. Leaves silky hairy especially on midrib below; margin entire...20. F. francisci
Leaves not silky hairy; margin lobed towards apex..40. F. moderata
74. Leaves hairy only on the midrib below...48. F. pubinervis
Leaves hairy all over the lower or both surfaces..75
75. Indumentum whitish. Very tall trees with smooth whitish bole and tall
buttresses.........1. F. albipila
Indumentum brownish or yellowish. Small trees.....76
76. Leaf margin revolute........77
Leaf margin plane.........78
77. Syconia with long lateral bracts....33. F. macrostyla (in part)
Syconia without lateral bracts.........61. F. subfulva
78. Leaves with hairy glands in the axils of lateral veins.....2. F. androchaete
Leaves without hairy glands in the axils of lateral veins.79
79. Leaves up to 8 cm wide; lateral veins 46 pairs...23. F. glandulifera
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Leaves 715.5 cm wide; lateral veins more than 6 pairs...53. F. ruficaulis

80. Stranglers..........81
Climbers, creepers, epiphytes, or scandent shrubs.116
82. Leaves thick-coriaceous, cardboard like; margin revolute70. F. xylophylla (in part)
Leaves very much thinner; margin plane.....83
83. Leaf base rounded....................
F. borneensis Kochummen (in part)
(of Borneo)
Gard. Bull. Sing. 50 (1998) 202. Type: SAN 131862, Borneo, Sabah, Labuk Sugut district, Telupid (holotype KEP).
Monoecious strangler. Young twigs yellowish, grooved. Stipules ovate or lanceolate, c.

1.5 0.5 cm, glabrous or hairy. Leaves coriaceous, glabrous, drying to chocolate-brown;
elliptic or oblong, 6.510.5 1.55 cm, base rounded or cuneate, margin entire, plane,
apex acute; midrib sunken above; lateral veins 46 pairs, curving and joining to form
looped intramarginal vein, distinct below, invisible above, basal pair reaching about 1/3
the length of the blade; intercostal venation reticulate, visible below, invisible above;
petiole 12 cm long, glabrous, drying black. Syconia borne on twigs behind the leaves,
ripening red, sessile, oblong or ellipsoid-oblong, 11.2 0.7 cm, surface rugose with
distinct ridges, apex truncate; basal bracts triangular with acute tip, persistent.
Endemic to Borneo; very rare, known only from six collections including the type from
Sabah and Sarawak. From Sabah recorded for Mt. Kinabalu area, while in Sarawak it is
known from Kapit and Miri (Lambir Hill NP) districts. Lowland to lower montane
forests, up to 1300 m altitude, on ultrabasic or organic soils.
Leaf base cuneate.....84
84. Petiole appressed hairy.....
F. subgelderi Corner
(resembling F. gelderi, a synonym of F. pellucido-punctata)
Gard. Bull. Sing. 17 (1960) 386, l.c. (1965) 17; Kochummen l.c. 157; Anderson l.c. 261; Whitmore, Tantra & Sutisna
l.c. 256; Turner l.c. 354; Coode et al. (eds.) l.c. 218. Type: SFN 29244, Singapore (holotype SING).
Large, monoecious strangler. Twigs slender, irregularly ridged, appressed hairy. Stipules
lanceolate, 1.21.5 cm long, silky hairy outside, caducous. Leaves coriaceous, glabrous;
elliptic to narrowly obovate, 914 2.54.5 cm; base cuneate, margin entire, plane, apex
acute; midrib sunken above; lateral veins 5 pairs, with shorter veins in between, raised
below, faint above, looping and joining near the leaf margin, basal pair reaching up to 1/2
the length of the blade; intercostal venation reticulate, faintly visible below, invisible
above; petiole 1.52.5 cm long, appressed hairy, channelled on the adaxial side. Syconia
axillary, sessile, ellipsoid to oblong, 11.5 0.91.1 cm; surface rugose on drying, apex
flat to slightly depressed; basal bracts prominent, persistent; lateral bracts absent.
Peninsular Malaysia and Borneo. Widely distributed in Sabah, very rare in Sarawak. In
lowland mixed dipterocarp forest. Also in Brunei and Kalimantan.
Petiole glabrous................85
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85. Syconia 11.2 cm diameter......

F. pisocarpa Blume
(Greek, pison = pea, karpos = fruit; the pea-shaped syconia)
Bijdr. Fl. Ned. Ind. (1825) 454; King l.c. (1887) 48; Ridley l.c. (1924) 336; Corner l.c. 17 (1960) 394, l.c. (1965) 20;
Kochummen l.c. 153; Anderson l.c. 260; Whitmore, Tantra & Sutisna l.c. 252; Turner l.c. 353; Coode et al. (eds.) l.c.
217. Type: Blume s.n., Java, Mt. Gede (holotype L?). Synonyms: Urostigma pisocarpum (Blume) Miq., Fl. Ind.-Bat. 1,
1 (1859) 344; U. cycloneuron Miq., Fl. Ind. Bat., Suppl. (1861) 438; F. pseudorubra Miq. l.c. (1867) 264; F. cycloneura
(Miq.) King l.c. (1887) 31; F. microstoma Wall. ex King l.c. (1887) 38.
Monoecious strangler. Twigs yellowish, angled. Stipules lanceolate, c. 1 cm long, hairy,

caducous. Leaves coriaceous, glabrous; obovate or elliptic, 916 38.8 cm, base
cuneate, margin entire, plane, apex obtuse or acute; midrib sunken above; lateral veins 4
6 pairs, looping and joining near the leaf margin, raised below, visible above, basal pair
reaching more than 1/2 the length of the blade; intercostal venation reticulate, visible on
both surfaces; petiole 0.72 cm long, drying black, glabrous. Syconia borne in pairs on
twigs behind the leaves, sessile, subglobose, 11.2 cm diameter, apex umbonate with tiny
perforation; basal bracts persistent; lateral bracts absent.
Thailand, Sumatra, Peninsular Malaysia, Java, and Borneo. Common in Sabah, rare in
Sarawak. In lowland forest by streams to hill forest up to 1000 m altitude. Also in
Kalimantan.
Syconia 0.41 cm diameter..........86
86. Syconia oblong or ellipsoid-oblong.....F. borneensis (in part)
Syconia broadly ellipsoid or subglobose..........87
87. Perianth of male flower consists of 3 tepals. Syconia in axillary pairs, ellipsoid-globose, apex
umbonate, rarely depressed......................
F. sumatrana (Miq.) Miq. (in part)
(of Sumatra)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 263; King l.c. (1887) 40; Backer & Bakhuizen f. l.c. 35; Corner l.c. (1965) 18;
Kochummen l.c. 158; Anderson l.c. 261; Whitmore, Tantra & Sutisna l.c. 256; Turner l.c. 354. Basionym: Urostigma
sumatranum Miq., Pl. Jungh. (1851) 49. Type: Junghuhn s.n. (= RHL Sheet No. 9081881575), Sumatra, Tobing
(holotype L). Synonyms: U. zollingeriana Miq. in Zollinger, Syst. Verz. 2 (1854) 91; U. monadenum Miq. l.c. (1861)
437; F. zollingeriana (Miq.) Miq. l.c. (1867) 264; F. pseudocamptophylla Valeton in Koorders & Valeton, Bijdr.
Booms. Java 11 (1906) 105.
Monoecious strangler. Twigs pale white to pale brown, grooved. Stipules oblong to
lanceolate, acute, up to 1.5 cm long, usually glabrous, caducous. Leaves thick-coriaceous,
glabrous; elliptic or oblong, 4.510 26 cm, base cuneate to rounded, margin entire,
plane, apex acute; midrib sunken above (flat in var. subsumatrana); lateral veins 45
pairs, raised below, very faint above, basal pair reaching more than 1/2 the length of the
blade; intercostal venation reticulate, visible below, invisible above; petiole glabrous,
0.71.5 cm long. Syconia axillary, in pairs, sessile, broadly ellipsoid, 0.40.9 cm
diameter, apex umbonate, rarely depressed; basal bracts prominent, persistent; lateral
bracts absent. Perianth of male flowers consists of 3 tepals.
Burma, Indo-China, Sumatra, Peninsular Malaysia, Java, Borneo, the Philippines,
Sulawesi, and Sumbawa. Uncommon in Sabah and Sarawak. In lowland and hill forests,
up to 800 m altitude. Also in Kalimantan.
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Three varieties are recognised in Sabah and Sarawak, viz. var. sumatrana, var.
subsumatrana (Gagnep.) Corner and var. microcyse Corner (cf. Corner l.c. 17 (1960)
390).
Perianth of male flowers consists of 2 tepals. Syconia borne on twigs behind the leaves,
subglobose, apex rounded........................
F. dulitensis Kochummen
(of Dulit Range, Sarawak)
Gard. Bull. Sing. 50 (1998) 205. Type: Dayang Awa & Yii S. 46743, Borneo, Sarawak, Dulit Range, Belaga district
(holotype KEP; isotype SAR).
Monoecious strangler. Twigs dark brown. Stipules oblong, acute, c. 1 cm long, glabrous,
caducous. Leaves coriaceous, glabrous; elliptic to oblong, 46.5 23.5 cm, base
broadly cuneate, margin entire, plane, apex acute; midrib sunken above; lateral veins 34
pairs, raised below, invisible above, basal pair reaching more than 1/2 the length of the
blade; intercostal venation reticulate, visible below, invisible above; petiole glabrous,
0.51 cm long. Syconia borne on twigs behind the leaves, yellowish when ripe,
subglobose, c. 0.8 cm diameter, apex rounded, sessile, with 3 large basal lanceolate
bracts. Perianth of male flowers consists of 2 tepals.
Endemic to Borneo; very rare, known only by the type from Dulit Range, Belaga district,
Sarawak; in hill forest, up to 800 m altitude.
88. Leaf margin revolute............89
89. Leaves drying blackish; apex obtuse or rounded; lateral veins invisible. Confined to
peatswamp forest..................
F. tristaniifolia Corner
(with leaves resembling those of Tristania, Myrtaceae)
l.c. 258; Turner l.c. 355. Type: SFN 36698, Peninsular Malaysia, Johore, Pontian (holotype SING).
Monoecious strangler. Stipules up to 2.2 cm long, glabrous. Leaves thick-coriaceous,

drying blackish, glabrous; obovate, 58.5 1.55 cm, base cuneate, margin entire,
revolute, apex rounded; midrib flat above; lateral veins 79 pairs, almost invisible, basal
pair reaching to about 1/3 the length of the blade; petiole glabrous, 0.51.2 cm long.
Syconia axillary, in pairs, sessile; subglobose, 0.81 cm across; basal bracts covering
1/31/2 of the receptacle surface; lateral bracts absent.
Sumatra, Peninsular Malaysia and Borneo. In Borneo rare, recorded in Sarawak only from
peatswamp forest.
Leaves drying not blackish; apex acute or acuminate; lateral veins visible.....90
90. Leaves with 12 glands in the axils of basal pairs of lateral veins below. Syconia in clusters
on the twigs behind the leaves..........
F. uniglandulosa Wall. ex Miq.
(Latin, uni- = one, glandulosus = gland-bearing; the basal pair of lateral veins)
In Hooker, Lond. J. Bot. 7 (1848) 277; Corner l.c. (1965) 79; Kochummen l.c. 160; Anderson l.c. 261; Whitmore,
Tantra & Sutisna l.c. 258; Turner l.c. 355; Coode et al. (eds.) l.c. 219. Type: Sine coll., s.n. (= RHL Sheet No.
9081881706), Penang (Prince of Wales Island) (K, L). Synonyms: F. subsubulata Miq. l.c. (1867) 225; F.
uniglandulosa Wall. ex Miq. var. latior Miq. l.c. (1867) 277; F. uniglandulosa Wall. ex Miq. var. parvifolia Miq. l.c.
(1867) 277, syn. nov.; F. copelandii C.B. Robinson, Philip. J. Sc. 3 (1908) Bot. 176.
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Dioecious strangler. Leaves thin- to thick-coriaceous, glabrous, drying greenish yellow;

elliptic or oblong, 521 2.510.5 cm, base cuneate, margin entire, revolute, apex
acuminate, acumen 12 cm long; midrib raised above; lateral veins 47 pairs, with
shorter veins in between, forming a wide angle with the midrib, with a dark brown gland
in the axil of each basal lateral vein below, looping and joining to form a distinct
intramarginal vein, distinctly raised below, visible above; intercostal venation finely
reticulate-areolate, often distinctly visible below, invisible above; petiole 0.71.5 cm
long. Syconia in clusters on the twigs behind the leaves, glabrous, oblong to subglobose,
0.50.7 cm diameter, ostiole often sunken, base tapered into slender stalk 0.30.5 cm
long.
Burma, Thailand, Sumatra, Peninsular Malaysia, Borneo, Sulawesi, and the Philippines.
Very common in Sabah and Sarawak. In lowland and hill forests, including limestone
habitat, up to 900 m altitude. Also in Brunei and Kalimantan.
Leaves without glands in the axils of basal pairs of lateral veins below. Syconia in pairs,
axillary or borne on twigs behind the leaves........91
91. Lateral veins 46 pairs; leaf margin thickened....
F. sundaica Blume
(of the Sundalands)
Bijdr. Fl. Ned. Ind. (1825) 450; Backer & Bakhuizen f. l.c. 35; Corner l.c. (1965) 18; Kochummen l.c. 158; Anderson
l.c. 261; Whitmore, Tantra & Sutisna l.c. 256; Turner l.c. 354; Coode et al. (eds.) l.c. 218; Argent et al. (eds.) l.c. 439.
Type: Blume s.n. (= RHL Sheet No. 908188554), Java, Tjiandjur (holotype L). Synonyms: F. rubescens Blume l.c. 453;
F. indica L. var. sundaica (Blume) Miq. l.c. (1851) 50; Urostigma pyrifolium Miq. l.c. (1859) 338; U. sundaicum
(Blume) Miq. l.c. (1859) 343; U. rubescens (Blume) Miq. l.c. (1859) 338; U. pseudorubrum Miq. l.c. (1859) 343; F.
pyrifolia (Miq.) Boerl., Handl. Fl. Ned. Ind. 3 (1900) 369.
Monoecious strangler. Twigs grey-brown to dark brown, grooved. Stipules lanceolate, 2.5
cm long, velvety hairy outside or glabrous, caducous. Leaves glabrous; elliptic, oblong or
narrowly obovate, 6.521 2.89 cm, base cuneate, margin thickened, entire or lobed,
revolute, apex acute; midrib flat or sunken (var. impressicostata) above; lateral veins 46
pairs, visible on both surfaces, curving and joining near leaf margin, with 23 shorter
veins in between, basal veins reaching about 1/2 or more the length of the blade, without
glands in the axils below; intercostal venation reticulate, visible below, faint above;
petiole 1.53 cm long, channelled on the adaxial side. Syconia in pairs, axillary, sessile,
orange-yellow when ripe, oblong, 1.52 1.22 cm, strongly irregularly wrinkled on
drying, apex disc-like; basal bracts large with round apex, persistent.
Assam, Burma, Indo-China, Thailand, Sumatra, Peninsular Malaysia, Java, and Borneo.
Widely distributed in Sabah and Sarawak, from lowland to lower montane forests, up to
1200 m altitude. Also in Brunei and Kalimantan.
Three varieties are recognised in Sabah and Sarawak, viz. var. sundaica, var.
impressicostata Kochummen, and var. beccariana (King) Corner (cf. Corner l.c. 1965,
and Kochummen l.c. (1998) 217).
Lateral veins 610 pairs; leaf margin not thickened....92
92. Syconia subglobose to globose, 1.21.5 11.5 cm...
F. callophylla Blume
(Greek, kallos = beautiful, phullon = leaf; with beautiful leaves)
Bijdr. Fl. Ned. Ind. (1825) 445; Backer & Bakhuizen f. l.c. 24; Corner l.c. (1965) 22; Kochummen l.c. 142; Whitmore,
Tantra & Sutisna l.c. 242; Turner l.c. 349; Coode et al. (eds.) l.c. 212. Type: Blume s.n. (= RHL Sheet No. 9081861220),
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Java, Tjiandjur, Mt. Sallashie (holotype L). Synonyms: F. tylophylla Hassk., Cat. Hort. Bog. (1844) 75; Urostigma
clusioides Miq. l.c. (1847) 579; Artocarpus venenosa Zoll. & Moritzi var. tylophylla (Hassk.) Miq. l.c. (1854) 89; U.
callophyllum (Blume) Miq. l.c. (1859) 349; F. clusioides (Miq.) Miq. l.c. (1867) 286.
Monoecious strangler. Twigs grey-brown, flat and grooved, glabrous. Stipules lanceolate,
acute, 2.53 cm long, glabrous. Leaves glabrous, drying not blackish; elliptic to oblong
12.524 59.5 cm, base cuneate, margin faintly revolute and lobed, apex acute; midrib
raised above; lateral veins 810 pairs, joining near the leaf margin to form distinct
intramarginal vein, with 23 shorter veins in between, distinct below, faint above, basal
pair without glands in the axils below; intercostal venation faint; petiole 24.5 cm long.
Syconia in pairs, borne on twigs behind the leaves, sessile, subglobose to globose, 1.21.5
11.5 cm, apex umbonate; surface rugose, with persistent large basal bracts.
Indo-China, Hong Kong, Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, the
Philippines, and Sulawesi. Very rare in Sabah, known by 2 collections, SAN A 67 and
SAN 23615, from Sandakan district and Kinabatangan district, respectively; not yet
reported for Sarawak. Also in Brunei and Kalimantan.
Syconia oblong, up to 1 0.6 cm...........
F. chewii Kochummen
(W.L. Chew, 1932, formerly plant taxonomist, Singapore Botanic Gardens)
Gard. Bull. Sing. 50 (1998) 203. Type: RSNB 1972, Borneo, Sabah, Mt. Kinabalu (holotype SAN; isotype SING).
Monoecious strangler. Twigs greyish yellow, finely ridged. Stipules ovate-lanceolate,

acute, 1.51.7 cm long, hairy outside, caducous. Leaves thick-coriaceous, glabrous,
drying not blackish, shiny above; elliptic or oblong, (2.7)611 (1.5)3.55.5 cm, base
rounded or broadly cuneate, margin entire, revolute, apex acute; midrib raised above
with distinct groove in the middle; lateral veins 68 pairs, with shorter veins in between,
curving and joining near the leaf margin, faintly visible on both surfaces, basal veins
reaching about 1/2 the length of the blade, without glands in the axils below; intercostal
venation reticulate, faint below, invisible above; petiole 0.71.7 cm long, drying black.
Syconia solitary on the twigs behind the leaves, sessile, oblong, c. 1 0.6 cm, apex
truncate with disc-like apical bracts; surface irregularly wrinkled; basal bracts persistent,
with rounded apex.
Endemic to Borneo. Very rare, known from the type only from Mt. Kinabalu in Sabah, at
about 1300 m altitude. This species is near to F. sumatrana but the syconia are oblong.
93. Lateral veins numerous, closely spaced, parallel, running straight towards the leaf margin...94
Lateral veins fewer, distantly spaced, curved towards the leaf margin........95
94. Leaves 48 1.83.8 cm. Syconia globose; basal bracts tiny, concealed by the syconia.
Branches drooping.......................7. F. benjamina (in part)
Leaves 1015 3.58.5 cm. Syconia subglobose; basal bracts prominent. Branches not
drooping...........
F. stricta (Miq.) Miq.
(Latin, strictus = drawn close together; probably referring to the syconia)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 288; Corner l.c. (1965) 21; Whitmore, Tantra & Sutisna l.c. 255; Turner l.c. 354.
Basionym: Urostigma strictum Miq. l.c. (1851) 50. Type: Junghuhn 10 (= RHL Sheet No. 9081881495), Java (holotype
L).
Monoecious strangler. Twigs crooked, not drooping, brown, 0.20.3 cm thick. Leaves
glabrous, drying to straw colour; elliptic, 1015 3.58.5 cm, base cuneate, margin
entire, plane, apex acute; midrib flat above; lateral veins numerous, close and parallel,
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very faint on both surfaces, with 23 shorter veins in between; intercostal venation
invisible; petiole 11.2 cm long, channelled on the adaxial side, drying yellowish. Syconia
on twigs behind the leaves, sessile, subglobose, 11.2 cm diameter, irregularly wrinkled
on drying, apex umbonate; basal bracts ovate, prominent, persistent.
Andaman Islands, S China, Indo-China, Sumatra, Peninsular Malaysia, Java, Borneo,
Sulawesi, and the Philippines. Very rare in Sabah and Sarawak; in lowland forest. Not
known in Brunei.
95. Petiole up to 2 cm long.....96
Petiole more than 2 cm long.......100
96. Apex of leaf obtuse......
F. binnendykii (Miq.) Miq. (in part)
(S. Binnendijk, 18211883, sometime adjunct horticulturist, Bogor Botanic Gardens,
Indonesia)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 288; King l.c. (1887) 41; Merrill l.c. (1921) 221; Ridley l.c. (1924) 336; Backer &
Bakhuizen f. l.c. 35; Corner l.c. (1965) 20; Kochummen l.c. 141; Anderson l.c. 257; Whitmore, Tantra & Sutisna l.c.
241; Turner l.c. 349. Basionym: Urostigma binnendykii Miq. l.c. (1859) 341. Type: unknown. Synonyms: U. peracutum
Miq. l.c. (1859) 343; F. peracuta (Miq.) Miq. l.c. (1867) 306.
Monoecious strangler, all parts glabrous. Twigs grey-brown, angled and ridged. Stipules
lanceolate, acute, c. 1.2 cm long, persistent. Leaves thin- to thick-coriaceous, drying
reddish brown or greenish yellow; elliptic or oblong, 411.5 25.5 cm, base rounded or
cuneate, margin entire, plane, apex obtuse; midrib raised above; lateral veins 45 pairs,
distantly spaced, curved, arching and joining near the leaf margin, faint on both surfaces;
intercostal venation reticulate, visible below; petiole 0.71 cm long, drying blackish,
channelled on the adaxial side. Syconia in pairs on twigs behind the leaves, sessile,
depressed globose, c. 0.5 cm diameter, with perforated apex; basal bracts small,
persistent.
Peninsular Malaysia, Java and Borneo. Rare in Sabah and Sarawak. In lowland to hill
forests, up to 1000 m altitude. Not known in Brunei.
Three varieties are recognised in Sabah and Sarawak, viz. var. binnendykii, var. coriacea
Corner, and var. latifolia Corner (cf. Corner l.c. (1965) 20).
Apex of leaf acute................97
97. Leaf blade asymmetric..........64. F. tinctoria (in part)
Leaf blade symmetric...........98
98. Syconia depressed globose with perforated apex......F. binnendykii (in part)
Syconia globose, ellipsoid, or subglobose, apex umbonate, flat to convex, without apical
perforation............99
99. Twigs reddish brown. Basal bracts of syconia small.......
F. delosyce Corner
(Greek, delos = striking, syce = fig-tree)
Gard. Bull. Sing. 17 (1960) 391, l.c. (1965) 19; Kochummen l.c. 144; Anderson l.c. 258;
Whitmore, Tantra & Sutisna l.c. 243; Coode et al. (eds.) l.c. 213; Turner l.c. 349. Type: SFN 28607, Peninsular
Malaysia, Johore (holotype SING). Synonym: F. delosyce Corner var. obtusa Corner l.c. 17 (1960) 391. Type:
Kostermans 5681, Borneo, E Kutei (BO), syn. nov.
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Monoecious strangler. Twigs reddish brown, slender, c. 0.2 cm thick. Stipules glabrous,
ovate to lanceolate with caudate tip c. 1 cm long. Leaves glabrous; elliptic, oblong, or
narrowly obovate, 3.55 1.52.8 cm, base symmetric, cuneate, margin entire, plane,
apex acute, rarely obtuse or rounded; midrib flat above; lateral veins 45 pairs, distantly
spaced, curved, looping and joining near the leaf margin, very faint below, invisible
above, basal pair reaching up to 1/2 of the blade; intercostal venation very faint below,
invisible above; petiole 0.50.8 cm long. Syconia borne on the twigs behind the leaves,
sessile, ellipsoid or subglobose, c. 0.5 cm diameter, ripening orange-yellow, apex
umbonate; basal bracts small, acute, persistent.
Peninsular Malaysia and Borneo. Common in Sabah, not yet recorded for Sarawak. In
lowland forest including swamp forest. Also in Brunei and Kalimantan.
Twigs pale whitish. Basal bracts of syconia large.........F. sumatrana (in part)
100. Plants with abundant free hanging aerial roots....101
Plants without or with a few free hanging aerial roots.....103
101. Lateral veins steeply ascending. Syconia globose.....37. F. microcarpa (in part)
Lateral veins forming a broad angle with the midrib. Syconia oblong or globose..102
102. Syconia oblong, sessile..................
F. drupacea Thunb.
(Latin, drupaceus = drupe-like; the syconia)
Diss. Fic. (1786) 6; Miquel l.c. (1867) 286; King l.c. (1887) 21; Corner l.c. 17 (1960) 380, l.c. (1965) 13; Turner l.c.
350. Type: Thunberg s.n., no locality (UPS). Synonyms: F. chrysocoma Blume l.c. 443; F. pilosa Reinw. ex Blume
l.c. 446; F. payapa Blanco, Fl. Filip. (1837) 683; F. vidaliana Warb. in Perkin, Fragm. Fl. Philip. 3 (1905) 197.
Large, monoecious strangler, with abundant free hanging aerial roots. Stipules reddish
hairy. Leaves at first reddish brown hairy, becoming completely glabrous later; elliptic,
oblong-elliptic or obovate, 826 313 cm, base cuneate to rounded, margin entire,
plane, apex acute; midrib raised above; lateral veins 710 pairs, distantly spaced,
curved, forming a broad angle with the midrib, with few shorter veins in between, raised
on both surfaces, basal pair short; intercostal venation subscalariform, visible below,
invisible above; petiole 14.5 cm long. Syconia axillary, sessile, oblong, 1.53.5 1.22
cm, ripening orange to red; apex disc-like; basal bracts small, concealed by the syconia.
Sri Lanka, S India, Burma, Indo-China, Thailand, Malesia, Solomon Islands, and
Queensland. Very rare in Sabah and Sarawak. In lowland forest by streams. Not known
Syconia globose, peduncled...............
Ficus glaberrima Blume var. bracteata Corner
(Latin, glaberrimus = completely without hairs; the leaves; bracteatus = with bracts, the
syconia)
Gard. Bull. Sing. 17 (1960) 388, l.c. (1965) 17. Type: Hoogland 3748, New Guinea, Garara (holotype LAE).
Synonyms: F. travancorica King l.c. (1887) 28; F. adamii Elmer, Leafl. Philip. Bot. 4 (1911) 1258; F. villamilii Merr.
ex Sata, Monogr. (1944) 183.
Monoecious strangler with abundant free hanging aerial roots. Leaves glabrous; elliptic
or ovate-elliptic, 617 26 cm, base cuneate, margin entire, plane, apex acute; midrib
raised above; lateral veins 715 pairs, distantly spaced, curved, forming a broad angle
with the midrib, with shorter intermediate veins; intercostal venation invisible; petiole
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1.25 cm long. Syconia often solitary, ripening dull orange-yellow to purple-black,

globose, 0.71 cm diameter, apex umbonate; peduncle 0.31 cm long; basal bracts 3,
with distinct round apices.
India, Borneo, the Philippines, and New Guinea. Very rare in Borneo, known only by a
single collection, Purseglove 5183, from Sg. Mayeng, Tatau district, Sarawak.
103. Basal pair of lateral veins distinct and departing at an acute angle from the
midrib........10
4
Basal pair of lateral veins not distinct and departing at a wide angle from the
midrib.......114
104. Syconia covered with irritant hairs.....
F. cucurbitina King
(Latin, cucurbitinus = resembling pumpkin or gourd; the syconia)
Ann. Roy. Bot. Gard. Cal. 1, 1 (1887) 22; Merrill l.c. (1921) 222; Corner l.c. (1965) 14; Kochummen l.c. 144;
Whitmore, Tantra & Sutisna l.c. 243; Turner l.c. 349. Type: Beccari PB 3436, Borneo, Sarawak (FI). Synonym: F.
elliptifolia Merr., Philip. J. Sc. 18 (1921) 55.
Monoecious strangler, without free hanging aerial roots. Twigs reddish brown, strongly
wrinkled on drying. Stipules lanceolate, reddish hairy. Leaves glabrous; oblong to
narrowly obovate, 1118.5 59 cm, base rounded, margin entire, plane, apex acute;
midrib flat above; lateral veins 814 pairs, distantly spaced, curved, faintly looping near
the leaf margin, with short intermediate veins, distinct on both surfaces, basal pair
distinct, departing at an acute angle from the midrib; intercostal venation reticulate,
visible on both surfaces; petiole 1.53 cm long, drying black. Syconia borne on twigs
behind the leaves, sessile, ripening yellow, oblong, 3.54.5 23 cm, with truncate
apex, surface irregularly wrinkled and covered with long irritant hairs.
Peninsular Malaysia, Borneo and the Philippines. Rare in Sabah and Sarawak. Not
known in Brunei. In lowland and hill mixed dipterocarp forests.
Syconia not covered with irritant hairs.....105
105. Syconia sessile..........106
Syconia with distinct peduncles...........111
106. Syconia 23 cm diameter.........107
Syconia 0.51.8 cm diameter...108
107. Leaves thick-coriaceous, apex obtuse. Syconia ellipsoid or oblong......
F. stupenda Miq.
(Latin, stupendus = wonderful, marvellous; probably referring to the syconia)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 286; Backer & Bakhuizen f. l.c. 31; Corner l.c. (1965) 15; Kochummen l.c. 157;
Anderson l.c. 261; Whitmore, Tantra & Sutisna l.c. 255. Type: Zollinger s.n. (= RHL Sheet No. 9081881488), Java,
Mt. Salak (holotype L). Synonym: Urostigma giganteum Miq. l.c. (1854) 90; F. stupenda Miq. var. minor Corner l.c.
17 (1960) 384. Type: Goodenough 3399, Singapore (SING), syn. nov.
Monoecious strangler, without free hanging aerial roots. Twigs grey-brown, strongly
ridged, with prominent leaf scars. Stipules caducous, ovate, c. 1.3 cm long, finely hairy.
Leaves thick-coriaceous, glabrous; elliptic or oblong, 13.525.5 714 cm, base
broadly cuneate to rounded, margin entire, plane, apex obtuse; midrib raised above;
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lateral veins 79 pairs, distantly spaced, curved, raised below, visible above, joining
near the leaf margin, basal pair distinct, departing from the midrib at an acute angle,
reaching up to 1/3 the length of the blade; intercostal venation finely reticulate, faint on
both surfaces; petiole 2.73.5 cm long, channelled on the adaxial side. Syconia borne on
twigs behind the leaves, sessile, ellipsoid or oblong, 2.55 1.53 cm, surface rugose,
not covered by irritant hairs, apex umbonate; basal bracts persistent.
Sumatra, Peninsular Malaysia, Java, and Borneo. Rare in Sabah and Sarawak. Not
known in Brunei. In lowland to lower montane forests, up to 1500 m altitude.
Leaves thinner, apex acute. Syconia ovoid........
F. annulata Blume (in part)
(Latin, annulatus = marked with ring-like swelling; the syconia)
Bijdr. Fl. Ned. Ind. (1825) 448; King l.c. (1887) 25; Merrill l.c. (1921) 220; Ridley l.c. (1924) 33; Burkill l.c. 1020;
Backer & Bakhuizen f. l.c. 31; Corner l.c. (1965) 12, l.c. (1988) 538; Kochummen l.c. 140; Anderson l.c. 256;
Whitmore, Tantra & Sutisna l.c. 240; Turner l.c. 348. Type: Blume s.n. (= RHL Sheet No. 908186925), Java, Mt. Salak
(holotype L). Synonyms: F. flavescens Blume l.c. 449; F. valida Blume l.c. 449; Urostigma annulatum (Blume) Miq.
l.c. (1854) 90; U. biverrucellum Miq. l.c. (1860) 436; F. annulata (Blume) Miq. var. flavescens (Blume) King l.c.
(1887) 26; F. annulata (Blume) Miq. var. valida (Blume) King l.c. (1887) 26; F. balabacensis Quisumbing, Philip. J.
Sc. 41 (1930) 316.
Monoecious strangler, without free hanging aerial roots. Twigs grey, ridged. Stipules
lanceolate, 1.53.5 cm long, acute, glabrous or velvety hairy outside, caducous or
persistent. Leaves thin-coriaceous, drying chocolate-brown, glabrous or hairy below;
elliptic, 1032 311 cm, base wedge-shaped, margin entire, plane, apex acute; midrib
raised above; lateral veins 716 pairs, distantly spaced, with short intermediate veins,
curved, looping and joining near the leaf margin, faint above, distinct below, basal pair
distinct, departing at an acute angle from the midrib; intercostal venation reticulate,
visible below, very faint above; petiole 16 cm long, drying black. Syconia borne on
twigs behind the leaves, golden yellow, ovoid or ellipsoid, 2.53.5 23 cm, apex
umbonate, surface irregularly rugose on drying, not covered by irritant hairs; sessile or
with peduncle up to 2 cm long, with ring-like swelling towards the apex; basal bracts 3,
tiny, persistent.
Burma, Indo-China, Yunan, Thailand, Sumatra, Peninsular Malaysia, Java, Borneo,
Sulawesi, and the Philippines. Common in Sabah, rare in Sarawak. Not known in
Brunei. Lowland and hill forests, up to 600 m altitude.
108. Syconia oblong, apex perforated. Petiole 13 cm long......
F. pellucido-punctata Griff.
(Latin, pellucidus = translucent but not hyaline, punctatus = dotted; the syconia)
Notula 4 (1854) 394; Corner l.c. (1965) 20; Kochummen l.c. 153; Anderson l.c. 260; Whitmore, Tantra & Sutisna l.c.
252; Turner l.c. 352; Coode et al. (eds.) l.c. 216. Type: unknown. Synonyms: F. gelderi Miq. l.c. (1867) 216; F. indica
L. var. gelderi (Miq.) King l.c. (1887) 39; F. everettii Elmer, Leafl. Philip. Bot. 2 (1908) 538.
Monoecious strangler, without free hanging aerial roots. Twigs grey-brown, grooved.
Stipules lanceolate, c. 1.5 cm long, softly hairy outside, caducous. Leaves glabrous;
oblong to elliptic, 5.517 3.55.8 cm, base cuneate, margin entire, plane, apex acute;
midrib flat above; lateral veins 57 pairs, distantly spaced, with 34 short intermediate
veins, faintly raised below, faint to invisible above, curved, looping and joining near the
leaf margin, basal pair distinct and departing at an acute angle from the midrib;
intercostal venation invisible; petiole 13 cm long. Syconia axillary, oblong, 11.5
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0.50.8 cm, sessile; surface strongly wrinkled on drying, not covered by irritant hairs;
basal bracts persistent; apex with small aperture.
Assam, Indo-China, Thailand, Sumatra, Peninsular Malaysia, Borneo, and the
Philippines. Widely distributed in Sabah, rare in Sarawak; in lowland mixed dipterocarp
forest. Also in Brunei and Kalimantan.
Syconia globose, apex not perforated. Petiole 2.56 cm long.....109
109. Syconia up to 0.7 cm diameter.......
F. subtecta Corner
(Latin, sub- = somewhat, tectus = hidden or concealed; the syconia somewhat hidden by
the basal bracts)
Gard. Bull. Sing. 17 (1960) 386, l.c. (1965) 16; Kochummen l.c. 158; Turner l.c. 354. Basionym: F. procera Reinw.
ex Blume (1825), non Salisb. 1796. Type: Blume s.n. (= RHL Sheet No. 9081881391), Java (holotype L). Synonym:
Urostigma procerum (Blume) Miq. l.c. (1859) 336.
Monoecious strangler, without free hanging aerial roots. Twigs dark brown, strongly
rugose. Stipules lanceolate, up to 2.5 cm long, appressed hairy outside, caducous. Leaves
glabrous; elliptic or oblong, 721.5 3.58.5 cm, base broadly cuneate, margin entire,
plane, apex acute; midrib flat above; lateral veins 57 pairs, distantly spaced, curved,
with many short intermediate veins, looping and joining near the leaf margin, visible on
both surfaces, basal pair reaching 1/3 the length of the blade, departing from the midrib
at an acute angle; intercostal venation reticulate, visible on both surfaces; petiole 2.53
cm long, channelled on the adaxial side. Syconia axillary, sessile, globose, c. 0.7 cm
diameter; surface rugose on drying, not covered by irritant hairs; apex umbonate, not
perforated; basal bracts large, persistent.
Indo-China, Sumatra, Peninsular Malaysia, and Borneo. Widely distributed in Sabah,
rare in Sarawak. Not known in Brunei and Kalimantan. In lowland to lower montane
Syconia more than 1 cm diameter........110
110. Basal bracts enclosing 3/4 of the syconium...
F. involucrata Blume
(Latin, involucratus = having an involucre or ring of bracts around the base; the syconia)
Bijdr. Fl. Ned. Ind. (1825) 447; King l.c. (1887) 34; Backer & Bakhuizen f. l.c. 34; Corner l.c. (1965) 18. Type: Blume
s.n., Java, Mt. Salak (L?). Synonyms: Urostigma involucratum (Blume) Miq. l.c. (1859) 334; F. macrocalyx Miq. l.c.
(1867) 217.
Monoecious strangler, without free hanging aerial roots. Twigs c. 0.7 cm thick, greybrown, strongly ridged. Stipules lanceolate, acute, 1.52.5 cm long, glabrous, caducous.
Leaves glabrous; oblong to narrowly obovate, 14.520 47 cm, base cuneate, margin
entire, plane, apex acute; midrib raised above; lateral veins 810 pairs, distantly spaced,
curved, with short intermediate veins, faintly raised on both surfaces, basal pair
reaching to 1/3 the length of the blade, departing from the midrib at an acute angle;
intercostal venation finely reticulate, distinct below, faint above; petiole 3.56 cm long,
flattened and wrinkled. Syconia axillary, sessile, surface not covered by irritant hairs,
globose, c. 1.8 cm diameter, apex umbonate, not perforated; basal bracts persistent,
woody, large, enclosing about 3/4 of the syconium.
Sumatra, Peninsular Malaysia, Java, Borneo, Bali, and the Philippines. Very rare in
Sabah, known only by two collections, KEP 36672 from Labuk Sugut district, and SAN
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A 3683 from Tawau district in Sabah; in lowland mixed dipterocarp forest. Not known in
Brunei, Kalimantan and Sarawak.
Basal bracts of syconia shorter, not enclosing the syconium.........
F. crassiramea (Miq.) Miq.
(Latin, crassus = thick, ramus = branch or twig; with stout twigs)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 286; Corner l.c. 17 (1960) 384, l.c. (1965) 16, l.c. (1988) 540; Kochummen l.c.
144; Anderson l.c. 258; Whitmore, Tantra & Sutisna l.c. 243; Turner l.c. 349; Coode et al. (eds.) l.c. 213. Basionym:
Urostigma crassirameum Miq. l.c. (1851) 48. Type: Junghuhn 5 (= UH Sheet No. 36869), Java, Mt. Ungaran
(holotype U; isotype L). Synonyms: F. procera Reinw. ex Blume var. crassiramea (Miq.) King l.c. (1887) 36; F.
crassicalyx Elmer, Leafl. Philip. Bot. 9 (1937) 3457.
Monoecious strangler, without free hanging aerial roots. Twigs grey-brown or dark
brown, strongly ridged. Stipules ovate to lanceolate, c. 1.8 0.8 cm, glabrous, persistent.
Leaves glabrous; elliptic to oblanceolate, 11.518.5 37 cm, base attenuate, margin
entire, plane, apex obtuse or acute; midrib flat above; lateral veins 78 pairs, with a
number of short intermediate veins, distantly spaced, curved, joining near the leaf
margin, faintly visible on both surfaces, basal pair reaching up to 1/3 the length of the
leaf blade, departing from the midrib at an acute angle; intercostal venation reticulate,
faint above, distinct below; petiole 35.5 cm long, flat and wrinkled on drying. Syconia
borne on twigs behind the leaves, sessile, globose, c. 1.8 cm diameter, ripening orange to
red, apex umbonate, not perforated, surface wrinkled, not covered by irritant hairs;
basal bracts stiff, short, persistent.
Burma, Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, the Philippines, and New
Guinea. Uncommon in Sabah and Sarawak. In lowland forest. Also in Brunei and
Kalimantan.
111. Stipules c. 3.5 cm long, drying pinkish. Syconia in clusters on twigs behind the leaves, up to
0.7 cm diameter......................
F. longistipulata Kochummen
(Latin, longi- = long, stipulatus = stipulate; with long stipules)
Gard. Bull. Sing. 50 (1998) 208. Type: Argent et al. SAN 441987, Borneo, Sabah, Palum Tambun (holotype SAN).
Monoecious strangler, without free hanging aerial roots. Twigs brownish, irregularly
ridged. Stipules lanceolate, acute, c. 3.5 cm long, drying pinkish, persistent. Leaves
glabrous; elliptic to oblong, 11.513.5 56 cm, base rounded or broadly cuneate,
margin entire, plane, apex acute; midrib flat above, drying pinkish below; lateral veins
1215 pairs, distantly spaced, with short intermediate veins, curving and joining near
the leaf margin, distinct below, visible above, basal veins distinct, short, departing from
the midrib at an acute angle; intercostal venation reticulate, faint below, invisible above;
petiole 23 cm long. Syconia in clusters on twigs behind the leaves, subglobose, 0.50.7
cm diameter, surface not covered by irritant hairs, apex swollen and prominently
umbonate; basal bracts small, persistent; peduncle c. 1 cm long, sparsely hairy.
Endemic to Borneo. Very rare, known only by the type collection from Sabah. In
lowland forest, up to 150 m altitude.
Stipules up to 3 cm long. Syconia 13 cm diameter, not in clusters........................112
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112. Midrib strongly keeled below......68. F. virens (in part)

Midrib rounded below..................113
113. Intercostal venation distinctly areolate below; lateral veins indistinct. Syconia
subglobose..............
F. dubia Wall ex King
(Latin, dubius = doubtful; inference unknown)
Ann. Roy. Bot. Gard. Cal. 1, 1 (1887) 46; Ridley l.c. (1924) 333; Corner l.c. (1965) 17; Kochummen l.c. 146;
Anderson l.c. 258; Whitmore, Tantra & Sutisna l.c. 245; Turner l.c. 350; Coode et al. (eds.) l.c. 214. Type: Wall. Cat.
4561, Penang (K?).
Monoecious strangler, without free hanging aerial roots. Twigs brownish, irregularly
wrinkled on drying. Stipules lanceolate, acute, c. 1.5 cm long. Leaves glabrous, drying
chocolate-brown; elliptic or oblong, 814 4.57 cm, base rounded, margin entire,
plane, apex cuspidate with short acumen; midrib flat above, rounded below; lateral veins
79 pairs, distantly spaced, faint on both surfaces, curved, looping and joining near the
leaf margin, basal pair distinct, departing from the midrib at an acute angle; intercostal
venation finely areolate, visible on both surfaces; petiole 23 cm long. Syconia axillary,
solitary, red with greenish yellow spots when ripe, surface not covered by irritant hairs,
irregularly wrinkled on drying, subglobose, 2.5 3.5 1.52 cm; peduncle up to 0.6 cm
long; basal bracts 3, rounded.
Sumatra, Peninsular Malaysia and Borneo. Rare in Sabah and Sarawak. In lowland to
lower montane forests, up to 1200 m altitude. Also in Brunei and Kalimantan.
Intercostal venation reticulate; lateral veins distinct. Syconia ovoid..
..........F. annulata (in part)
114. Petiole hairy............
F. kerkhovenii Valeton
(A.E. Kerkhoven, 18581924, sometime administrator of the Ardjasari tea plantation, W
Java)
In Koorders & Valeton, Bijdr. Booms. Java 11 (1906) 84; Corner l.c. (1965) 15, l.c. (1988) 541; Kochummen l.c. 149;
Whitmore, Tantra & Sutisna l.c. 247; Turner l.c. 351. Type: Kerkhoven s.n., Java, Mt. Malabar, Andjasari (BO?, L?).
Synonym: F. lamaoensis Merr., Philip. J. Sc. 18 (1923) 55.
Monoecious strangler, without free hanging aerial roots. Twigs dark brown. Stipule not
seen. Leaves glabrous; elliptic or oblong, 717 36 cm, base cuneate, margin entire,
plane, apex acute; midrib raised above, rounded below; lateralveins 1012, distantly
spaced, curved, with intermediate veins in between, finely raised on both surfaces, basal
pair indistinct, departing from the midrib at a broad angle; intercostal venation
reticulate, faintly visible on both surfaces; petiole 1.52.5 cm long, whitish hairy,
grooved on the adaxial side. Syconia axillary, bright red when ripe, sessile, subglobose,
c. 1 cm diameter, apex convex; basal bracts small, persistent.
Sumatra, Peninsular Malaysia, Java, Borneo, and the Philippines. Very rare in Sabah and
Sarawak; recorded from lowland mixed dipterocarp forest. Not known in Brunei.
Petiole glabrous............115
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115. Lateral veins with 23 short intermediate veins in between. Syconia sessile....
F. palaquiifolia Corner
(having leaves resembling those of Palaquium, Sapotaceae)
Gard. Bull. Sing. 17 (1960) 399, l.c. (1965) 24; Whitmore, Tantra & Sutisna l.c. 251. Type: SFN 27413, Borneo,
Sabah, Mt. Kinabalu (holotype SING).
Monoecious strangler, without free hanging aerial roots. Twigs dark brown, 0.40.5 cm
thick. Stipules oblong, up to 2.4 cm long, glabrous. Leaves coriaceous, glabrous; elliptic
to subovate, 712 36.5 cm, base cuneate, margin entire, plane, apex acute; midrib
raised above; lateral veins 713 pairs, distantly spaced, with 23 short intermediate
veins in between, curved, looping and joining near the leaf margin, distinct below, faint
above, basal pair indistinct, departing from the midrib at a wide angle; intercostal
venation invisible; petiole 1.83.8 cm long, glabrous. Syconia axillary, in pairs, sessile,
subglobose to pear-shaped, 0.91 cm diameter, surface rough on drying, apex umbonate;
basal bracts rounded, persistent.
Endemic to Borneo. Very rare, known only in Sabah from Mt. Kinabalu by 3 collections
and from Tawau district by one collection (SAN 88206). In lower montane forest, at
about 1200 m altitude.
Lateral veins without short intermediate veins in between. Syconia with peduncles 0.10.3
cm long.......
F. concinna (Miq.) Miq.
(Latin, concinnus = neat, pretty, elegant; probably referring to the syconia)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 286; Corner l.c. (1965) 8; Turner l.c. 349. Basionym: Urostigma concinnum Miq.
l.c. (1847) 570. Type: unknown. Synonyms: F. parvifolia Miq. l.c. (1867) 286; F. affinis Wall. ex Kurz, J. Roy. As.
Soc. Beng. 42 (1873) 105; F. glabella Blume var. concinna (Miq.) King l.c. (1887) 50.
Monoecious strangler, without free hanging aerial roots. Twigs slender, 0.10.2 cm
thick. Stipules lanceolate, often finely hairy along the edges. Leaves elliptic to oblong;
79 23.5 cm, base cuneate, margin entire, plane, apex acute; midrib raised above;
lateral veins 1015 pairs, distantly spaced, curved, without intermediate veins in
between, faint on both surfaces, basal pair indistinct, departing from the midrib at a
wide angle; intercostal venation reticulate, equally prominent as the lateral vein; petiole
1.52.5 cm long, glabrous, slender. Syconia in pairs, axillary or borne on twigs behind
the leaves, ripening white to pink to purple and black, subglobose, 0.40.5 cm diameter,
apex slightly umbonate, internal bristles absent; peduncle 0.10.3 cm long; basal bracts
falling off early.
India, Burma, S China, Indo-China, Thailand, Andaman Islands, Peninsular Malaysia,
Borneo, and the Philippines. Very rare in Borneo, known only by a single collection,
KEP 36377, from Gaya island, Semporna district, Sabah.
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116. Climbers, creepers, or scandent shrubs............117

Epiphytes..........165
117. Leaves glabrous or sometimes pubescent with hook-like or long-straight hairs.....118
Leaves pubescent at least below with short, rough or soft hairs......145
118. Leaf margin toothed.................119
Leaf margin entire, plane or revolute...120
119. Midrib sunken above; leaf apex obtuse. Stipules persistent...
F. tulipifera Corner
(Latin, tulipiferus = bearing tulip-like flowers; referring to the wine-red perianth of the
male flowers)
Gard. Bull. Str. Settl. 10 (1939) 150, l.c. (1965) 62; Whitmore, Tantra & Sutisna l.c. 258. Type: Carr SFN 27763,
Borneo, Sabah, Mt. Kinabalu (holotype SING).
Dioecious creeper. Stipules ovate, 0.41.4 cm long, glabrous, persistent. Leaves

glabrous; elliptic to lanceolate or oblong, 3.57 1.23 cm, base cuneate, margin finely
toothed, apex obtuse; midrib sunken above; lateral veins 811 pairs, looping near the leaf
margin, raised below, sunken above; intercostal venation reticulate with fine areoles,
prominent below; petiole 0.20.9 cm long. Syconia on tubercles arising from stem,
solitary, sessile, depressed globose, c. 3 3.5 cm, apex depressed, smooth to pimply
with small lenticels, sparsely hairy; basal bracts 3.
Endemic to Borneo. Very rare, known only from Mt. Kinabalu in Sabah; in montane
forest.
Midrib flat to slightly raised above; leaf apex acuminate. Stipules caducous.......
F. subsidens Corner
(Latin, subsidens = sunken or depressed; probably referring to the upper part of the
syconia)
Gard. Bull. Sing. 17 (1960) 454, l.c. (1965) 64; Whitmore, Tantra & Sutisna l.c. 256. Type: Carr SFN 26443, Borneo,
Sabah, Mt. Kinabalu (holotype SING).
Dioecious creeper with long internodes. Twigs 0.20.4 cm thick. Stipules caducous.
Leaves glabrous; elliptic, 1722 911.5 cm, base broadly cuneate, margin faintly
toothed, apex acuminate, acumen c. 1 cm long; midrib flat to slightly raised above;
lateral veins 810 pairs, with many short intermediate veins, with a gland in the axil of
each basal lateral vein below; petiole 37 cm long. Syconia axillary or borne on older
branches, rose-red when ripe, subglobose, 0.60.8 cm diameter, tapered into a stalk 0.5
0.8 cm long; with 23 small lateral bracts.
Endemic to Borneo, very rare, known only by 2 collections including the type from Mt.
Kinabalu in Sabah. In lower montane forest near streams, at about 1500 m altitude.
120. Midrib sunken or with distinct groove through the centre above.....121
Midrib raised or flat above...129
121. Leaf apex rounded....122
Leaf apex acute.....123
122. Syconia peduncled, subglobose, c. 0.5 cm diameter, apex umbonate. Confined to montane
forest...........
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F. detonsa Corner
(meaning and inference unknown)
Gard. Bull. Sing. 18 (1960) 18, l.c. (1965) 58; Whitmore, Tantra & Sutisna l.c. 244. Type: Clemens 33456, Borneo,
Sabah, Mt. Kinabalu (holotype BM).
Dioecious climber; stem dark brown. Leaves glabrous; ovate or elliptic, 411 2.26
cm, base broadly cuneate, margin entire, revolute, apex rounded or obtuse; midrib
sunken above; lateral veins 48 pairs, faintly impressed above, raised below; intercostal
venation reticulate, distinct below, invisible above; petiole 0.83 cm long. Syconia
axillary, in pairs, reddish when ripe, subglobose, c. 0.5 cm diameter, apex umbonate;
peduncle 0.20.3 cm long; basal bracts acute.
Endemic to Borneo. In Sabah rare, known only from Mt. Kinabalu in lower montane to
upper montane forests at 16002200 m altitude; not known in Sarawak and Brunei. Also
Syconia sessile, oblong, c. 0.7 cm diameter, apex truncate. Confined to lowland kerangas
forest...........................
F. kerangasensis Kochummen
(of kerangas forest)
Gard. Bull. Sing. 50 (1998) 207. Type: Ilias S. 38595, Borneo, Sarawak, Samarahan district, Sabal FR (holotype KEP;
isotype SAR).
Monoecious climber. Twigs blackish. Stipules lanceolate, acute, 11.4 cm long,

caducous. Leaves glabrous, drying dark brown or blackish; obovate, 67.5 2.54 cm,
base cuneate, margin entire, revolute, apex rounded; midrib sunken above; lateral veins
68 pairs, with short intermediate veins, curving and joining near the leaf margin, faintly
visible below, invisible above, basal pair extending up to 1/3 the length of the blade;
intercostal venation reticulate, faintly visible below, invisible above; petiole 11.5 cm
long, drying black. Syconia borne on twigs behind the leaves, yellowish green when ripe,
sessile, oblong, 0.81 0.7 cm, surface rugose, apex truncate with distinct disc; basal
bracts obtuse, persistent.
Endemic to Borneo. Very rare, known only by the type from Sabal FR, Samarahan
district, in Sarawak; in kerangas forest.
123. Leaf margin revolute............124
Leaf margin plane.............126
124. Stipules persistent. Intercostal venation finely areolate below. Syconia 67 cm diameter,
borne on the stem...........................
F. grandiflora Corner
(Latin, grandi- = large; florus = flower; with large flowers)
Gard. Bull. Str. Settl. 10 (1939) 133, l.c. (1965) 60; Whitmore, Tantra & Sutisna l.c. 246; Coode et al. (eds.) l.c. 215.
Type: Clemens 30290, Borneo, Sabah, Mt. Kinabalu (holotype SING).
Dioecious climber. Stipules lanceolate, 11.7 cm long, persistent. Leaves glabrous;

elliptic, 1016 47 cm, base cuneate, margin entire, revolute, apex acute; midrib
sunken above, keeled below; lateral veins 69 pairs, with short intermediate veins, faintly
visible on both surfaces, basal pair reaching to 1/3 the length of the blade; intercostal
venation finely areolate below; petiole 1.22 cm long. Syconia borne on the stem,
glabrous, marbled, 67 cm diameter.
Endemic to Borneo. Very rare, known only from Mt. Kinabalu, in hill forest, at about
1000 m altitude.
236
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Stipules caducous. Intercostal venation reticulate below. Syconia borne on twigs behind
the leaves, up to 1.5 cm diameter...............125
125. Basal bracts of syconia caducous.......
F. cornerii Kochummen
(E.J.H. Corner, 19061996, formerly prominent Professor of Tropical Botany,
University of Cambridge, U.K.)
Gard. Bull. Sing. 50 (1998) 204. Type: SAN 119744, Borneo, Sabah, Keningau district (holotype KEP; isotype SAN).
Monoecious climber. Twigs yellowish brown, strongly grooved. Stipules caducous.

Leaves glabrous, yellowish green on drying; elliptic to oblong, 1220 46.5 cm, base
cuneate, margin entire, revolute, apex acute; midrib raised above with distinct sunken
groove through the centre; lateral veins 57 pairs, with many short veins in between,
looping and joining near the leaf margin to form a looped intramarginal vein, distinct
below, very faint above, basal pair reaching to almost 1/2 the length of the blade;
intercostal venation reticulate; petiole 2.54 cm, wrinkled on drying. Syconia borne on
twigs behind the leaves, sessile, yellowish when ripe, drying reddish brown, globose,
1.21.5 cm diameter, surface smooth, apex slightly depressed, ostiole closed by 2 bracts;
basal bracts 3, small, caducous, concealed by the syconium base which is thickened and
ring-like with white edge.
Endemic to Borneo. Recorded from Sabah, Sarawak and Brunei. In lowland and hill
mixed dipterocarp forests, up to 800 m altitude.
Basal bracts of syconia persistent...................
F. soepadmoi Kochummen
(E. Soepadmo, 1937, coordinator and chief editor of the Tree Flora of Sabah and
Sarawak Project)
Gard. Bull. Sing. 50 (1998) 212. Type: S. 38376, Borneo, Sarawak, Miri district, Lambir Hill NP (holotype KEP;
isotype SAR).
Monoecious climber. Twigs dark brown, angled. Stipules lanceolate, c. 1.5 cm long,
appressed hairy outside, caducous. Leaves glabrous, drying greenish yellow; oblong or
oblanceolate, 913 2.83.7 cm, base cuneate, margin entire, revolute, apex acute;
midrib sunken above; lateral veins 57 pairs, with a number of short intermediate veins,
curving and joining near the leaf margin to form looped intramarginal vein; intercostal
venation reticulate, distinct below, very faint to inconspicuous above; petiole 11.5 cm
long, drying black, sparsely hairy. Syconia borne on twigs behind the leaves, sessile, in
pairs, pale green when ripe, subglobose, 0.60.8 cm diameter, apex faintly depressed,
ostiole closed by 2 apical bracts, surface rugose with five prominent ridges from apex to
base; basal bracts with rounded apices, persistent.
Endemic to Borneo. Very rare, known only by the type from Sarawak, in lower montane
forest by streams, at about 1400 m altitude. A sterile collection from Brunei, BRUN
5332, from Sg. Belalong, a juvenile stage collection, probably belongs here.
126. Stipules persistent. Syconia sessile, base not narrowed into a long stalk......................
F. paracamptophylla Corner
(Greek, near or resembling F. acamptophylla)
Gard. Bull. Sing. 17 (1960) 387, l.c. (1965) 17; Whitmore, Tantra & Sutisna l.c. 251; Coode et al. (eds.) l.c. 216.
Type: Anderson S. 9103, Borneo, Sarawak, Kuching district, Semengoh FR (holotype SAR).
237
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Monoecious climber. Twigs pale whitish, grooved. Stipules finely hairy outside,
lanceolate, up to 3.5 cm long, persistent. Leaves glabrous, thick-coriaceous; oblong or
lanceolate, 1028 3.510 cm, base rounded to subcordate, margin entire, plane, apex
acute; midrib sunken above, prominent below; lateral veins 811 pairs, faintly raised on
both surfaces, looping and joining near the leaf margin; intercostal venation reticulate,
distinct below, faint above; petiole 1.54 cm long, irregularly wrinkled on drying.
Syconia in pairs on twigs behind the leaves, sessile, subglobose to ellipsoid, 1.31.6 cm
diameter, yellowish when ripe, surface rugose on drying; base not narrowed into a long
stalk, apex disc-like; basal bracts with rounded apices, persistent.
Endemic to Borneo. Uncommon in Sarawak; very rare in Sabah, known by two
collections, SAN 105173 from Beaufort district and SAN 7765 from Sipitang district. In
lowland mixed dipterocarp forest by streams, including kerangas forest. Also in Brunei
and Kalimantan.
Stipules caducous. Base of syconia narrowed into a long stalk.......127
127. Petiole 2.55 cm long. Intercostal venation distinctly areolate below. Syconia apex
umbonate................
F. apiocarpa (Miq.) Miq.
(Greek, apion = pear, karpos = fruit; the pear-shaped syconia)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 269; King l.c. (1888) 70; Merrill l.c. (1921) 220; Ridley l.c. (1924) 338; Burkill
l.c. 1020; Corner l.c. (1965) 60; Kochummen l.c. 140; Whitmore, Tantra & Sutisna l.c. 240; Turner l.c. 348; Coode et
al. (eds.) l.c. 21. Basionym: Urostigma apiocarpa Miq. l.c. (1861) 440. Type: HB 796 (= UH Sheet No. 39621),
Sumatra, Seboga (holotype U; isotype L). Synonyms: F. tetangis Miq. l.c. (1861) 432; Synoecia apiocarpa (Miq.)
Miq. l.c. (1867) 313.
Dioecious climber. Twigs brownish, almost flat, grooved. Stipules lanceolate, c. 1 cm

long, acute, glabrous, caducous. Leaves glabrous; elliptic, ovate or narrowly obovate,
1523 69.5 cm, base rounded to broadly cuneate, margin entire, plane, apex acute;
midrib sunken above; lateral veins 45 pairs, raised below, faint above; intercostal
venation finely reticulate to areolate, very distinct below, invisible above; petiole 2.55
cm long. Syconia axillary, pear-shaped, 46.5 1.52.5 cm, smooth, apex umbonate,
base tapered into a stalk 2.53.5 cm long.
Sumatra, Peninsular Malaysia and Borneo. Rare in Sabah, recorded from Papar and
Beaufort districts only; more common in Sarawak. In lowland mixed dipterocarp forest,
including peatswamp forest. Also in Brunei and Kalimantan.
Two varieties are recognised, viz. var. apiocarpa and var. villosa Corner (cf. Corner l.c.
1965).
Petiole 0.62 cm long. Intercostal venation not distinctly areolate below. Apex of syconia
not umbonate............128
128. Petiole 0.61.2 cm long. Hairs on leaves hook-like...............
F. recurva Blume (in part)
(Latin, recurvus = recurved; the leaf margin)
Bijdr. Fl. Ned. Ind. (1825) 457; Merrill l.c. (1921) 226; Ridley l.c. (1924) 335; Burkill l.c. 1030; Corner l.c. (1965) 56;
Whitmore, Tantra & Sutisna l.c. 252; Turner l.c. 353; Coode et al. (eds.) l.c. 217. Type: Blume s.n. (= RHL Sheet No.
908283226), Java, Bogor, Parang (holotype L). Synonyms: F. strigosa Blume l.c. 441; F. leptocarpa Steud., Nom.
Bot. ed. 2(1840) 636; F. spanogheana Miq. l.c. (1848) 451; F. villipes Miq. l.c. (1848) 451; F. samarensis Merr.,
Philip. J. Sc. 18 (1921) 60.
238
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Dioecious climber, often rooting from stem. Stipules ovate to lanceolate, sparsely or
densely hairy, caducous. Leaves glabrous or pubescent with hook-like hairs, often pitted
on the upper surface; ovate, lanceolate, elliptic or oblong, 316 1.56 cm, base
rounded, truncate or narrowly cordate, margin entire, plane, apex acute; midrib sunken
above; lateral veins 37 pairs, raised below, very faint above, basal pair reaching to 1/2
the length of the blade; intercostal venation not distinctly areolate, faint to distinct
below, faint to invisible above; petiole 0.61.2 cm long. Syconia in clusters on twigs
behind the leaves, glabrous or hairy, depressed globose, 0.50.9 cm diameter, base often
narrowed and stalk-like, sessile or shortly peduncled; apex truncate with distinct ostiole;
basal bracts large, persistent, hairy.
Thailand, Sumatra, Peninsular Malaysia, Java, Lombok, Borneo, and the Philippines.
Uncommon in Sabah and Sarawak. In lowland to lower montane forests, up to 1600 m
altitude. Also in Brunei and Kalimantan.
Five varieties are recognised in Sabah and Sarawak, viz. var. bridelioides Corner, var.
elegantior Corner, var. lasiocarpa Corner, var. recurva, and var. ribesoides Wall. ex
Miq. (cf. Corner l.c. (1965) 56).
Petiole 12 cm long. Hairs on leaves straight................
F. sagittata Koen. ex Vahl
(Latin, sagittatus = shaped like an arrow-head with two equal sharp basal lobes directed
down wards; the leaves)
Symb. Bot. 1 (1790) 83; Backer & Bakhuizen l.c. 22; Corner l.c. (1965) 55; Kochummen l.c. 155; Whitmore, Tantra
& Sutisna l.c. 253; Turner l.c. 353. Type: unknown. Synonyms: F. compressicaulis Blume l.c. 439; F. ramentacea
Roxb., Fl. Ind., ed. Carey, 3 (1832) 547; F. crininervia Miq. l.c. (1861) 432; F. subrigida Miq. l.c. (1861) 433; F.
leptocarpa Steud. var. borneensis Miq., var crassa Miq., var. subglabra Miq., var. timorensis Miq. l.c. (1867) 280.
Dioecious climber. Stipules lanceolate, densely long-hairy, caducous. Leaves glabrous

above, sparsely long-hairy on midrib and veins below, hairs straight; sagittate or elliptic
or oblong, 718 2.58 cm, base cuneate or truncate, margin entire, plane, apex acute;
midrib sunken above; lateral veins 68 pairs, sunken above, distinctly raised below;
intercostal venation invisible on both surfaces or scalariform and visible below; petiole
12 cm long, channelled on the adaxial side. Syconia axillary, in pairs or solitary,
occasionally in clusters, orange or red when ripe, sessile or stalked, glabrous or hairy,
depressed globose, 0.31 cm diameter, constricted at the base into a stalk, apex with a
distinct ostiole; basal bracts small, persistent.
Sikkim, Assam, Chittagong, Burma, Andaman Islands, Thailand, Indo-China, S China,
Hainan, Sumatra, Peninsular Malaysia, Java, Borneo, the Philippines, Sulawesi, Maluku,
Timor, Key Is., and Caroline Islands. Rare in Sabah and Sarawak; in lowland and hill
mixed dipterocarp forests by streams. Not known in Brunei.
Three varieties are recognised in Sabah and Sarawak, namely var. adhaerens (Miq.)
Corner, var. oligosperma (Miq.) Corner, and var. sagittata (cf. Corner l.c. 18, 1960).
129. Leaf margin revolute.........................130
Leaf margin plane.............137
239
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130. Leaf base rounded or subcordate......131

Leaf base cuneate.........136
131. Leaf apex obtuse or rounded; petiole up to 1 cm long. Syconia up to 0.3 cm
diameter..........
F. excavata King
(Latin, excavatus = hollowed out in a curve; the apical part of syconia)
Ann. Roy. Bot. Gard. Calc. 1, 2 (1888) 127; Merrill l.c. (1921) 223; Ridley l.c. (1924) 344; Corner l.c. (1965) 57;
Kochummen l.c. 146; Whitmore, Tantra & Sutisna l.c. 245; Turner l.c. 350. Syntypes: Beccari PB 1368, Borneo (FI);
Kings coll. 5404, 5985, Peninsular Malaysia, Perak (CAL, K?).
Dioecious, scandent shrub. Young twig soft-hairy. Leaves glabrous; ovate to orbicular,
33.75 cm long, base rounded, margin entire, revolute, apex obtuse to rounded; midrib
raised above; lateral veins 35 pairs; intercostal venation reticulate, prominent below;
petiole 0.51 cm long, hairy. Syconia sessile, in clusters, axillary, depressed globose, c.
0.3 cm diameter, orange red when ripe; basal bracts 3, persistent.
Peninsular Malaysia and Borneo. Rare in Sabah and Sarawak; in lowland to hill forests,
up to 900 m altitude. Not known in Brunei.
Leaf apex acute; petiole up to 10 cm long. Syconia up to 6 cm diameter........132
132. Petiole 410 cm long. Syconia up to 6 cm diameter, with warty surface......
F. densechini Corner
(Latin, densus = dense, echinus = slender, prickle- or spine-like appendages; syconia
covered with dense prickles)
Borneo, Sabah, Mt. Kinabalu (holotype SING; isotype K?).
Dioecious root-climber. Twigs pale yellow, hollow, with distinct ring-like scars. Stipules
lanceolate, 1.54.5 cm long, caducous. Leaves glabrous; ovate, 1218 710.5 cm, base
rounded or shallowly cordate, margin entire, revolute, apex acute; midrib raised above;
lateral veins 48 pairs, raised on both surfaces; intercostal venation reticulate, faint with
fine distinct areoles; petiole 410 cm long. Syconia axillary or borne on twigs behind the
leaves, solitary or in pairs, ripening dark brown with paler markings, ellipsoid, c. 7 6
cm, surface armed with short, slender prickles, apex umbonate with 5 large bracts
around the ostiole, base tapered into a stalk; peduncle 0.20.5 cm long; basal bracts 3.
Endemic to Borneo. Very rare, known only from Sabah by 2 collections. In lowland to
lower montane forests, up to 1600 m altitude.
Petiole up to 3.5 cm long. Syconia up to 2 cm diameter......133
133. Leaf with a distinct gland in the axil of each of the basal lateral veins below. Syconia arising
from tubercles on stem and branches.....
F. allutacea Blume
(Latin, allutaceus, mispelling of alutaceus = leather-coloured, pale brown; the drying
leaves)
Bijdr. Fl. Ned. Ind. (1825) 457; King l.c. (1888) 131; Ridley l.c. (1924) 344; Backer & Bakhuizen f. l.c. 22; Corner l.c.
(1965) 58; Kochummen l.c. 139; Whitmore, Tantra & Sutisna l.c. 239; Turner l.c. 348. Type: Blume s.n. (= RHL Sheet
No. 908188212), Java, Mt. Salak (holotype L). Synonyms: F. teysmanniana Miq. l.c. (1859) 319; F. allutacea Blume
var. teysmanniana (Miq.) King l.c. (1888) 131; F. areolata Elmer, Leafl. Philip. Bot. 4 (1911) 1252; F. alutacia
Backer, Bekn. Fl. Java 6 (1948) 21.
240
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Dioecious, scandent shrub. Twigs warty. Stipules ovate to lanceolate, c. 1 cm long,

hairy. Leaves glabrous, upper surface smooth, shining, grey-brown on drying; elliptic,
oblong or ovate, 1017.5 cm long, base rounded, margin entire, revolute, apex acute;
midrib raised above; lateral veins 67 pairs, distinct below, faint to invisible above,
basal veins short, with a black gland in the axil of each vein below; intercostal venation
reticulate, minutely areolate, distinct below, invisible above; petiole 23 cm long.
Syconia solitary, in pairs, or in fascicles on tubercles arising from stem and branches,
globose, 0.71.2 cm diameter, smooth, reddish when ripe; peduncle c. 2.5 cm long; basal
bracts 3, united.
Sumatra, Peninsular Malaysia, Java, Borneo, the Philippines, Sulawesi, and Maluku. In
Sabah and Sarawak very rare, known only from Mt. Kinabalu in Sabah; in hill mixed
dipterocarp forest up to 900 m altitude. Not known in Brunei.
Leaf without glands in the axils of basal lateral veins below. Syconia axillary...134
134. Lower leaf surface scabrous (rough to the touch)......
F. sabahana Kochummen
(of Sabah)
Gard. Bull. Sing. 50 (1998) 211. Type: Ashik SAN 114907, Borneo, Sabah, Pensiangan district, Pandewan, Mesopo R.
(holotype SAN).
Dioecious root-climber. Twigs flat, greyish yellow. Stipules ovate to lanceolate, acute, c.
2 cm long, subpersistent. Leaves glabrous, greyish yellow on drying, undersurface
scabrous; ovate to lanceolate, 12.518.5 710 cm, base cordate or subcordate, margin
entire, revolute, apex acute; midrib flat above; lateral veins 34 pairs, prominently raised
below, flat or faintly raised above, basal pair reaching more than 1/2 the length of the
blade, without glands in the axils below; intercostal venation scalariform or reticulate,
distinct below, faint above; petiole 12 cm long, sparsely hairy. Syconia in clusters,
axillary, globose, 0.61.1 cm diameter, surface smooth on drying, apex depressed, base
narrowed into a cylindrical stalk; basal bracts densely long-hairy.
Endemic to Borneo. Known only by 3 collections from Sabah, SAN 83817 and SAN
118638 from Keningau district, and the type. In disturbed lowland mixed dipterocarp
forest.
Lower leaf surface smooth.......................135
135. Syconia in clusters, base narrowed into a stalk......F. recurva (in part)
Syconia solitary, base not narrowed into a stalk but with distinct peduncle......
F. diandra Corner
(Greek, diandrus = two-stamened; the male flowers)
Gard. Bull. Sing. 19 (1962) 393, l.c. (1965) 60; Whitmore, Tantra & Sutisna l.c. 244. Type: Corner S. 13740, Borneo,
Sarawak (holotype CGE).
Dioecious climber. Twigs dark brown, ridged. Stipules lanceolate, acute, up to 1.2 cm
long. Leaves glabrous, lower surface smooth; elliptic to ovate or lanceolate, 414 2
8.5 cm, base rounded to narrowly cordate, margin entire, revolute, apex acute; midrib
flat above; lateral veins 35 pairs, faint above, curving and joining near the leaf margin,
without glands in the axils of basal pair below; intercostal venation finely reticulate or
areolate; petiole 0.43.5 cm long. Syconia solitary, axillary, subglobose, 1.52 cm
diameter, apex faintly umbonate; surface reddish brown, verrucose; peduncle up to 1 cm
long; basal bracts 3, caducous.
241
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Endemic to Borneo. Very rare, known only by three collections, S. 13740, S. 38570 and
S. 46239 from Kuching district in Sarawak. In lowland forest, including limestone
habitat.
136. Petiole 1.52.5 cm long. Leaves strongly pitted below. Syconia borne on stout, leafless,
finger-like twigs arising from stem, 25 cm diameter.......................
F. iliaspaiei Kochummen
(Ilias Paie, plant collector of the Sarawak Herbarium)
Gard. Bull. Sing. 50 (1998) 207. Type: Ilias S. 42527, Borneo, Sarawak, path to G. Silantek (holotype SAR; isotype
CGE).
Dioecious climber. Twigs reddish brown. Stipules subpersistent. Leaves glabrous, lower
surface distinctly pitted; elliptic, 1012 2.54 cm, base cuneate, margin entire,
revolute, apex acute; midrib raised above; lateral veins 56 pairs, very faintly visible on
both surfaces; intercostal venation reticulate, areolate, distinct below; petiole 1.72 cm
long. Syconia borne on stout, leafless, finger-like twigs arising from the stem, greenish
with white dots when ripe, ellipsoid, c. 7 2.5 cm, apex acute, base tapered into a stalk
up to 1 cm long; peduncle 2.53 cm long; basal bracts tiny.
Endemic to Borneo. Very rare, known only from the type locality in Sarawak. In
lowland mixed dipterocarp forest, at c. 200 m altitude.
Petiole up to 1 cm long. Leaves not pitted below. Syconia on twigs and branches behind the
leaves, subglobose, c. 0.5 cm diameter..
F. gamostyla Kochummen
(Greek, gamo- = united, stulos = style; flowers with united styles)
Gard. Bull. Sing. 50 (1998) 206. Type: SAN 107123, Borneo, Sabah, Kinabatangan district, Tongod (holotype SAN).
Dioecious climber. Twigs dark brown, hollow. Stipules lanceolate, acuminate, c. 0.4 cm
long, caducous. Leaves glabrous, lower surface not pitted; obovate or elliptic, 711
2.55.5 cm, base cuneate, margin entire, revolute, apex acute; midrib flat above; lateral
veins 56 pairs, with shorter veins in between, basal pair short, raised below, very faint
above; intercostal venation finely reticulate, distinct below, invisible above; petiole 0.7
1 cm long. Syconia borne on twigs and branches behind the leaves, in clusters, ripening
red, subglobose, c. 0.5 cm diameter, apex umbonate with slight depression in the centre;
peduncle 11.3 cm long; basal bracts persistent.
Endemic to Borneo. Very rare, known only from the type locality in Sabah. In lowland
mixed dipterocarp forest.
137. Lateral veins more than 15 pairs, closely spaced, straight towards the leaf margin......
F. subcordata Blume
(Latin, sub- = somewhat, cordatus = heart-shaped; the leaf)
Bijdr. Fl. Ned. Ind. (1825) 440. Type: Zippelius 1328 (= RHL Sheet No. 9081882159), Lesser Sunda Islands, Timor
(L).
Only var. malayana is present in Sabah and Sarawak.

var. malayana Corner
(of Malaya)
Gard. Bull. Sing. 17 (1960) 396; l.c. (1965) 21; Kochummen l.c. 157; Whitmore, Tantra & Sutisna l.c. 255; Turner l.c.
354. Type: Curtis s.n., Peninsular Malaysia, Selangor (holotype SING).
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Tall, monoecious climber. Twigs c. 0.6 cm thick. Leaves glabrous, drying greenish
yellow; elliptic, 10.519 3.88.5 cm, base cuneate, margin entire, plane, apex acute;
midrib raised above; lateral veins more than 15 pairs, closely spaced, straight towards
the leaf margin, with shorter veins in between, raised on both surfaces, joining near the
leaf margin to form distinct looped intramarginal vein; intercostal venation invisible;
petiole 1.52 cm long, drying yellow. Syconia subcylindrical, 3.55 22.5 cm; basal
bracts fused and forming a ring-like structure.
Peninsular Malaysia and Borneo. In Sabah and Sarawak, very rare, known by a single
collection SAN A 4365 from Mt. Trus Madi in Sabah, and by three collections from
Lambir Hill NP (RDH 109), Kapit (S. 28380) and G. Santubong (Corner s.n.) in
Sarawak. In hill mixed dipterocarp forest, up to 800 m altitude. Not known in Brunei.
Lateral veins less than 15 pairs, well-spaced, curved towards the leaf margin....138
138. Lateral veins up to 5 pairs....139
Lateral veins more than 5 pairs....143
139. Leaf apex cuspidate....
F. heteropleura Blume
(Greek, heteros = different, uneven, pleuron = rib, nerve; the unevenly veined leaf
blade)
Bijdr. Fl. Ned. Ind. (1825) 466; Corner l.c. (1965) 77; Kochummen l.c. 148; Whitmore, Tantra & Sutisna l.c. 247;
Turner l.c. 351. Type: Blume s.n. (= RHL Sheet No. 9081882147), Java (holotype L). Synonyms: F. radicans Roxb.,
Fl. Ind., ed. Carey, 3 (1832) 536; F. urophylla Wall. ex Miq. l.c. (1848) 429; F. intermedia Griff., Ic. Pl. As. 4 (1854)
t. 556; F. caudatifolia Warb. in Perkin, Fragm. Fl. Philip. 3 (1905) 194; F. eucaudata Elmer, Leafl. Philip. Bot. 1
(1906) 40.
Dioecious, woody climber. Twigs brownish, slender, glabrous or hairy, rounded. Leaves
glabrous or hairy beneath, drying reddish brown; elliptic to narrowly obovate, 614
2.55.5 cm, base cuneate, slightly asymmetric, margin entire, plane, apex cuspidate,
acumen 23 cm long; midrib flat above, sometimes appear impressed; lateral veins 24
pairs, well-spaced, curved, raised below, faint to invisible above, looping and joining
near the leaf margin, basal pair extending to more than 1/2 the length of the blade;
intercostal venation scalariform or reticulate, distinct below, invisible above; petiole 0.5
1.5 cm long. Syconia axillary, rough hairy, globose, 0.40.8 cm diameter, with small
ostiole at the apex; peduncle slender, 0.31 cm long.
Assam, Bhutan, Bengal, Burma, Indo-China, Hainan, Thailand, Sumatra, Peninsular
Malaysia, Java, Borneo, the Philippines, Sulawesi, and Ambon. Common in Sabah and
Sarawak. In lowland to lower montane forests, up to 1800 m altitude. Not known in
Brunei.
Two varieties are recognised in Sabah and Sarawak, viz. var. heteropleura and var. hirta
Corner. The latter differs from the former in having reddish brown hairy twigs and leaf
undersurface, and shorter peduncle up to 0.3 cm long.
Leaf apex rounded or obtuse....140
140. Leaf base asymmetric.......141
Leaf base symmetric.........142
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141. Stipules persistent. Syconia borne on stem, velvety hairy, 48 cm diameter....................

F. punctata Thunb.
(Latin, punctatus = dotted; the lower leaf surface)
Diss. Fic. (1786) 9; Merrill l.c. (1921) 226; Ridl. l.c. (1924) 338; Backer & Bakhuizen f. l.c. 21; Corner l.c. (1965) 61;
Kochummen l.c. 154; Whitmore, Tantra & Sutisna l.c. 252; Turner l.c. 353; Coode et al. (eds.) l.c. 217. Type:
Thunberg s.n., Java (UPS). Synonyms: F. falcata Thunb. l.c. 5; F. macrocarpa Blume l.c. 459.
Dioecious woody creeper. Stipules, twigs and petioles finely appressed hairy. Stipules
persistent. Leaves glabrous, drying yellowish green, lower surface tesselate with minute
stomatal pits; obovate or elliptic, 25 1.53.5 cm, base cuneate, often distinctly
asymmetric, margin entire, plane, apex rounded; midrib flat above; lateral veins 35
pairs, well-spaced, curved, faintly visible below, invisible above; intercostal venation
reticulate, visible below only; petiole 0.21 cm long. Syconia arising from woody
tubercles on the stem and older branches, globose to pear-shaped, 511 48 cm,
rugose, finely velvety hairy; basal bracts 3, triangular; peduncle up to 2 cm long.
Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, and Sulawesi. Uncommon in
Sabah and Sarawak. In lowland to lower montane forests. Also in Brunei and
Kalimantan.
Stipules caducous. Syconia axillary, glabrous, 0.50.6 cm diameter....
F. callicarpides Corner
(Greek, callicarpides = somewhat similar to F. callicarpa, a synonym of F. aurantiacea)
Gard. Bull. Sing. 18 (1960) 17, l.c. (1965) 57; Whitmore, Tantra & Sutisna l.c. 242. Type: Anderson S. 9890, Borneo,
Sarawak, Miri (holotype SAR).
Dioecious, slender climber. Twigs 0.10.15 cm thick, hairy. Stipules c. 0.3 cm long,
glabrous, caducous. Leaves glabrous; elliptic to obovate, 1.23.5 0.42 cm, base
asymmetric, one side cuneate and the other subcordate, margin entire, plane, apex
rounded or obtuse; midrib raised above; lateral veins 34 pairs, well-spaced, curved,
basal pair reaching to 1/2 the length of the blade; intercostal venation invisible; petiole
0.20.4 cm long, hairy. Syconia axillary, in pairs, glabrous, red when ripe; depressed
globose, 0.50.6 cm diameter; basal bracts 3; peduncle 0.10.15 cm long.
Endemic to Borneo. Known only from Sarawak, where it is locally common in
peatswamp forest.
142. Syconia subglobose, up to 0.5 cm diameter.......
F. disticha Blume
(Latin, distichus = arranged in two opposite rows; the leaves)
Bijdr. Fl. Ned. Ind. (1825) 458; Miquel l.c. (1867) 294; Backer & Bakhuizen f. l.c. 21; Corner l.c. (1965) 58;
Kochummen l.c. 145; Turner l.c. 350. Type: Blume s.n. (= RHL Sheet No. 908188547), Java, Mt. Salak (holotype L).
Synonyms: F. disticha Blume var. brunneinervia Hochr., Candollea 2 (1925) 332.
Dioecious creeper. Two minutely notched; midrib flat above; lateral veins 45 pairs,
well-spaced, curved, visible below, invisible above, basal pair reaching to 1/2 the length
of the blade; intercostal venation finely reticulate, areolate, distinct below; petiole 0.31
cm long, channelled on the adaxial side. Syconia in pairs, axillary, yellowish green when
ripe, subglobose, c. 0.5 cm diameter; peduncle 0.20.3 cm long; basal bracts caducous.
Burma, Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, the Philippines,
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Sulawesi, and Maluku. Rare in Borneo, known only from Mt. Kinabalu in Sabah; in hill
mixed dipterocarp to lower montane forests, at 5001800 m altitude.
Syconia ellipsoid to pear-shaped, c. 10 3 cm, with white lines and spots......
F. aurantiacea Griff.
(Latin, auranticeus = orange; the ripe syconia)
Notul. 4 (1854) 394, t. 535; King l.c. (1888) 67; Ridley l.c. (1924) 338; Backer & Bakhuizen f. l.c. 21; Corner l.c.
(1965) 61; Kochummen l.c. 140; Whitmore, Tantra & Sutisna l.c. 240; Turner l.c. 348. Type: unknown. Synonyms:
Synoecia sumatrana Miq. l.c. (1859) 329; F. gibbosa Blume var. pygmea Miq. l.c. (1860) 431; F. callicarpa Miq. l.c.
(1867) 268.
Dioecious root-climber, with tufts of short adventitious roots. Stipules lanceolate, acute,
up to 1.3 cm long, caducous. Leaves glabrous; elliptic to narrowly obovate, 612 1
7.5 cm, base broadly cuneate, symmetric, margin entire, plane, apex rounded; midrib
faint and flat above; lateral veins 34 pairs, well-spaced, curved, faintly raised below,
almost invisible above; intercostal venation areolate, distinct and prominent below,
invisible above; petiole 0.71 cm long. Syconia on twigs behind the leaves or on woody
tubercles arising from stem and branches, glabrous, with pale lines and spots, ellipsoid
or pear-shaped, c. 10 3 cm, ripening orange to carrot-red, apex with projecting
bracts; peduncle c. 2 cm long.
Burma, Thailand, Indo-China, Sumatra, Peninsular Malaysia, Java, Borneo, the
Philippines, Bali, and Sumbawa. Rare in Sabah and Sarawak; in lowland mixed
dipterocarp to lower montane forests, up to 1700 m altitude. Not known in Brunei.
Three varieties are recognised in Sabah and Sarawak, viz. var. angustifolia Corner, var.
aurantiacea, and var. parvifolia Corner. Var. angustifolia and var. parvifolia differ from
var. aurantiacea by the smaller leaves, while var. angustifolia can be distinguished from
var. parvifolia by its narrower leaves.
143. Stipules curving away from the twigs. Leaves with glands in the axils of basal pair of lateral
veins...............
F. subulata Blume.
(Latin, subulatus = awl-shaped; the buds)
Bijdr. Fl. Ned. Ind. (1825) 461; Merrill l.c. (1921) 226; Ridley l.c. (1924) 329; Backer & Bakhuizen f. l.c. 20; Corner
l.c. (1965) 76; Kochummen l.c. 158; Anderson l.c. 261; Whitmore, Tantra & Sutisna l.c. 256; Turner l.c. 354. Type:
Blume s.n. (= RHL Sheet No. 9081882180), Java, in montane forest near Parang (holotype L). Synonyms: F. saxatalis
Blume l.c. 460; F. ellipsoidea Miq. l.c. (1867) 230; F. sikkimensis Miq. l.c. (1867) 225; F. confusa Elmer, Leafl.
Philip. Bot. 1 (1906) 47; F. driveri Elmer, Leafl. Philip. Bot. 7 (1914) 2397.
Dioecious climber. Twigs slender, 0.20.3 cm thick, slightly angled. Stipules narrowly
lanceolate, 1.52.5 cm long, almost always curving away from the twigs. Leaves
glabrous; elliptic to oblong, 6.526 2.510 cm, base cuneate, asymmetric, margin
entire, plane, apex acute; midrib flat above; lateral veins 615 pairs, well-spaced,
curved, with shorter veins in between, faintly visible on both surfaces, often with a gland
in each axil of the basal pair; intercostal venation reticulate, faint; petiole 0.31.2 cm
long. Syconia on twigs behind the leaves, in clusters, ovoid or globose, 0.40.7 cm
diameter; peduncle up to 0.5 cm long; with many bracts at the base of peduncles.
Sikkim, S China, Malesia and Solomon Islands. Common in Sabah and Sarawak; in
lowland mixed dipterocarp to lower montane forests, including limestone forest, up to
1700 m altitude. Not known in Brunei.
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Stipules not curving away from the twigs. Leaves without glands in the axils of basal pair of
lateral veins.......144
144. Leaf base cuneate. Syconia ovoid, 1.52 cm diameter......
F. depressa Blume
(Latin, depressus = flattened from above; the syconia)
Bijdr. Fl. Ned. Ind. (1825) 450; Miquel l.c. (1867) 286; Backer & Bakhuizen f. l.c. 31; Corner l.c. (1965) 12;
Kochummen l.c. 145; Whitmore, Tantra & Sutisna l.c. 244; Turner l.c. 350. Type: Blume s.n., Java, Mt. Salak
(holotype L?). Synonyms: F. pruniformis Blume l.c. 451; F. johnsoni Elmer l.c. (1906) 190; F. iwahigensis Elmer l.c.
(1912) 1381.
Monoecious climber. Twigs grey-white to grey-brown, 0.40.5 cm thick, with distinct

stipular scars. Stipules not curving away from the twigs, lanceolate, acute, c. 1 cm long.
Leaves glabrous; elliptic, 7.516.5 37.5 cm, base cuneate, margin entire, plane, apex
acute; midrib flat or raised above; lateral veins 610 pairs, well-spaced, curved, with
short intermediate veins in between, joining near the leaf margin, raised below, faint
above, without glands in the axils of basal pair; intercostal venation reticulate, faintly
visible on both surfaces; petiole 1.52.5 cm long. Syconia axillary, greenish yellow
when ripe, ovoid to ellipsoid, 23 1.52 cm, surface strongly and irregularly wrinkled
on drying; peduncle 1.53.5 cm long.
Sumatra, Peninsular Malaysia, Java, Borneo, the Philippines, and Bali. Common in
Sabah, rare in Sarawak; in lowland and hill mixed dipterocarp forests, including swamp
forest, up to 600 m altitude. Not known in Brunei.
Leaf base rounded to subcordate. Syconia subglobose, 1.21.5 cm diameter.......
F. globosa Blume
(Latin, globosus = round like a ball; the syconia)
Bijdr. Fl. Ned. Ind (1825) 449; Miquel l.c. (1867) 285; King l.c. (1887) 27; Merrill l.c. (1921) 224; Ridley l.c. (1924)
333; Burkill l.c. 1026; Backer & Bakhuizen f. l.c. 33; Corner l.c. (1965) 12; Kochummen l.c. 147; Whitmore, Tantra &
Sutisna l.c. 246; Turner l.c. 351; Coode et al. (eds.) l.c. 215. Type: Blume s.n. (= RHL Sheet No. 908188756), Java,
Parang (holotype L). Synonyms: Urostigma onustum Wall. ex Miq. l.c. (1847) 575; U. globosum (Blume) Miq. l.c.
(1859) 337; U. manok Miq. in Zollinger l.c. (1854) 90; F. manok (Miq.) Miq. l.c. (1867) 261.
Monoecious climber. Twigs grey-brown, striate. Stipules glabrous, narrowly lanceolate,

acute, c. 3 cm long, not curving away from the twigs. Leaves glabrous, drying to
chocolate-brown; oblong, 8.519 2.58 cm, base broadly rounded to slightly
subcordate, margin entire, plane, apex acute; midrib raised above; lateral veins c. 10
pairs, well-spaced, curved, raised below, faint to invisible above, looping and joining
near the leaf margin, basal pair short, without glands in the axils; intercostal venation
reticulate, faintly visible on both surfaces; petiole 1.54.5 cm long. Syconia axillary,
subglobose, 1.21.5 cm diameter, irregularly wrinkled on drying, apex umbonate.
Burma, Thailand, Sumatra, Peninsular Malaysia, Java, and Borneo. Common in Sabah;
rare in Sarawak, known by a single collection, S. 34786, from the Dulit Range in Belaga
district; in lowland and hill mixed dipterocarp forests, up to 500 m altitude. Also in
145. Leaf margin toothed.........F. tulipifera (in part)
Leaf margin entire, plane or revolute.......146
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Midrib raised or flat above.......158
147. Lateral veins flat or raised above...
F. laevis Blume
(Latin, laevis = smooth; the syconia)
Bijdr. Fl. Ned. Ind. (1825) 437; King l.c. (1888) 128; Ridley l.c. (1924) 344; Backer & Bakhuizen f. l.c. 29; Corner l.c.
(1965) 53; Kochummen l.c. 149; Anderson l.c. 259; Whitmore, Tantra & Sutisna l.c. 248; Turner l.c. 351. Type:
Blume s.n. (= RHL Sheet No. 908188162), Java, Mt. Salak (holotype L). Synonyms: Pogonotrophe emodi Wall. ex
Miq. l.c. (1848) 73; P. emodi Wall. ex Miq. var. oblongata Miq. l.c. (1859) 330; P. emodi Wall. ex Miq. var. cordata
Miq. l.c. (1859) 330; F. subpedunculata Miq. l.c. (1867) 293.
Dioecious climber. Leaves soft-hairy or rarely glabrous; ovate or oblong, 1216 910
cm, base broadly cuneate or rounded, margin entire, plane, apex acute; midrib sunken
above; lateral veins up to 5 pairs, raised below, raised or flat above; intercostal venation
scalariform, faint on both surfaces; petiole 37.5 cm long. Syconia axillary, solitary or in
pairs, globose, 1.52.5 cm diameter, glabrous or hairy, apex depressed; peduncle 1.52
cm long, hairy or glabrous.
India, Sri Lanka, Burma, S China, Indo-China, Thailand, Sumatra, Peninsular Malaysia,
Java, and Borneo. Rare in Sabah and Sarawak; in lowland and hill mixed dipterocarp
forests, up to 1000 m altitude. Not known in Brunei.
Two varieties are recognised in Sabah and Sarawak, viz. var. laevis and var. tomentosa
King. Var. tomentosa differs from var. laevis by the hairy leaves below.
Lateral veins sunken above.......148
148. Leaf apex obtuse....149
Leaf apex acute or acuminate........151
149. Lateral veins 712 pairs.........
F. sarawakensis Corner
(of Sarawak)
Blumea 20 (1972) 427. Type: Anderson & Ilias S. 28606, Borneo, Sarawak, Kapit district, Bt. Tibang (holotype CGE;
isotype SAR).
Large, dioecious root-climber. Twigs flat, young ones hairy. Stipules up to 1 cm long,
subpersistent. Leaves sparsely rough hairy on both sides; lanceolate to elliptic, 915
2.63.8 cm, base weakly asymmetric, cuneate, margin entire, revolute, apex obtuse;
midrib sunken above; lateral veins 712 pairs, sunken above, raised below; intercostal
venation areolate, prominent beneath; petiole up to 3.5 cm long. Syconia crowded on
leafless shoots arising from the stem (young syconium-bearing shoots often persistently
and densely stipulate), ripening red, subglobose to ovoid, c. 4 1 cm, base tapered into
a stalk 0.31.2 cm long, apex acute; peduncle absent or up to 0.5 cm long; basal bracts
caducous; internal bristles sparse, white or brownish.
Endemic to Borneo. Confined to Sarawak, where it is widely distributed in lowland
mixed dipterocarp to lower montane forests, up to 1300 m altitude.
Lateral veins 47 pairs.............150
150. Leaf distinctly asymmetric, with auriculate and cordate base on each side...................
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F. ruginervia Corner
(Latin, ruginervius = with wrinkled veins; the leaves)
Gard. Bull. Sing. 18 (1960) 24, l.c. (1965) 61; Whitmore, Tantra & Sutisna l.c. 253; Turner l.c. 353; Coode et al.
(eds.) l.c. 217. Type: Corner SFN 28446, Peninsular Malaysia, Johore (holotype SING). Synonyms: F. callicarpa
Miq. var. crassinervia Corner l.c. (1939) 118.
Dioecious creeper, with flat stem. Twigs hairy. Leaves soft-hairy; elliptic to obovate, 2
8 14.7 cm, base distinctly asymmetric, one side auriculate and the other broadly
cordate, margin entire, plane, apex obtuse to rounded; midrib sunken above; lateral
veins 37 pairs, sunken above; intercostal venation reticulate, distinct below; petiole
0.31 cm long, hairy. Syconia on tubercles arising from the stem, bright red when ripe,
subglobose, 410 38.5 cm, tapered towards the base; basal bracts tiny, with pointed
apex, persistent; peduncle distinct.
Sumatra, Peninsular Malaysia and Borneo. Rare in Sarawak, not yet reported for Sabah.
In lowland mixed dipterocarp forest. Also in Brunei and Kalimantan.
Leaf symmetric, base cuneate, rounded or truncate...........
F. consociata Blume
(Latin, consociatus = united, connected; the syconia)
Bijdr. Fl. Ned. Ind. (1825) 447; King l.c. (1887) 33; Merrill l.c. (1921) 222; Ridley l.c. (1924) 331; Burkill l.c. 1023;
Backer & Bakhuizen f. l.c. 34; Corner l.c. (1965) 15; Kochummen l.c. 144; Anderson l.c. 257; Whitmore, Tantra &
Sutisna l.c. 243; Turner l.c. 349; Coode et al. (eds.) l.c. 213. Type: Blume s.n. (= RHL Sheet No. 908186949), Java,
Mt. Salak (holotype L). Synonym: Urostigma consociatum (Blume) Miq. l.c. (1854) 91; F. consociata Blume var.
murtoni King l.c. (1887) 34, syn. nov.
Monoecious climber. Twigs greyish brown, strongly ridged, youngest reddish brown
hairy. Stipules lanceolate, c. 1.5 cm long, reddish brown hairy, caducous. Leaves reddish
brown soft-hairy below, glabrescent; oblong to narrowly obovate, 713 36 cm, base
symmetric, cuneate to rounded or truncate, margin entire, plane, apex obtuse; midrib
sunken above; lateral veins 57 pairs, looping and joining near the leaf margin, raised
below, sunken above, especially the basal veins, basal pair reaching almost 1/2 the
length of the blade; intercostal venation reticulate, distinct below, invisible above;
petiole 0.82 cm long, hairy, glabrescent. Syconia on twigs behind the leaves, sessile,
ripening yellow, subglobose, 11.2 cm diameter, apex umbonate, surface smooth or
faintly wrinkled; basal bracts large and persistent.
Burma, Cambodia, Thailand, Sumatra, Peninsular Malaysia, Java, and Borneo. Widely
distributed in Sabah and Sarawak; in lowland mixed dipterocarp forest. Also in Brunei
and Kalimantan.
151. Leaf base cuneate..........152
Leaf base rounded or subcordate..........154
152. Leaves rough hairy below. Syconia solitary, without basal bracts....
F. parietalis Blume
(Latin, parietalis = borne on the wall; inference unknown)
Bijdr. Fl. Ned. Ind. (1825) 462; Merrill l.c. (1921) 225; Ridley l.c. (1924) 330; Masamune l.c. 248; Corner l.c. (1965)
77; Backer & Bakhuizen f. l.c. 25; Kochummen l.c. 153; Anderson l.c. 260; Whitmore, Tantra & Sutisna l.c. 251;
Turner l.c. 352; Coode et al. (eds.) l.c. 216. Type: Blume s.n. (= RHL Sheet No. 908188287), Java, Mt. Parang (holotpe
L). Synonyms: F. cerasiformis Desf., Cat. Hort. Paris ed. 3 (1829) 413; F. grandifolia Wall. ex Miq. l.c. (1848) 432;
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F. phlebophylla Miq. l.c. (1867) 293; F. rufipila Miq. l.c. (1867) 293; F. tabing Miq. l.c. (1867) 293.
Dioecious climber. Twigs yellowish, angled, youngest reddish brown hairy. Stipules
lanceolate, c. 0.5 cm long, reddish brown hairy, caducous. Leaves glabrous above, rough
hairy below; oblong or elliptic, 928 3.810 cm, base cuneate, margin entire, plane,
apex acuminate, acumen 1 cm long or more; midrib sunken above; lateral veins 45
pairs, steeply ascending, raised below, sunken above, basal pair reaching more than 1/2
the length of the blade; intercostal venation scalariform, raised below, faint above;
petiole 0.51.5 cm long, sparsely hairy. Syconia solitary, axillary, ripening orange,
rough hairy, globose, 0.71.3 cm diameter, without basal bracts; peduncle 11.5 cm
long.
Indo-China, Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, and the Philippines.
Widely distributed in Sabah and Sarawak; in lowland and hill mixed dipterocarp forests,
including peatswamp forest, usually by streams. Also in Brunei and Kalimantan.
Leaves soft-hairy below. Syconia in clusters, with persistent basal bracts..........153
153. Stipules caducous...........
F. supperforata Corner (in part)
(meaning and inference unclear)
Gard. Bull. Sing. 18 (1960) 16, l.c. (1965) 57; Whitmore, Tantra & Sutisna l.c. 257; Coode et al. (eds.) l.c. 218. Type:
Mondi 29, Borneo, Kalimantan, Pontianak (holotype L).
Dioecious climber. Twigs 0.20.3 cm thick, hairy. Stipules hairy, caducous. Leaves
thick-coriaceous, sparsely soft-hairy below; elliptic, 918 48 cm, base cuneate or
narrowly subcordate, margin entire, plane, apex acute; midrib sunken above; lateral
veins 57 pairs, sunken above, raised below, with a number of shorter intermediate veins
in between, basal pair reaching to 1/31/2 the length of the blade; intercostal venation
minutely reticulate; petiole 1.23 cm long, appressed hairy. Syconia axillary or borne on
twigs behind the leaves, in fascicles of 26, minutely hairy, sessile, tapered at the base
into a stalk 0.40.6 cm long, subglobose, 0.70.9 cm diameter, apex depressed; basal
bracts persistent.
Endemic to Borneo. Very rare in Sarawak, known by 4 collections from peatswamp
forest; not yet reported for Sabah. Also in Brunei.
Stipules persistent.......
F. spiralis Corner
(Latin, spiralis = spiral; the slender stems spiralling around the supporting trunk of host
tree)
Gard. Bull. Sing. 18 (1960) 13, l.c. (1965) 56. Type: Purseglove 4642, Borneo, Sarawak, Lundu district, Semantan,
Kg. Pueh (holotype CGE).
Dioecious climber. Twigs densely reddish brown hairy. Stipules lanceolate, 1.23 cm
long, hairy, persistent. Leaves rigid-chartaceous, pale glaucous and sparsely soft-hairy
below; elliptic to obovate, 2842 814 cm, base cuneate, margin entire, plane, apex
acute; midrib sunken above; lateral veins 1014 pairs, raised below, sunken above,
curving and joining near the leaf margin; intercostal venation scalariform, raised below,
sunken above; petiole 12 cm long, densely reddish brown hairy. Syconia subsessile, in
clusters, axillary, depressed globose, 0.60.7 cm diameter, reddish brown hairy; basal
bracts persistent.
Endemic to Borneo. Confined to Sarawak and reported for Kapit, Kuching, Lundu, and
Miri (G. Mulu NP) districts. In lowland mixed dipterocarp forest by streams.
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154. Stipules persistent...........

F. villosa Blume (in part)
(Latin, villosus = shaggy with fairly long soft straight hairs; the vegetative parts of the
plant)
Bijdr. Fl. Ned. Ind. (1825) 441; Miquel l.c. (1848) 451; Merrill l.c. (1921) 228; Ridley l.c. (1924) 345; Masamune l.c.
251; Corner l.c. (1965) 56; Backer & Bakhuizen f. l.c. 22; Kochummen l.c. 161; Whitmore, Tantra & Sutisna l.c. 259;
Turner l.c. 355. Type: Blume s.n. (= RHL Sheet No. 908188153), Java, Mt. Salak (holotype L). Synonyms: F. barbata
Wall. ex Miq. l.c. (1848) 441; F. hirsuta Wall. ex Miq. l.c. (1859) 317.
Dioecious climber. Stem, twigs, stipules, petioles and undersurface of leaves densely
reddish brown, woolly or appressed soft-hairy. Stipules oblong, c. 3 cm long, persistent.
Leaves ovate to lanceolate, 1120 4.59 cm, base rounded to subcordate, margin
entire, plane, apex acute; midrib sunken above, often densely hairy in young leaves;
lateral veins 56 pairs, raised below, sunken above, looping and joining near the leaf
margin; intercostal venation scalariform, raised below, sunken above; petiole 1.22 cm
long. Syconia on twigs behind the leaves, or borne on branches or stem, obovoid or
subglobose, 0.61.2 0.61 cm, smooth, apex umbonate or not; peduncle up to 0.5 cm
long; basal bracts absent.
Andaman Islands, Assam, Burma, Indo-China, Thailand, Sumatra, Peninsular Malaysia,
Java, and Borneo. Very rare in Sabah and Sarawak; in lowland and hill mixed
dipterocarp forests, up to 1200 m altitude. Also in Kalimantan.
Three varieties are recognised, viz. var. adpressa Corner (l.c. 18 (1960) 12), var.
subglobosa Corner (l.c. 18 (1960) 13), and var. villosa. Var. adpressa differs from var.
subglobosa and var. villosa by its appressed hairs on the midrib and petiole. Var.
subglobosa differs from var. villosa by its smaller, subglobose, non-umbonate syconia.
Stipules caducous.........155
155. Leaf margin revolute..........
F. grossivenis Miq.
(Latin, grossus = thick, coarse, vena = vein; the coarsely veined leaves)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 227; Merrill l.c. (1921) 224; Masamune l.c. 246; Corner l.c. (1965) 55;
Whitmore, Tantra & Sutisna l.c. 246; Coode et al. (eds.) l.c. 215. Type: De Vriese s.n. (= RHL Sheet No.
9081881299), Maluku, Seram (L). Synonyms: F. propinqua Merr., Publ. Govt. Lab. Philip. 29 (1905) 8; F. lagunensis
Merr., Philip. J. Sc. 9 (1914) Bot. 273; F. jaroensis Merr. l.c. (1914) 268.
Large, dioecious, woody climber. Young stem and twigs covered with long brown soft
hairs. Stipules caducous. Leaves soft-hairy below; ovate to elliptic, 10.520 4.510
cm, base rounded, margin entire, revolute, apex acute; midrib sunken above; lateral
veins 58 pairs, raised below, sunken above, basal pair short; intercostal venation sunken
above; petiole 13 cm long, covered with long soft hairs. Syconia axillary, subglobose,
0.71 cm diameter, smooth; peduncle c. 0.5 cm long; basal bracts caducous.
Borneo, the Philippines, Sulawesi, and Maluku. Very rare in Sabah and Sarawak, known
from Sabah by SAN A 4447 from Keningau district, and from Sarawak by S. 50508 from
Bario; in hill to lower montane forests, at 9001200 m altitude. Also in Brunei and
Kalimantan.
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156. Syconia in pairs, glabrous, not tapered towards the base...

F. forstenii Miq. (in part)
(E.A. Forsten, 18111843, zoologist and elected member of the Commission for Natural
Sciences for the Dutch East Indies)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 214; King l.c. (1887) 29; Merrill l.c. (1921) 223; Corner l.c. (1965) 16;
Kochummen l.c. 147; Whitmore, Tantra & Sutisna l.c. 245; Turner l.c. 350. Type: Forsten s.n. (= RHL Sheet No.
908186979), Sulawesi, Tondano (holotype L). Synonym: F. palawanensis Merr., Publ. Govt. Lab. Philip. 29 (1905)
11.
var. villosa Corner

(Latin, villosus = shaggy hairy; the lower leaf surface)
l.c. 245; Turner l.c. 350. Type: Endert 3344, Borneo, Kalimantan, W Kutai (holotype L).
Monoecious climber. Twigs grey-brown, irregularly wrinkled, finely hairy. Stipules

lanceolate, c. 1.5 cm long, caducous, appressed hairy outside. Leaves shining above,
soft-hairy below; oblong, c. 21 11 cm, base rounded or subcordate, margin entire,
plane, apex acute; midrib sunken above; lateral veins 810 pairs, sunken above, raised
below, distinctly looping near the leaf margin; petiole 22.5 cm long, sparsely hairy.
Syconia sessile, in pairs, glabrous, not tapered towards the base; basal bracts persistent.
Peninsular Malaysia, Borneo and the Philippines. In Borneo, so far recorded only for
Sabah; in lowland mixed dipterocarp forest.
Syconia in clusters, glabrous or hairy, base tapered.....157
157. Lateral veins faintly sunken above................F. supperforata (in part)
Lateral veins distinctly sunken above........
F. lanata Blume
(Latin, lanatus = clothed with woolly hairs; the twigs, petiole and leaf under surface)
Bijdr. Fl. Ned. Ind. (1825) 441; Miquel l.c. (1867) 294; King l.c. (1888) 137; Corner l.c. (1965) 57; Backer &
Bakhuizen f. l.c. 23; Whitmore, Tantra & Sutisna l.c. 248; Coode et al. (eds.) l.c. 215. Type: Blume s.n. (= RHL Sheet
No. 908188153), Java, Mt. Salak (holotype L).
Dioecious climber. Twigs, petioles and leaf undersurface densely soft-hairy. Stipules
caducous. Leaves ovate or lanceolate, 12.519 5.58.5 cm, base rounded, margin
entire, plane, apex acute; midrib sunken above; lateral veins 46 pairs, raised below,
distinctly sunken above, basal pair reaching more than 1/2 the length of the blade;
intercostal venation scalariform, raised below, invisible above; petiole 11.5 cm long,
hairy. Syconia in clusters, axillary, subglobose, 0.40.7 cm diameter, apex perforated,
base tapered into a stalk; peduncle absent.
Sumatra, Borneo and Java. In hill to lower montane forests, at 8001600 m altitude.
Two varieties are recognised in Sabah and Sarawak, namely var. foveolata Corner (l.c.
18 (1960) 16) and var. lanata. The former differs from the latter by its minutely pitted
leaf undersurface.
158.
Petiole 0.20.4 cm long. Leaves cordate or auriculate at the base....

F. barba-jovis Corner
(Latin, barba = beard, jovis = latinised spelling for Jupiter; bearded with tufts of long
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weak hairs resembling that of Jupiters beard; the syconia)

Borneo, Sabah, Mt. Kinabalu, near Kg. Koung (holotype SING).
Dioecious root-climber. Young twigs, petioles and leaf blades sparsely rough-hairy.
Stipules lanceolate, 0.41 cm long, finely appressed hairy. Leaves elliptic or lanceolate,
28.5 13.5 cm, base strongly asymmetric, cordate to auriculate on one side and
cuneate on the other side, margin entire, plane, apex acute; midrib flat or faintly raised
above; lateral veins 69 pairs, raised below, faint to invisible above, arching and joining
near the leaf margin to form distinct intramarginal vein; intercostal venation reticulate,
distinctly visible below; petiole 0.20.4 cm long, brown hairy. Syconia solitary, arising
from short woody tubercles on the stem, rose-pink when ripe, densely hairy, ellipsoid, c.
7 5 cm, apex incompletely closed; peduncle c. 0.5 cm long, hairy; basal bracts 3,
triangular, acute.
Endemic to Borneo. Very rare, known only from Mt. Kinabalu in Sabah; in hill mixed
dipterocarp forest, at about 500 m altitude.
Petiole (0.2)0.72(4.8) cm long. Leaves not cordate or auriculate at the base....159
159. Lateral veins sunken above......160
Lateral veins flat or raised above.....162
160. Leaf margin revolute. Syconia c. 7.5 5 cm.....
F. carri Corner
(C.E. Carr, 18921936; sometime manager of rubber plantations in Malacca and Pahang,
Peninsular Malaysia)
Borneo, Sabah, Mt. Kinabalu, Bundu Tuhan (holotype SING).
Dioecious root-climber, with greyish white stem. Leaves thick-coriaceous, reddish

brown soft-hairy below; elliptic to lanceolate or ovate-lanceolate, 1021 4.89 cm,
base rounded, symmetric, margin entire, revolute, apex acute; midrib flat above; lateral
veins 610 pairs, looping and joining near the leaf margin, raised below, sunken above;
intercostal venation reticulate, sunken above; petiole 2.54.8 cm long, brown-hairy.
Syconia axillary or borne on short woody tubercles arising from the stem; ellipsoid, c.
7.5 5 cm, with distinct neck, ripening olive-brown to purplish brown, velvety whitehairy, surface rugose, apex depressed, base narrowed into a short stalk; peduncle 0.20.3
cm long; basal bracts 3.
Endemic to Borneo, restricted to Mt. Kinabalu in Sabah; in lower montane forest, at
2001700 m altitude.
Leaf margin plane. Syconia up to 1 cm long....161
161. Indumentum comprising hairs with hook-like tips. Syconia-bearing branches not hanging
down...............................................
F. uncinulata Corner
(Latin, uncinulatus = set with fine hooks or barbs; the leaves, twigs and syconia)
Gard. Bull. Sing. 18 (1960) 15, l.c. (1965) 56; Kochummen l.c. 160; Whitmore, Tantra & Sutisna l.c. 258; Turner l.c.
355; Coode et. al. (eds.) l.c. 219. Type: van Steenis 1156, Borneo, Kalimantan, Natuna Island, G. Ranai (holotype L).
Dioecious climber. Twigs, leaves and syconia pubescent with hook-like hairs. Twigs 2
2.5 cm thick, brown. Stipules lanceolate, up to 1 cm long, soft-hairy. Leaves sparsely
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soft-hairy on midrib and veins above, densely soft-hairy below; ovate, elliptic or
lanceolate, 9.518 38 cm, base broadly cuneate, margin entire, plane, apex acute;
midrib raised above; lateral veins 45 pairs, with a number of short intermediate veins
in between, distinct below, sunken above, basal pair reaching more than 1/2 the length of
the blade; intercostal venation reticulate, visible below, faint above; petiole 0.71.5 cm
long, hairy. Syconia borne on twigs behind the leaves, in pairs or in fascicles,
subglobose, 0.70.9 cm diameter, with depressed apex, base narrowed into a stalk 0.2
0.9 cm long.
Peninsular Malaysia and Borneo. Rare, but locally frequent in Sabah and Sarawak,
especially on Mt. Kinabalu. In lowland to hill mixed dipterocarp forests, up to 1000 m
altitude. Also in Brunei and Kalimantan.
Hairs straight. Syconia-bearing branches hanging down....
F. pendens Corner
(Latin, pendens = hanging down from its support; the syconia-bearing branches)
353. Type: Purseglove 4168, Peninsular Malaysia, Pahang, Frasers Hills (holotype SING).
Dioecious, slender climber. Syconia-bearing branches hanging down. Twigs, leaves and
syconia densely covered with reddish brown, rigid, straight hairs. Leaves glabrous
above, hairy below; ovate, elliptic or lanceolate, 4.515 1.87 cm, base subcordate or
cuneate, margin entire, plane, apex acute; midrib raised above; lateral veins 37 pairs,
raised below, sunken above, basal pair reaching to 1/2 the length of the blade; petiole
0.21.5 cm long, hairy. Syconia axillary, in fascicles, sessile or subsessile, subglobose,
0.50.6 cm diameter, apex depressed, base sometimes narrowed into a stalk up to 0.1 cm
long.
Sumatra, Peninsular Malaysia and Borneo. Very rare in Sabah and Sarawak, recorded for
Mt. Kinabalu in Sabah, and Kuching district in Sarawak only. In hill mixed dipterocarp
and lower montane forests, up to 1300 m altitude. Not known in Brunei.
162. Leaf base broadly cuneate......
F. urnigera Miq.
(Latin, urna = urn or a water-pot, -ger = bearing; with pitcher-shaped syconia)
In Zollinger, Syst. Verz. 2 (1854) 92, l.c. (1859) 318; Corner l.c. (1965) 55; Kochummen l.c. 160; Whitmore, Tantra
& Sutisna l.c. 259; Coode et al. (eds.) l.c. 219. Type: Zollinger s.n. (= RHL Sheet No. 9081882002), Java, Bogor
(holotype L). Synonyms: F. recurva Blume var. urnigera (Miq.) King l.c. (1888) 132; F. ramentacea Roxb. var.
urnigera (Miq.) Backer, Bekn. Pl. Java (em. ed.) 6 (1948) 23.
Dioecious climber. Twigs dark brown to reddish brown, slightly grooved. Stipules
lanceolate, velvety hairy, caducous. Leaves drying chocolate-brown, sparsely soft-hairy
on the lower surface, sometimes also on the upper surface; ovate or elliptic, 6.510.5
5.5 cm, base broadly cuneate, margin entire, plane, apex acute; midrib raised above;
lateral veins 34 pairs, raised below, flat above, with few short intermediate veins in
between, basal pair reaching more than 1/2 the length of the blade; intercostal venation
scalariform, visible below, invisible above; petiole 0.51.2 cm long, sparsely hairy.
Syconia borne on twigs behind the leaves, subglobose to urceolate, c. 0.5 cm diameter,
flat and depressed at the apex; basal bracts large, persistent, covered with long hairs.
Lower Burma, Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, and the
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Philippines. Very rare in Sabah and Sarawak, known by one collection each, SAN
127816, from Penampang district in Sabah, and S. 45685 from Kuching district in
Sarawak; in lowland and hill mixed dipterocarp forests. Also in Brunei and Kalimantan.
Leaf base rounded or subcordate..........163
163. Stipules persistent.........F. villosa (in part)
Stipules caducous.........164
164. Leaf apex acute.......F. recurva (in part)
Leaf apex usually obtuse........
F. trichocarpa Blume
(Greek, tricho = hairy, karpos = fruit; the hairy syconia)
Bijdr. Fl. Ned. Ind. (1825) 458; Corner l.c. (1965) 59; Backer & Bakhuizen f. l.c. 22; Kochummen l.c. 159; Whitmore,
Tantra & Sutisna l.c. 257; Turner l.c. 355. Type: Blume s.n., Java, Mt. Pangrango (holotype L?). Synonyms: F.
filiformis Blume l.c. 442; Pogonotrophe piperifolia Miq. l.c. (1854) 93; P. borneensis Miq. l.c. (1859) 330; F.
piperifolia (Miq.) Miq. l.c. (1867) 293; F. piperifolia var. borneensis (Miq.) Miq. l.c. (1867) 293.
Dioecious climber. Twigs glabrous or hairy. Stipules lanceolate, glabrous or rusty

velvety hairy, caducous. Leaves rough-hairy on both surfaces or soft hairy below; ovate,
obovate or oblong, 617 39 cm, base often asymmetric, rounded to subcordate or
rarely cuneate, margin entire, plane, apex rounded or obtuse, rarely acute; midrib flat
above; lateral veins 47 pairs, raised on both surfaces; intercostal venation scalariform
to reticulate, distinct below, faint above; petiole 12 cm long, glabrous or hairy. Syconia
borne on twigs behind the leaves, subglobose, 0.71 cm diameter, velvety hairy to
glabrous, apex umbonate; basal bracts persistent; peduncle 0.51 cm long.
Indo-China, Thailand, Sumatra, Peninsular Malaysia, Java, and Borneo. Rare in Sabah
and Sarawak; in lowland and hill mixed dipterocarp forests, mainly by streams. Also in
Kalimantan.
Two varieties are recognised in Sabah and Sarawak, viz. var. piperifolia Corner (l.c. 18
(1960) 19) and var. trichocarpa. The former can be distinguished from the latter by its
thinly appressed hairy leaves with acute apex, and its larger syconia.
165. Midrib forked...........166
Midrib not forked.........167
166. Lateral veins 45 pairs. Syconia hairy.......
F. deltoidea Jack var. subhirsuta Kochummen
(Latin, deltoideus = deltoid, equilateral-triangular, the leaves; subhirsutus = covered
with somewhat coarse and stiff long erect hairs; the syconia)
Gard. Bull. Sing. 50 (1998) 214. Type: Yii S. 48452, Borneo, Sarawak, Kapit district, Batang Baleh (holotype KEP;
isotype SAR).
Dioecious epiphyte. Twigs dark brown, scaly. Leaves sparsely yellowish hairy below;
obovate, 2.23.5 1.72 cm, base attenuate, margin entire, plane, apex rounded; midrib
forked near the apex; lateral veins 45 pairs, curving and joining near the leaf margin,
distinct below, faint above; petiole 0.30.6 cm long. Syconia axillary, solitary, red when
ripe, ellipsoid, c. 0.3 0.2 cm, rough-hairy; peduncle c. 0.2 cm long; basal bracts
persistent, hairy.
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Endemic to Borneo, very rare, known only by the type collection from Sarawak. In hill
mixed dipterocarp forest, at c. 900 m altitude.
Lateral veins 12 pairs. Syconia usually glabrous, very rarely hairy.....................
F. deltoidea Jack var. recurvata Kochummen
(Latin, recurvatus = recurved; the leaf margin)
Gard. Bull. Sing. 50 (1998) 213. Type: Othman et al. S. 48969, Borneo, Sarawak, Bintulu district, Tubau (holotype
KEP; isotypes SAN, SAR).
Dioecious epiphyte. Leaves thick-coriaceous; deltoid, 7.512 410 cm, base attenuate,
margin entire, revolute, apex rounded; midrib forked, sunken above on the lower half;
lateral veins 1 pair; intercostal venation reticulate, visible below, invisible above; petiole
13 cm long, channelled on the adaxial side. Syconia glabrous, black on drying,
subglobose, c. 1.2 cm diameter, apex umbonate; peduncle short.
Endemic to Borneo. Confined to Sarawak, so far recorded for Bintulu and Kuching
districts. In lower montane forest.
167. Leaves hairy below.................
F. bracteata (Wall. ex Miq.) Miq.
(Latin, bracteatus = with bracts; the syconia)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 285; King l.c. (1887) 23; Merrill l.c. (1921) 221; Ridley l.c. (1924) 331; Burkill
l.c. 1021; Masamune l.c. 243; Corner l.c. (1965) 15; Kochummen l.c. 141; Whitmore, Tantra & Sutisna l.c. 242.
Basionym: Urostigma bracteatum Wall. ex Miq. l.c. (1847) 576. Type: Wall. Cat. 4498, Penang (K?).
Monoecious epiphyte, with free hanging aerial roots. Twigs yellowish, irregularly
angled, youngest reddish brown hairy. Stipules ovate or lanceolate, 23 cm long, densely
reddish brown hairy. Leaves reddish brown hairy below; oblong, 1520 710 cm, base
subpeltate, rounded to subcordate, margin entire, plane, apex cuspidate; midrib not
forked, sunken above; lateral veins 89 pairs, raised below, sunken above, looping and
joining near the leaf margin to form distinct intramarginal vein, basal pair reaching about
1/3 the length of the blade; intercostal venation reticulate, raised below, almost invisible
above; petiole 1.52 cm long, reddish hairy. Syconia axillary, subtended by lanceolate,
acuminate persistent bracts, sessile, covered with brown hairs, globose, c. 1.2 cm
diameter (immature), wrinkled, apex with prominent disc; basal bracts large with blunt
apices, persistent; internal bristles abundant.
Sumatra, Peninsular Malaysia and Borneo. Very rare in Borneo, known only by a single
collection, S. 34097, from Lubok Antu district in Sarawak. In lowland mixed dipterocarp
forest by rocky streams.
Leaves glabrous on both surfaces.........168
168. Leaf margin lobed..............
F. deltoidea Jack var. motleyana (Miq.) Corner (in part)
(James D. Motley, ?1859 engineer and plant collector, mainly in Borneo)
Gard. Bull. Sing. 17 (1960) 424, l.c. (1965) 38, l.c. (1969) 308; Kochummen l.c. 145; Whitmore, Tantra & Sutisna l.c.
244; Coode et al. (eds.) l.c. 214. Basionym: F. motleyana Miq. l.c. (1867) 228. Type: De Vriese s.n. (= RHL Sheet No.
9081881866), Borneo (isotype L). Synonym: F. landonii Sym., J. Mal. Br. Roy. As. Soc. 14 (1936) 359.
Dioecious epiphyte. Twigs crooked. Leaves glabrous; elliptic to lanceolate, 5.518 26

cm, base cuneate, margin distinctly lobed, apex acuminate; midrib not forked, sunken
above; lateral veins 45 pairs, faint above; intercostal venation reticulate, equally
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prominent as lateral veins below; petiole 0.53 cm long. Syconia oblong, 12 0.51
cm, with short peduncles.
Peninsular Malaysia, Sumatra and Borneo. Common in Sabah and Sarawak. Also in
Brunei and Kalimantan. In lowland swampy forest and in limestone habitats.
Leaf margin entire, plane or revolute.......169
169.
Leaf without glands in the axils of basal lateral veins.......

F. acamptophylla Miq.
(Greek, akamptos = stiff, phullon = leaf; with stiff leaves)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 264; King l.c. (1887) 40; Merrill l.c. (1921) 220; Ridley l.c. (1924) 335;
Masamune l.c. 242; Corner l.c. (1965) 20; Whitmore, Tantra & Sutisna l.c. 239; Coode et al. (eds.) l.c. 211.
Basionym: Urostigma acamptophyllum Miq. l.c. (1860) 439. Type: Teijsmann HB 3457, Sumatra, Bangka (holotype
U; isotype L). Synonym: F. pachyphylla King l.c. (1887) 32.
Monoecious epiphyte. Young twigs purplish hairy at first, becoming glabrous, angled.
Stipules c. 1 0.4 cm, hairy outside. Leaves thick-coriaceous, glabrous, faintly glaucous
below, drying chocolate-brown; oblong, oblanceolate or narrowly obovate, 412 1.86
cm, base rounded or broadly wedge-shaped, margin entire, distinctly revolute, apex
rounded or obtuse; midrib not forked, sunken above; lateral veins 78 pairs, joining near
the leaf margin to form looped intramarginal vein, distinct below, faint above, without
glands in the axils of basal pair; intercostal venation reticulate, visible on both surfaces;
petiole 0.51.5 cm long. Syconia borne on twigs behind the leaves, pale orange when
ripe, usually in pairs, sessile, subglobose to oblong, 0.81 cm diameter, slightly
wrinkled, apex umbonate, with an ostiole; basal bracts ovate, acute, persistent.
Sumatra, Borneo and St. Barbe Is. Rare in Sabah, widely distributed in Sarawak; in
lowland forest, including beach, swamp, kerangas and secondary forests. Also in Brunei
and Kalimantan.
Leaf with glands in the axils of basal lateral veins below....170
170. Leaves elliptic, apex acute. Syconia oblong to subglobose...
F. oleifolia King var. impressicostata Kochummen
(Latin, oleifolius = with leaves like that of olive treeOlea, Oleaceae, impressus =
sunken, costa = midrib; the leaves with impressed midrib)
Gard. Bull. Sing. 50 (1998) 216. Type: Ilias S. 40961, Borneo, Sarawak, Kapit district, Melinau (holotype KEP;
isotypes SAN, SAR).
Dioecious epiphyte. Leaves glabrous, elliptic, base cuneate, margin entire and plane,
apex acute; midrib not forked, sunken above; lateral veins very faint to invisible, basal
pair with 12 glands in each axil below; petiole 0.20.3 cm long. Syconia oblong to
subglobose, 0.30.4 cm diameter; peduncle 0.50.8 cm long.
Endemic to Borneo. Reported for Sarawak only. Rare, known by 2 collections, S. 21943
and S. 37304, from Melinau, Kapit district. In hill mixed dipterocarp to lower montane
forests, at 7001300 m altitude.
Leaves narrowed obovate, apex rounded. Syconia globose...............
F. oleifolia King var. dodonaeifolia (Gagnep.) Corner
(Latin, dodonaefolius = with leaves resembling those of Dodonaea, Sapindaceae)
Gard. Bull. Sing. 17 (1960) 425, l.c. (1965) 38; Whitmore, Tantra & Sutisna l.c. 251; Coodeet al. (eds.) l.c. 216.
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Basionym: F. dodonaeifolia Gagnep., Not. Syst. 4 (1927) 90. Type: unknown.
Dioecious epiphyte. Leaves glabrous, sessile or with petiole 0.20.3 cm long; narrowly
obovate, 25 0.40.8 cm, base attenuate, margin entire, revolute, apex rounded;
midrib not forked, sunken above; lateral veins invisible to very faintly visible below,
basal pair with 12 glands in each axil below. Syconia axillary or borne on twigs behind
the leaves, globose, 0.30.4 cm diameter, apex umbonate; peduncle 11.5 cm long.
Endemic to Borneo; recorded for Sarawak only, where it is widely distributed. In
lowland to hill mixed dipterocarp forests, including kerangas forest, up to 1200 m
altitude.
1. Ficus albipila (Miq.) King

(Latin, albus = white, pilus = hairs; the whitish hairs on the twigs and leaves)
Ann. Roy. Bot. Gard. Calc. 1, 2 (1888) 179; Corner l.c. 17 (1960) 408, l.c. (1965) 29; Backer & Bakhuizen f. l.c. 30;
Kochummen l.c. 139; Whitmore, Tantra & Sutisna l.c. 239; Turner l.c. 348. Basionym: Covellia albipila Miq. l.c. (1861) 434.
Type: Teijsmann HB 3545 (= UH Sheet No. 47547 B), Sumatra, Palembang, Muaradua (holotype U). Synonyms: Morus
leucophylla Miq. l.c. (1861) 415; F. mollis Miq. var. albipila (Miq.) Miq. l.c. (1867) 296; F. mallotoides Valeton ex Backer,
Blumea 6 (1948) 304; F. microtricherinos Backer, Bekn. Fl. Java 6 (1948) 40; F. albipila (Miq.) King var. glabra Corner l.c. 17
(1960) 408, Whitmore, Tantra & Sutisna l.c. 239; type: Motley 613, Borneo, Banjarmasin (CGE), syn. nov.
Monoecious deciduous tree, up to 40 m tall; buttresses up to 6 m high. Bark whitish, smooth

resembling that of Koompassia excelsa (Leguminosae). Twigs 0.20.4 cm thick, whitish hairy.
Stipules 0.61.2 cm long, densely appressed hairy, free, paired. Leaves spirally arranged, thincoriaceous, drying yellowish brown, sparsely whitish hairy below; lanceolate, 810 3.55 cm,
base symmetric, broadly cuneate to truncate, margin entire, plane, apex acute; midrib raised
above; lateral veins c. 10 pairs, with short veins in between, faint but visible on both surfaces,
basal veins 24 pairs, short, departing at a broad angle from the midrib, without a gland in each
axil below; intercostal venation reticulate, faint on both surfaces; petiole slender, 3.55.5 cm
long. Syconia paired or solitary, axillary or borne on twigs behind the leaves, ripening yellow to
pink and red, depressed globose, 11.5 cm across, densely to sparsely whitish hairy; peduncle up
to 0.8 cm long; basal bracts 3, ovate, obtuse, 11.5 mm long, appressed hairy, caducous; internal
bristles abundant. Tepals 47, free, red, glabrous. Achenes ellipsoid, c. 1.2 mm long, smooth,
with small ridge along the styles side.
Distribution. S India, Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, Sulawesi, Lesser
Sunda Islands, New Guinea, New Britain, New Ireland, and Queensland, Australia. In Borneo
extremely rare (probably not collected because of its large size). So far known only by a single
collection, Reza Azmi RA 75, from Sg. Resang near Kg. Sukau, Kinabatangan district, in Sabah,
and from Banjarmasin in Kalimantan.
Ecology. In lowland forest by streams. This tree is a nesting place for honey bees, just like
Koompassia excelsa, with which it shares similarity in external bark features and height.
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2. Ficus androchaete Corner

(Greek, andro = male, chaete = long hair; the hairy stamens)
Gard. Bull. Sing. 17 (1960) 438, l.c. (1965) 49; Anderson l.c. 256; Primack l.c. 71; Whitmore, Tantra & Sutisna l.c. 240; Coode
et al. (eds.) l.c. 211. Type: Corner BRUN 5320, Borneo, Brunei, Amo, Ulu Belalong (holotype CGE).
Dioecious tree, up to 10 m tall, 10 cm diameter. Twigs 0.30.4 cm thick, dark brown, slightly
angled, with distinct stipular scars. Stipules lanceolate, acute, c. 0.6 0.1 cm, brownish hairy
outside. Leaves spirally arranged, coriaceous, sparsely brownish rough-hairy on both surfaces;
oblong or obovate, 1126 3.59.5 cm, base wedge-shaped or broadly rounded, margin entire,
plane, apex acuminate, acumen 11.5 cm long; midrib flat above; lateral veins 46 pairs, with
shorter veins in between, curving and joining near the leaf margin to form intramarginal vein,
basal pair reaching up to 1/2 the length of the blade, departing at an acute angle from the midrib,
with a hairy gland in each axil below; intercostal venation reticulate, visible above, distinct
below; petiole 26.5 cm long, brownish rough-hairy, drying yellowish and wrinkled. Syconia
axillary, ripening yellow to dark orange, pear-shaped, 0.81.4 0.81.1 cm, apex umbonate;
peduncle slender, 0.81.1 cm long, with tiny persistent bracts at its apex; basal bracts ovate, 23
mm long; internal bristles numerous, up to 1.5 mm long. Tepals 34, white, brownish hairy.
Achenes c. 1.3 mm long, tuberculate, keeled.
Vernacular names. Sarawakkumpan (Iban), lunok (Murut).
Distribution. Endemic to Borneo. Rare with localised distribution in Sabah and Sarawak. Also
Ecology. In lowland and hill mixed dipterocarp forests, up to 1000 m altitude, mainly by
streams.
3. Ficus ashtonii Kochummen

(P.S. Ashton, 1934, sometime forest botanist at the Forestry Departments of Brunei and
Sarawak; Professor of Dendrology, Harvard University, U.S.A.)
Gard. Bull. Sing. 50 (1998) 201. Type: Dayang Awa & Lee S. 47846, Borneo, Sarawak, Limbang (holotype KEP; isotypes CGE,
K, L, SAN, SAR).
Monoecious tree, up to 30 m tall, 60 cm diameter. Twigs yellowish brown, irregularly ridged.

Stipules ovate to lanceolate, acute, c. 2.5 cm long, caducous. Leaves spirally arranged,
coriaceous, glabrous; elliptic, 14.517.5 6.59 cm, base broadly cuneate, margin entire, plane,
apex acute; midrib sunken above; lateral veins 67 pairs, curving and joining near the leaf
margin, raised below, faintly raised above, basal pair reaching more than 1/3 the length of the
blade, departing at an acute angle from the midrib; intercostal venation scalariform, distinct
below, invisible above; petiole 2.23.7 cm long, wrinkled on drying. Syconia axillary, ripening
orange to deep red, sessile, oblong, c. 2 1 cm, apex flat with disc-like bracts; basal bracts large,
united and cup-shaped, with rounded apex, persistent. Tepals 3, lanceolate, acuminate. Achenes
ovoid, c. 2 mm long, with a few irregular protrusions.
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Distribution. Endemic to Borneo. Rare, known only from Bario and Limbang districts in
Sarawak.
4. Ficus aurata (Miq.) Miq.

(Latin, auratus = flecked with gold; the golden yellow hairs on the leaves)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 271; Corner l.c. (1965) 49, l.c. (1988) 546; Kochummen l.c. 140; Anderson l.c. 257; Primack
l.c. 72; Whitmore, Tantra & Sutisna l.c. 240; Turner l.c. 348; Coode et al. (eds.) l.c. 212. Basionym: Covellia aurata Miq. l.c.
(1861) 433. Syntypes: Teijsmann HB 3426, Sumatra, Bangka, Djebus (U); Teijsmann HB 3713, Sumatra, Palembang, Ogan Ulu
(U); Teijsmann HB 4517, Sumatra, Lampong (U). Synonyms: F. aurata (Miq.) Miq. var. densiserra Miq. l.c. (1867) 271; F.
chrysocarpa (non Reinw. ex Blume) King l.c. (1888) 151; F. chrysocarpa (non Reinw. ex Blume) King var. undulata Gibbs, J.
Linn. Soc. Bot. 42 (1914) 137.
Dioecious tree, up to 10 m tall, 10 cm diameter. Bark whitish, smooth. Young twigs, stipules,
lower leaf surface, petioles, and syconia densely velvety with golden yellow hairs, or rarely
rough-hairy (var. brevipilosa). Twigs 0.20.5 cm thick, brownish, ridged, with distinct stipular
scars. Stipules lanceolate, acute, up to 1.5 cm long. Leaves spirally arranged, thin-coriaceous,
sparsely rough hairy above; elliptic, narrowly oblong or obovate, (5.5)10.528 (3)415 cm,
base symmetric, cuneate, margin distinctly toothed from the base to apex, apex acute; midrib
slightly raised above; lateral veins 46 pairs, with shorter veins in between, faint above, raised
below, basal pair reaching almost 1/2 the length of the blade, departing at an acute angle from
the midrib, with a faint gland in each axil below; intercostal venation scalariform to reticulate,
distinctly raised below, faint to invisible above; petiole 0.77 cm long, wrinkled on drying.
Syconia axillary or borne on twigs behind the leaves, ripening yellow, sessile, globose, up to 1(
2) cm diameter, with umbonate apex, surface strongly wrinkled on drying, velvety yellowish
hairy; basal bracts 3, ovate, acute, 23.5 mm long; internal bristles numerous. Tepals 34,
glabrous or densely hairy, yellowish. Achenes ellipsoid, up to 1 mm long, keeled, tuberculate.
Vernacular names. Sarawakkumpan, tempan (Iban).
Distribution. Indo-China, Sumatra, Peninsular Malaysia, and Borneo. Common in Sabah and
Ecology. In lowland to lower montane forests, up to 1300 m altitude; growing in a variety of
soils and habitats, including secondary forest.
Key to varieties
1. Hairs on twigs, petioles and syconia sparse, up to 1 mm long................
var. brevipilosa Corner
(Latin, brevi = short, pilosus = hairy with ascending hairs; the twigs petioles and syconia)
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Gard. Bull. Sing. 17 (1960) 437, l.c. (1965) 49; Whitmore, Tantra & Sutisna l.c. 240. Type: Bur. Sci. 602, Borneo,
Sabah (PNH?). Synonym: F. inaequipetiolata Merr., Philip. J. Sc. 21 (1922) 517.
Common in Sabah, not yet recorded for Sarawak.

Hairs on twigs, petioles and syconia 14 mm long...............2
2. Leaves drying dark brown...............
var. palawanensis Corner
(of Palawan)
Gard. Bull. Sing. 17 (1960) 437, l.c. (1965) 49; Whitmore, Tantra & Sutisna l.c. 240; Turner l.c. 348. Type: Bur. Sci.
77382, the Philippines, Palawan (holotype SING).
In Borneo very rare, known by a single collection from Sabah.

Leaves drying golden yellow.................3
3. Hairs 24 mm long. Leaves narrowly oblong..............
var. longipilosa Corner
(Latin, longus = long, pilosus = hairy with ascending hairs; the twigs, petioles and
syconia)
348; Coode et al. (eds.) l.c. 212. Type: SFN 32313, Peninsular Malaysia, Perak (holotype SING).
Sumatra, Peninsular Malaysia, and Borneo. Very rare in Sarawak, known by a single
collection, S. 3924, from Belaga district.
Hairs 12 mm long. Leaves obovate or elliptic.....................var. aurata
5. Ficus aurita Reinw. ex Blume

(Latin, auritus = with ear-like lobes; the leaf base)
Bijdr. Fl. Ned. Ind. (1825) 462; Corner l.c. (1965) 77; Whitmore, Tantra & Sutisna l.c. 241. Type: Reinwardt 1451 (= RHL Sheet
No. 9081861497), Maluku, Nusa Laut (holotype L). Synonym: F. hispidulosa Elmer, Leafl. Philip. Bot. 7 (1914) 2401.
Dioecious shrub or small tree, up to 7 m tall. Twigs up to 0.2 cm thick, angled, soft-hairy.
Stipules paired, lanceolate, up to 2 cm long, hairy, persistent or caducous. Leaves distichous,
thin-coriaceous, soft-hairy below; elliptic to lanceolate, 7.523.5 2.311 cm, base asymmetric,
cuneate, distinctly auriculate or not, margin slightly toothed or incised, apex acuminate, acumen
23 cm long; midrib raised above, with a gland at the base below; lateral veins 48 pairs, raised
below, faint above, basal pairs short, departing at a broad angle from the midrib; intercostal
venation reticulate, visible below, invisible above; petiole up to 1 cm long, hairy. Syconia
axillary, solitary, fulvous hairy, ripening orange to red, globose or ellipsoid, up to 0.6 cm
diameter, apex umbonate; lateral bracts 12; internal bristles absent. Tepals 34, white,
glabrous, entire or with toothed margin, free. Achenes up to 1 mm long, slightly compressed,
distinctly keeled at the apex.
Vernacular name. Sarawakpunoh (Kelabit).
Distribution. Borneo, the Philippines, Sulawesi, Maluku, and New Guinea. Very rare in Sabah
and Sarawak. Also in Kalimantan.
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Ecology. In lowland, including limestone habitat, to lower montane forests, up to 1600 m

altitude.
Key to varieties
Leaf base distinctly auriculate; stipules persistent...............var. aurita
Leaf base not auriculate; stipules caducous...............
var. auriculifera (Merr.) Corner
(Latin, auriculiferus = bearing small ear; the leaf base)
Gard. Bull. Sing. 17 (1960) 480, l.c. (1965) 78; Anderson l.c. 257; Whitmore, Tantra & Sutisna l.c. 241. Basionym: F.
auriculifera Merr., PEB (1929) 46. Type: Elmer 20468, Borneo, Sabah, Tawau district (holotype SING).
Very rare in Sabah and Sarawak. Also in Kalimantan.
6. Ficus beccarii King

Ann. Roy. Bot. Gard. Calc. 1, 2 (1888) 102; Merrill l.c. (1921) 221; Masamune l.c. 243; Corner l.c. (1965) 92; Kochummen l.c.
141; Anderson l.c. 257; Primack l.c. 73; Whitmore, Tantra & Sutisna l.c. 241. Type: Beccari PB 2900, Borneo, Sarawak
(holotype FI).
Dioecious treelet, up to 5 m tall. Twigs c. 0.2 cm thick, fulvous appressed hairy. Stipules linear
to lanceolate, hairy, c. 3.5 cm long, persistent. Leaves distichous, thin-coriaceous, fulvous
appressed hairy below; oblong, elliptic, lanceolate, or obovate, 2538 311 cm, base cuneate,
symmetric or asymmetric, margin entire and plane or faintly toothed near the apex, apex shortly
acuminate; midrib sunken above; lateral veins 912 pairs, raised below, faint above, basal veins
12 pairs, short, departing at a broad angle from the midrib; intercostal venation scalariform,
distinct below, faint to invisible above; petiole appressed yellowish hairy, 11.5 cm long.
Syconia solitary, sessile, borne on much divided, leafless, stipule-bearing branches arising from
the base of stem, depressed globose, 0.71.7 cm diameter, with many bracts on the surface; basal
bracts 3. Achenes up to 0.7 mm long, shallowly ridged.
Vernacular names. Sarawakbuah abung (Kelabit), entimau (Iban).
Distribution. Peninsular Malaysia and Borneo. Rare in Sabah and Sarawak. Var. beccarii also
Ecology. In lowland mixed dipterocarp to lower montane forests, up to 1800 m altitude.
Key to varieties
1. Leaves 36 cm broad............var. beccarii
Leaves 511 cm broad..............2
2. Leaf base asymmetric.............
var. asymmetrica Corner
(Latin, asymmetricus = asymmetric; the leaf base)
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Gard. Bull. Sing. 18 (1960) 60, l.c. (1965) 92; Whitmore, Tantra & Sutisna l.c. 241. Type: Corner s.n., Borneo,
Sarawak, Kuching district, Bt. Kuap (holotype CGE; isotype L).
Leaves oblong, elliptic or obovate, 2628 7.511 cm; base asymmetric with one side
narrowly cuneate and the other broadly subcordate, margin entire, apex acuminate,
acumen 2.54.5 cm long.
Endemic to Borneo. Very rare, only known by the type collection from Sarawak. In
lowland mixed dipterocarp forest.
Leaf base symmetric.......
var. latifolia Corner
(Latin, latifolius = broad-leaved)
Gard. Bull. Sing. 18 (1960) 60, l.c. (1965) 92; Whitmore, Tantra & Sutisna l.c. 241; Coode et al. (eds.) l.c. 212. Type:
BRUN 5305, Brunei, Sg. Belalong (holotype CGE).
Leaves elliptic or lanceolate, 510 cm broad; base symmetric, broadly cuneate, margin
entire and plane or slightly toothed towards apex, apex acute.
Endemic to Borneo. Rare in Sabah, in lowland to lower montane forests, up to 1800 m
altitude. Also in Brunei. Not known in Sarawak and Kalimantan.
7. Ficus benjamina L.
(Sanskrit, bani; the banyan tree)
Mant. 1 (1767) 129; Blume l.c. 456; Miquel l.c. (1867) 267; King l.c. (1887) 43; Merrill l.c. (1921) 221; Ridley l.c. (1924) 336;
Burkill l.c. 1021; Masamune l.c. 243; Backer & Bakhuizen f. l.c. 24; Corner l.c. (1965) 21, l.c. (1988) 539; Kochummen l.c. 141;
Anderson l.c. 257; Primack l.c. 74; Whitmore, Tantra & Sutisna l.c. 241; Argent et al. (eds.) l.c. 439. Type: Linn. Herb. (BM?).
Synonyms: F. nitida Thunb., Diss. Fic. (1785) 5; F. haematocarpa Blume ex Decne., Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834)
494; F. neglecta Blume ex Decne l.c. 494; Urostigma benjaminum (L.) Miq. in Hooker, Lond. J. Bot. 6 (1847) 583; U.
haematocarpum (Blume ex Decne.) Miq. l.c. (1847) 584; U. nitidum (Thunb.) Miq. l.c. (1847) 582; U. neglectum (Blume ex
Decne.) Miq., Fl. Ned. Ind. 1, 2 (1859) 347.
Monoecious tree or strangler, up to 30 m tall, 40 cm diameter, with few free hanging aerial roots
and drooping branches. Bark light grey. All parts glabrous. Twigs up to 0.2 cm thick, dark
brown, reddish brown when young, angled. Stipules paired, broadly lanceolate, acute, c. 1 cm
long, caducous. Leaves distichous, thin-coriaceous, withering yellow; elliptic to ovate-lanceolate,
48 1.83.8 cm, base cuneate, margin entire, plane, apex acute; midrib raised above, drying
yellowish on both surfaces; lateral veins numerous, closely spaced, almost parallel, visible on
both surfaces, joining near the leaf margin to form distinct intramarginal vein, basal pair short;
intercostal venation reticulate, equally prominent as lateral veins; petiole 0.51.5 cm long,
wrinkled and yellowish on drying. Syconia axillary, paired, sessile, ripening orange-yellow,
globose, c. 0.8 cm diameter, apex flat, drying pale yellowish with rugose surface; basal bracts 2
3, 0.51.5 mm long, concealed beneath the syconia; ostiole covered by 3 small apical bracts;
internal bristles none. Tepals 34, free. Achenes ovoid to reniform, longer than the style.
Vernacular names. Sarawakberingin, waringin (Malay), dunuk (Bidayuh).
Distribution. India, S China, Malesia, Solomon Islands, and N Australia. Common in Sabah and
Ecology. In lowland and hill forests, up to 600 m altitude, usually by streams; often planted in
urban areas.
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8. Ficus bruneiensis Corner
Fig. 6.
(of Brunei)
Gard. Bull. Sing. 17 (1960) 435, l.c. (1965) 48; Anderson l.c. 257; Primack l.c. 75; Whitmore, Tantra &
Sutisna l.c. 242; Coode et al. (eds.) l.c. 212. Type: Corner BRUN 5383, Borneo, Brunei, Amo, Ulu
Belalong (holotype CGE; isotype L).
Dioecious tree, up to 20 m tall, 45 cm diameter. Twigs glabrous, hollow, 0.81.5 cm

thick, reddish brown, shallowly ridged, with prominent stipular scars. Stipules lanceolate,
acute, 3.55.2 cm long, minutely appressed silky hairy, caducous. Leaves spirally arranged,
coriaceous, rough-hairy above, yellowish brown soft-hairy below; elliptic, ovate or oblong,
2037 1124 cm, base rounded or subcordate (distinctly cordate in young trees), margin
toothed, apex acute; midrib raised above; lateral veins 5 pairs, prominently raised below,
faint above, basal pair reaching more than 1/2 the length of the blade, with a gland
in each axil below; intercostal venation reticulate to scalariform, prominent below, faint
above; petiole 5.511 cm long, sparsely long-hairy. Syconia axillary, paired, sessile,
ripening red, oblong or ellipsoid, 23.5 2.53 cm, covered with rough long yellow
hairs; basal bracts 3, ovate, 610 mm long, caducous; internal bristles numerous, yellowish.
Tepals 34, yellowish to reddish, almost linear, densely long-hairy. Achenes oblong,
smooth, ridged on the side where style comes out.
Vernacular names. Sarawakkempan, kumpan, tempan (Iban).
Distribution. Endemic to Borneo. Common in Sarawak and known from Belaga, Kapit,
and Lubok Antu districts; not yet reported for Sabah. Also in Brunei.
Ecology. In lowland and hill mixed dipterocarp forests, up to 500 m altitude, usually
by streams.
9. Ficus brunneoaurata Corner

(Latin, brunneus = brown, auratus = flecked with gold; the yellowish brown indumentum)
Sutisna l.c. 242; Coode et al. (eds.) l.c. 212. Type: SFN 26608, Borneo, Sabah, Mt. Kinabalu, Menetendok
Gorge (holotype SING).
Dioecious tree, up to 15 m tall, 20 cm diameter. Twigs, petioles and leaf undersurface

yellowish brown hairy. Twigs 0.31 cm thick, dark brown, striate, with distinct stipular
scars. Stipules lanceolate, acute, 13.5 cm long, velvety hairy. Leaves spirally arranged,
coriaceous; ovate-elliptic, 1530 614 cm (lobed and up to 50 14 cm in juvenile
trees), base subcordate to cuneate, margin faintly toothed, apex acute; midrib raised above;
lateral veins 45 pairs, raised on both surfaces, with glands in the axils of lateral veins
below, basal veins 23 pairs, reaching more than 1/2 the length of the blade; intercostal
venation scalariform, visible on both surfaces; petiole 1.58 cm long. Syconia axillary
or borne on twigs behind the leaves, sessile, ripening red, short velvety yellowish brown
hairy, subglobose, 11.8 cm diameter; basal bracts 11.5 mm long; internal bristles
numerous, brownish. Tepals 34, reddish, densely hairy. Achenes keeled and tuberculate.
263
Vernacular names. Sabahtegong (Dusun). Sarawakkupan, tempan (Iban).

Distribution. Endemic to Borneo. Common in Sabah especially on Mt. Kinabalu, frequent
in Sarawak. Also in Brunei and Kalimantan.
Ecology. In lowland and hill mixed dipterocarp forests, up to 1100 m altitude, including
secondary forest.
10. Ficus callosa Willd.
Fig. 7.
(Latin, callosus = bearing hardened thickenings; probably referring to the syconia-bearing

branches and stem or to the densely sclerotic syconium wall)
Act. Acad. Berol. (1798) 102; Miquel l.c. (1867) 295; King l.c. (1887) 64; Merrill l.c. (1921) 221;
Masamune l.c. 243; Backer & Bakhuizen f. l.c. 32; Corner l.c. (1965) 29, l.c. (1988) 546; Kochummen
l.c. 142; Anderson l.c. 257; Primack l.c. 77; Whitmore, Tantra & Sutisna l.c. 242; Turner l.c. 349. Type:
unknown. Synonyms: F. scleroptera Miq., Pl. Jungh. (1851) 63; F. basidentula Miq. l.c. (1859) 314; F.
malunuensis Warb. in Perkin, Fragm. Fl. Philip. 3 (1905) 196; F. cordatifolia Elmer, Leaft. Philip. Bot.
4 (1911) 1250; F. longispathulata Sata, Monogr. (1944) 323.
Monoecious tree, up to 33 m tall, 90 cm diameter, without free hanging aerial roots;

buttresses up to 3 m tall. Bark pale grey to dark brown, smooth; inner bark pale orangebrown; sap watery, white. Sapwood orange to cream. Twigs 0.30.8 cm thick, yellowish
with dark brown lenticels. Stipules 1.23 cm long, caducous. Leaves spirally arranged,
coriaceous, glabrous; elliptic or oblong, 1427 5.514 cm, base symmetric, rounded
or broadly cuneate, margin entire, plane, apex rounded or obtuse; midrib raised above;
lateral veins 710 pairs, well-spaced, curved, departing from the midrib at a broad angle,
distinct on both surfaces, basal veins 23 pairs, distinct, reaching up to 1/3 the length
of the blade, without glands in the axils below; intercostal venation reticulate, visible
on both surfaces; petiole 3.57 cm long, drying yellowish. Syconia axillary or borne
on the twigs behind the leaves, ripening yellow, finely villous, glabrescent, pear-shaped,
1.52.5 1.22 cm, apex with tiny ostiole; peduncle slender, 23 cm long; basal bracts
3, 1.54 mm long, persistent; internal bristles none. Tepals 36, red, free, glabrous. Achenes
oblong, c. 1.5 mm long, slightly keeled.
Vernacular name. Sabahkayu ara (Malay).
Distribution. India, Sri Lanka, Andaman Islands, Burma, Indo-China, Thailand, Sumatra,
Peninsular Malaysia, Java, Borneo, the Philippines, Sulawesi, Maluku, and Lesser Sunda
Islands. Widely distributed in Sabah and Sarawak but nowhere abundant; often seen in
urban areas. Also in Kalimantan.
Ecology. In lowland forest, especially in coastal areas.
11. Ficus cereicarpa Corner

(Greek, ceras = with horn-like projection, karpos = fruit; the lateral bracts on the syconia)
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Sutisna l.c. 243; Coode et al. (eds.) l.c. 212. Type: Corner BRUN 5315, Borneo, Brunei, Amo, Sg.
Belalong (holotype CGE; isotype SAR).
Dioecious tree, up to 9 m tall. Twigs 0.41.8 cm thick, reddish brown, shallowly ridged,
with prominent stipular scars; young parts covered with white, silky, appressed or spreading
hairs. Stipules lanceolate, 1.75 cm long, appressed velvety hairy, subpersistent. Leaves
spirally arranged, thin-coriaceous, sparsely yellowish brown hairy below, especially on
midrib and lateral veins; elliptic, ovate or obovate, 1343 6.526 cm, base subcordate
or cordate, margin entire, plane, apex acute; midrib raised above; lateral veins 820
pairs, distinct on both surfaces, basal veins 24 pairs, short, departing at a broad angle
from the midrib, without glands in the axils below; intercostal venation scalariform to
reticulate, distinctly visible below; petiole 213.5 cm long, glabrous or hairy, wrinkled
on drying. Syconia borne in dense clusters on stout leafless branching twigs up to 30
cm long arising from the base of trunk, ripening brown to pink, finely hairy, pear-shaped,
2.53 cm long; peduncle 23 cm long; basal bracts 3, lateral bracts horn-like, thick and
many, apical bracts lanceolate, thick, many; internal bristles numerous, flexuous. Tepals
fused into a cup-shaped or tubular perianth, reddish. Achenes c. 1 mm long, rough to
smooth, slightly keeled.
Vernacular name. Sarawaksampandai (Iban).
Distribution. Endemic to Borneo. Frequent in Sabah and Sarawak. Also in Brunei and
Kalimantan.
Ecology. In lowland mixed dipterocarp to lower montane forests, up to 1600 m altitude,
usually by streams.
Key to varieties
Twig slender, 0.40.9 cm thick. Leaf base subcordate; hairs sparse, up to 4 mm long.
Syconia with peduncle up to 1 cm long.....................var. cereicarpa
Twig stout, up to 1.8 cm thick. Leaf base distinctly cordate; hairs dense, longer than
4 mm. Syconia with peduncle up to 3 cm long.
var. ashtonii Kochummen
Gard. Bull. Sing. 50 (1998) 213. Type: Ashton S. 17806, Borneo, Sarawak, Kapit district, Batang
Baleh, Ulu Selentang (holotype KEP; isotypes A, BO, CGE, K, L, SAR, SING).
Differs from the typical variety in the stouter twigs, very large cordate leaves which
are at least 20 cm broad, and in the long peduncled syconia.
Endemic to Borneo; recorded only for Sarawak by 3 collections, S. 45512 from Belaga
district, S. 40302 from Miri district (Lambir Hill NP), and the type from Batang
Baleh in Kapit district.
12. Ficus chaii Kochummen

(Paul P.K. Chai 1941, sometime Forest Botanist at the Herbarium, Forestry Department of
Sarawak)
Gard. Bull. Sing. 50 (1998) 203. Type: Chai S. 36002, Borneo, Sarawak, Kapit district, Sg. Kapit
(holotype KEP; isotypes SAN, SAR).
266
vi
Treelet. Twigs reddish brown, angled. Stipules lanceolate, up to 1 cm long, finely hairy
outside, caducous. Leaves spirally arranged, thin-coriaceous, glabrous, drying greenish
yellow; narrowly oblong, 13.523.5 1.52.2 cm, base cuneate, margin distantly toothed,
apex acuminate, acumen 12 cm long; midrib raised above; lateral veins 1321 pairs,
distinctly curving and joining near the leaf margin, visible below, very faint above, basal
pairs short; intercostal venation reticulate, visible below; petiole 11.5 cm long, yellowish
on drying. Syconia axillary, solitary or in pairs, ripening orange, pear-shaped, 0.50.8
0.30.4 cm; peduncle to 0.3 cm long; basal bracts tiny. Tepals 3, lanceolate, transparent.
Achenes not seen.
Distribution. Endemic to Borneo. Known only by the type specimens from Sarawak.
Ecology. In lowland mixed dipterocarp forest by stream.
13. Ficus chartacea (Wall. ex Kurz) King

(Latin, chartaceus = papery; the leaves)
Ann. Roy. Bot. Gard. Calc. 1, 2 (1888) 159; Ridley l.c. (1924) 350; Burkill l.c. 1022; Corner l.c. (1965)
47, l.c. (1988) 547; Kochummen l.c. 142; Anderson l.c. 257; Primack l.c. 78; Whitmore, Tantra &
Sutisna l.c. 243; Turner l.c. 349; Coode et al. (eds.) l.c. 213. Type: Wall. Cat. 4580, Peninsular Malaysia,
Penang (K?). Synonym: F. lamponga Miq. var. chartacea Wall. ex Kurz, Fl. Burma 2 (1877) 451.
Dioecious shrub or small tree, up to 12 m tall, without free hanging aerial roots. Bark
greyish white to dull brown. Young twigs and petioles pink. Twigs up to 0.2 cm thick.
Stipules lanceolate, 0.31 cm long, glabrous, caducous. Leaves spirally arranged, thincoriaceous, glabrous, with cream white or yellow dots on the upper surface; elliptic, 7.5
15 2.56 cm, base symmetric, cuneate, margin entire, plane, apex acute; midrib raised
above; lateral veins 46 pairs, well-spaced, curved, departing from the midrib at a broad
angle, distinct on both surfaces, basal pair distinct, extending up to 1/2 the length of
the blade, with a gland in each axil below; intercostal venation reticulate, visible on
both surfaces; petiole 13 cm long, slender. Syconia sessile or with short peduncle, in
pairs or clustered, axillary or borne on twigs behind the leaves, ripening yellow or orangebrown and then scarlet, globose, 0.60.8 cm diameter, apex umbonate; basal bracts 3,
tiny; internal bristles few, small. Tepals 34, free. Achenes ovoid, c. 1 mm long, tuberculate,
rather strongly keeled.
Distribution. Burma, Indo-China, Thailand, Peninsular Malaysia, and Borneo. Common
in Sabah, not yet reported for Sarawak. Also in Brunei and Kalimantan.
Ecology. In lowland forest, often by streams.
14. Ficus cumingii Miq.

(H. Cuming, 17911865, English traveller and naturalist; collected a large number of plant
specimens in S America, the Pacific Islands, the Philippines, Singapore, Peninsular Malaysia
and Sumatra)
268
In Hooker, Lond. J. Bot. 7 (1848) 235; King l.c. (1888) 92; Corner l.c. (1965) 66. Type: Cuming 1925,
the Philippines (K).
Dioecious shrub, up to 4 cm tall; latex white. Young shoots, petioles, peduncles and
undersurface of leaves rough-hairy. Twigs up to 0.2 cm thick, pale brown. Stipules paired,
up to 1 cm long, caducous. Leaves subopposite, scabrous; narrowly lanceolate, 630
1.25 cm, base cuneate to rounded, margin distantly faintly lobed or toothed, apex
obtuse; midrib raised above; lateral veins 1425 pairs, faintly raised below, basal pair
short, departing at a broad angle from the midrib, with 12 glands in the axils below;
intercostal venation reticulate; petiole c. 0.8 cm long. Syconia axillary, ripening yellow
to red, globose, c. 0.6 cm diameter, scabrous; peduncle 0.10.4 cm long; basal bracts
3; internal bristles numerous, short. Tepals 35, free. Achenes lenticular, up to 1.2 mm
long, smooth, slightly keeled.
Distribution. Formosa, Borneo, the Philippines, Sulawesi, and New Guinea. One variety
is present in Sabah.
Ecology. In lowland forest, including rocky coastal forest.
var. terminalifolia (Elmer) Sata
(having leaves resembling those of Terminalia, Combretaceae)
Monogr. (1944) 239; Corner l.c. (1965) 66; Whitmore, Tantra & Sutisna l.c. 243. Basionym: F.
terminalifolia Elmer, Leafl. Philip. Bot. 4 (1911) 318. Type: unknown. Synonym: F. producta Merr.,
En. Philip. Pl. 2 (1923) 62.
Young twigs and leaves covered with rough hairs; leaf margin faintly lobed or toothed.
Syconia axillary, globose, sessile, rough-hairy.
Formosa, Borneo, and the Philippines. Very rare in Borneo, known by 3 collections, SAN
103482, SAN 11158 and Kamis 4157 from Tawau district in Sabah.
15. Ficus deltoidea Jack
Fig. 8.
(Latin, deltoideus = deltoid, equilateral-triangular; the leaf shape)

Mal. Misc. 2 (1822) 71; Corner l.c. (1965) 37, Phil. Trans. Roy. Soc., London Ser. B, 256 (1969) 281,
l.c. (1988) 554; Backer & Bakhuizen f. l.c. 28; Kochummen l.c. 145; Primack l.c. 81; Whitmore, Tantra
& Sutisna l.c. 244; Turner l.c. 349; Coode et al. (eds.) l.c. 213. Type: Jack s.n., Sumatra (BM). Synonyms:
F. diversifolia Blume l.c. 456; F. diversifolia Blume var. latissima Miq. l.c. (1867) 268; F. diversifolia
Blume var. deltoidea (Jack) Ridl. l.c. (1924) 346.
Dioecious shrub, small tree or epiphyte, up to 3 m tall. Twigs up to 0.2 cm thick. Stipules
lanceolate, 0.151.5 cm long, caducous. Leaves spirally arranged, thin- to thick-coriaceous,
glabrous; deltoid, elliptic, obovate, or spathulate or rhomboid, 216.5 1.510 cm, base
tapered or cuneate, margin entire or lobed, revolute or plane, apex rounded, notched,
obtuse or acute; midrib usually forked near the middle and with a gland in the fork
or not forked but with a gland in the axils of basal lateral veins below, raised, flat
or sunken above; lateral veins up to 10 pairs, curving and joining near the leaf margin,
distinct or sometimes indistinct below, faint above; intercostal venation reticulate, equally
269
prominent as the lateral veins or not prominent or indistinct on both surfaces; petiole
0.23 cm long. Syconia in pairs or solitary, axillary, ripening yellow or red, oblong,
globose or ellipsoid, 0.32 0.21.2 cm, strongly umbonate at apex, glabrous or hairy;
peduncle 0.31.5 cm long or syconia almost sessile; basal bracts 3, c. 1 mm long, persistent;
internal bristles few, short. Tepals 34, red, glabrous, ovate to lanceolate, free. Achenes
oblong to kidney-shaped, 35 2 mm, with few faint lines or smooth.
Vernacular names. Sarawakara, kara (Iban), lunok (Murut).
Distribution. Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, and the Philippines.
Common and widely distributed in Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. In lowland, including peatswamp, kerangas and limestone to upper montane
Uses. A reputed traditional medicine. A decoction of leaves is taken after childbirth as
a tonic drink.
Notes. A variable species with 15 recognised varieties (cf. Corner l.c. 1969). Eight of
the varieties occur in Sabah and Sarawak.
Key to varieties
(The epiphytic varieties are annotated under the main key: leads 166 and 168)
1.
Leaf penninerved, apex acute; midrib not forked.............2

Leaf usually with one or 2 pairs of veins, rarely more, apex rounded; midrib forked..3
2.
Petiole up to 0.2 cm long. Midrib flat above. Common small trees of kerangas forest.
var. arenaria Corner (in part)
(Latin, arenarius = growing on sandy habitat)
Gard. Bull. Sing. 17 (1960) 421, l.c. (1965) 37, l.c. (1969) 302; Whitmore, Tantra & Sutisna
l.c. 244; Coode et al. (eds.) l.c. 213. Type: BRUN 5352, Borneo, Brunei, Kuala Belait (holotype
CGE).
Shrub up to 2 m tall. Leaves elliptic to narrowly obovate, 2.511 1.84 cm,

base cuneate, margin lobed, apex acute or rounded; midrib not forked or rarely
forked in some leaves towards the apex of the twig; lateral veins 35 pairs;
petiole up to 0.2 cm long. Syconia ellipsoid, 11.2 0.70.9 cm; peduncle 0.1
0.9 cm long.
Endemic to Borneo. Common in Sabah and Sarawak. Lowland kerangas forest.
Petiole very much longer than 0.2 cm. Midrib sunken above. Small trees of other
habitats.....
var. intermedia Corner
(Latin, intermedius = intermediate between var. lutescens and var. motleyana)
Gard. Bull. Sing. 17 (1960) 423, l.c. (1965) 37, l.c. (1969) 304; Kochummen l.c. 145; Whitmore,
Tantra & Sutisna l.c. 244. Type: Purseglove 4281, Peninsular Malaysia, Pahang, Frasers Hills
(holotype SING). Synonyms: F. kinabaluensis Stapf, FMK (1894) 226; F. burkillii Ridl. l.c.
(1924) 330.
Shrub up to 3 m tall. Leaves elliptic to spathulate, 310 1.55.5 cm, base cuneate,
270
vi
margin entire, plane, apex obtuse or rounded; midrib not forked, except where leaves
have rounded apex, sunken above; lateral veins 45 pairs, visible above, with a gland
in each axil of the basal pair; petiole 0.21.5 cm long. Syconia subglobose; peduncle
up to 1 cm long.
Peninsular Malaysia and Borneo. Common on Mt. Kinabalu in Sabah, not yet
reported for Sarawak. In lowland to hill forests, up to 1000 m altitude.
3.
Leaves deltoid, equally broad and long; midrib forking below the middle of the
blade......var. deltoidea
Leaves narrowly obovate, longer than broad; midrib usually forked above the middle
of the blade................4
4.
Petiole up to 0.2 cm long.........var. arenaria (in part)

Petiole at least 0.5 cm long...............5
5.
Leaves up to 3.5 cm broad. Syconia glabrous.....................

var. angustifolia (Miq.) Corner
(Latin, angustifolius = narrow-leaved)
Gard. Bull. Sing. 17 (1960) 420, l.c. (1965) 37, l.c. (1969) 300; Kochummen l.c. 145; Whitmore,
Tantra & Sutisna l.c. 244; Coode et al. (eds.) l.c. 213. Basionym: Synoecia diversifolia (Blume)
Miq. var. angustifolia Miq. l.c. (1851) 67. Type: unknown. Synonyms: F. ovoidea Jack l.c. 71;
F. spathulata Miq. l.c. (1848) 441; F. retusa L. var. ovoidea (Jack) Miq. l.c. (1867) 288; F.
diversifolia (Blume) Miq. var. ovoidea (Jack) King l.c. (1888) 140.
Lower Thailand, Sumatra, Peninsular Malaysia, and Borneo. Common in Sarawak,

not yet reported for Sabah. Also in Brunei and Kalimantan. In hill to lower montane
forests, at 8001300 m altitude, including limestone habitat.
Leaves more than 4 cm broad. Syconia hairy.........
var. borneensis Corner
(of Borneo)
Gard. Bull. Sing. 17 (1959) 422, l.c. (1965) 37, l.c. (1969) 303; Whitmore, Tantra & Sutisna
l.c. 244; Coode et al. (eds.) l.c. 214. Type: Endert 3920, Borneo, Kalimantan, W Kutai, Mt.
Kemul (holotype L).
Twigs 0.20.5 cm thick. Leaves obovate or oblong, 4.513 39 cm, base cuneate
to rounded, margin entire, revolute, apex rounded; midrib raised or sunken above,
forked from near the middle of the blade; lateral vein one pair; petiole 0.5
2.5 cm long. Syconia glabrous or hairy, ovoid or oblong, 0.81.2 cm diameter;
peduncle up to 0.6 cm long.
Endemic to Borneo. Frequent in Sarawak; very rare in Sabah, known by 2
collections, SAN 25867 and SAN 66909, from Beaufort district. In lowland mixed
dipterocarp to lower montane forests, up to 1500 m altitude.
16. Ficus diamantiphylla Corner

(Greek, diamante = decorated with sparkling spots, phullon = leaf)
Phil. Trans. Roy. Soc. London Ser. B, 259 (1970) 368; Anderson l.c. 258; Primack l.c. 83; Whitmore,
Tantra & Sutisna l.c. 244. Type: S. 22791, Borneo, Sarawak, Marudi district, Ulu Tutoh (holotype
SAR).
272
Dioecious tree up to 6 m high. Twigs hollow, finely ridged with raised lines, c. 1.2 cm
thick, long-woolly hairy. Leaves spirally arranged, rough-hairy on both surfaces, drying
yellowish; obovate, 2041.5 1320 cm, base subcordate, margin finely toothed, apex
acute; midrib sunken above, densely hairy; lateral veins 78 pairs, prominently raised
below, visible above; intercostal venation reticulate, raised below; petiole 77.5 cm long,
woolly hairy. Syconia yellow, ripening red, densely tomentose.
Distribution. Endemic to Borneo. Very rare, known only by the type collection from
Sarawak.
Ecology. In lowland mixed dipterocarp forest by stream.
17. Ficus endospermifolia Corner
Fig. 9.
(having leaves like that of Endospermum, Euphorbiaceae)

Gard. Bull. Sing. 17 (1960) 434, l.c. (1965) 48; Whitmore, Tantra & Sutisna l.c. 245. Type: SAN A
4492, Borneo, Sabah, Mt. Kinabalu, Tenompok (holotype K).
Dioecious tree, up to 25 m tall. Twigs 0.40.7 cm thick, brown hairy, strongly grooved,
with prominent petiolar and stipular scars. Stipules ovate, acute, 13 cm long, velvety
yellowish hairy. Leaves spirally arranged, coriaceous, densely velvety hairy on both
surfaces, denser below; broadly ovate, 1325 cm long and almost equally broad, base
truncate to cordate, margin faintly toothed, apex acute; midrib raised above; lateral veins
45 pairs, departing from near the base of midrib, strongly raised below, visible above,
basal pair reaching almost 1/2 the length of the blade, with a gland in each axil below;
intercostal venation scalariform, raised below, faint to invisible above; petiole 3.510 cm
long, yellowish, hairy. Syconia axillary, sessile, velvety hairy, ovoid to subglobose, 1.8
2.5 1.52 cm; basal bracts ovate, 35 mm long, caducous; internal bristles numerous,
yellow. Tepals 34, reddish, glabrous. Achenes oblong, 11.3 mm long, keeled, tuberculate.
Distribution. Endemic to Borneo, common in Mt. Kinabalu area in Sabah; very rare
in Sarawak, known by 2 collections S. 17688 from Ulu Anap, Tatau district, and S. 47783
from G. Pagon Periok, Limbang district. Not known in Brunei and Kalimantan.
Ecology. In hill mixed dipterocarp to lower montane forests, at 10001800 m altitude.
18. Ficus eumorpha Corner

(Greek, eu = good, morphus = shape; alluding to the shape of the leaves)
Sutisna l.c. 245. Type: SFN 27566, Borneo, Sabah, Tibobah R. (holotype SING).
Dioecious tree, up to 18 m tall. Twigs, leaves and syconia yellowish hairy. Twigs 0.4
0.5 cm thick. Stipules up to 1.5 cm long, hairy. Leaves spirally arranged, coriaceous,
yellowish soft-hairy below, upper surface rough to the touch; ovate, 820 6.514 cm,
base cordate or truncate, margin distinctly toothed, apex acute; midrib raised above;
lateral veins 57 pairs, raised on both surfaces, basal pair reaching almost 1/2 the length
273
of the blade, with a gland in each axil below; intercostal venation scalariform, raised
below, faint above; petiole 48.5 cm long, powdery yellow-hairy. Syconia axillary, sessile,
subglobose to ellipsoid, 1.61.7 1.41.5 cm, covered with long hairs, ripening yellowish
green; basal bracts 34 mm long; internal bristles numerous, 12 mm long. Tepals 3,
white, densely hairy. Achenes oblong, 11.2 mm long, slightly keeled, subtuberculate.
Vernacular name. Sarawaktempan (Iban).
Distribution. Endemic to Borneo. Rare in Sabah (Mt. Kinabalu) and Sarawak (Mt. Mulu)
but locally frequent. Also in Brunei and Kalimantan.
Ecology. In lower montane forest, at 13001500 m altitude.
19. Ficus fistulosa Reinw. ex Blume

(Latin, fistulosus = hollow throughout; the young twigs)
Bijdr. Fl. Ned. Ind. (1825) 470; King l.c. (1888) 114; Merrill l.c. (1921) 223; Ridley l.c. (1924) 343;
Burkill l.c. 1025; Backer & Bakhuizen f. l.c. 32; Corner l.c. (1965) 93; Kochummen l.c. 146; Anderson
l.c. 258; Primack l.c. 85; Whitmore, Tantra & Sutisna l.c. 245; Coode et al. (eds.) l.c. 214. Type: Reinwardt
s.n. (= RHL Sheet No. 9081881231), Java (holotype L). Synonym: F. condensa King l.c. (1888) 114.
Dioecious tree, up to 18 m, 20 cm diameter. Bark grey; inner bark yellowish white;

latex pale yellowish buff. Twigs 0.30.8 cm thick, hollow. Stipules up to 1.5 cm long,
caducous. Leaves spirally arranged or opposite, glabrous, coriaceous; ovate to lanceolate
or elliptic to oblong, 825 2.512 cm, base rounded, sometimes asymmetric, margin
entire, revolute, apex acute; midrib raised above; lateral veins 49 pairs, raised below,
faint above, basal pair short; intercostal venation reticulate or subscalariform, visible on
both surfaces; petiole 0.87 cm long, with subnodal gland. Syconia in clusters, axillary
or borne on stem and branches, depressed globose, 1.52.5 cm diameter, ripening greenish
yellow to pale yellow, base narrowed into a short stalk-like structure; peduncle 11.5
cm long; basal bracts 3, subtriangular, up to 1 mm long, persistent; internal bristles none.
Tepals 2 or 3, fused, tubular or collar-like. Achenes minutely tuberculate.
Vernacular names. Sarawakengkunoh, engkuroh, kara (Iban).
Distribution. Assam, Bengal, Burma, S China, Indo-China, Formosa, Hainan, Nicobar
Islands, Thailand, Sumatra, Peninsular Malaysia, Borneo, the Philippines, Java, and the
Lesser Sunda Islands. Common and widely distributed in Sabah and Sarawak. Also in
Ecology. In lowland to lower montane forests, up to 1500 m altitude, often by streams.
Key to varieties
Syconia borne on stem and branches. Leaves at least 5 cm wide. Generally in lowland
forest.........................var. fistulosa
274
vi
Syconia axillary. Leaves 23 cm broad. In hill and montane forests...........

var. tengerensis (Miq.) Kuntze
(of Mt. Tengger, E Java)
Rev. Gen. Pl. 1 626. Basionym: F. tengerensis Miq. l.c. (1867) 283.
Thailand, Sumatra, Peninsular Malaysia, Borneo (Brunei, Kalimantan, Sabah, and

Sarawak), and Java.
20. Ficus francisci H. Winkl.

(Franz Winkler, brother of H. Winkler)
In Engler, Bot. Jahrb. 49 (1913) 362; Corner l.c. (1965) 91; Anderson l.c. 258; Primack l.c. 86; Whitmore,
Tantra & Sutisna l.c. 246; Coode et al. (eds.) l.c. 214. Type: unknown.
Dioecious tree, up to 8 m tall, 10 cm diameter. Twigs c. 0.4 cm thick, solid, grey-brown,

striate, with distinct stipular and petiolar scars. Stipules lanceolate, c. 1 cm long, woolly
white-hairy, caducous. Leaves spirally arranged, thin-coriaceous, drying greenish, silky
hairy on the midrib; elliptic to narrowly obovate, 7.521.5 26 cm, base cuneate to
attenuate, margin entire, plane, apex acuminate, acumen 1.5 cm long; midrib flat above;
lateral veins 811 pairs, raised below, faint above, basal pair very short, with a gland
in each axil below; intercostal venation scalariform; petiole 1.54.5 cm long, sparsely
hairy. Syconia borne on finger-like leafless branches arising from stem, densely or sparsely
woolly pubescent, glabrescent, obovoid, c. 2 1 cm (immature), surface strongly ridged,
apex umbonate with constriction at the base; peduncle c. 0.7 cm long; basal bracts 3,
glabrous, persistent; internal bristles numerous, white to pale fawn. Perianth tubular.
Achenes lenticular, up to 0.9 mm long, smooth, slightly keeled.
Vernacular name. Sarawakentimau (Iban).
Distribution. Endemic to Borneo; rare in Sabah and Sarawak. Also in Brunei and
Kalimantan.
Ecology. In lowland mixed dipterocarp forest on alluvial soils.
21. Ficus fulva Reinw. ex Blume

(Latin, fulvus = tawny; the indumentum)
Bijdr. Fl. Ned. Ind. (1825) 478; Miq. l.c. (1867) 269; King l.c. (1888) 148; Merrill l.c. (1921) 223;
Ridley l.c. (1924) 341; Burkill l.c. 1026; Backer & Bakhuizen f. l.c. 30; Corner l.c. (1965) 46, l.c. (1988)
547; Kochummen l.c. 147; Anderson l.c. 258; Primack l.c. 87; Whitmore, Tantra & Sutisna l.c. 246;
Coode et al. (eds.) l.c. 214. Type: Blume 44 (= RHL Sheet No. 925250805), Java (holotype L). Synonyms:
F. chrysocarpa Reinw. ex Blume l.c. 475; F. reinwardtii Link & Otto, Ic. Pl. Rar. 1 (1828) 6; F. apiculata
Miq. l.c. (1854) 92; F. suborbicularis Miq. l.c. (1861) 425; F. flavidula Miq. l.c. (1867) 270; F. discolor
Miq. l.c. (1867) 221; F. patens Ridl., J. Str. Br. R. As. Soc. 86 (1922) 306.
Dioecious small tree, rarely up to 20 m tall and 20 cm diameter. Bark brown; inner
bark yellowish. Twigs 0.30.8 cm thick, stiff-hairy. Stipules lanceolate, c. 1 cm long,
276
acute, pale hairy outside. Leaves spirally arranged, thin-coriaceous, densely velvety
yellowish hairy below, sparsely hairy above; ovate, elliptic or obovate, 927 4.516
cm, base symmetric, usually cuneate, margin faintly toothed, apex shortly acute; midrib
flat above; lateral veins 46 pairs, raised below, faint above, basal pair reaching up to
2/3 the length of the blade, with a gland in each axil below; intercostal venation scalariform,
raised below, faint above; petiole 1.55 cm long, sparsely yellowish hairy. Syconia axillary,
in pairs, velvety yellowish hairy, ovoid to globose, 0.81 cm diameter, apex umbonate;
peduncle up to 0.5 cm long; basal bracts ovate to rounded, 23.5 mm long, persistent,
lateral bracts absent; internal bristles numerous, up to 1 mm long. Tepals 35, oblong,
glabrous. Achenes up to 1 mm long, minutely tuberculate.
Vernacular names. Sarawaklengan, tempan (Iban).
Distribution. Nicobar Islands, Thailand, Sumatra, Peninsular Malaysia, Java, Borneo,
Sulawesi, Maluku, Sumbawa, and Timor. Very common in Sabah and Sarawak. Also in
Ecology. In lowland primary, secondary and degraded forests.
22. Ficus geocharis Corner

(Greek, geos = earth or ground, charis = enchanting; the syconia)
Sutisna l.c. 246; Coode et al. (eds.) l.c. 214. Type: Clemens 21912, Borneo, Sarawak (holotype BM).
Dioecious tree, up to 13 m tall. Twigs, petioles, leaf undersurface and syconia yellowish
brown hairy. Twigs c. 0.3 cm thick, dark brown. Stipules lanceolate, 1.53.8 cm long,
persistent. Leaves distichous, membranous; elliptic to oblong or lanceolate, 1539 5
9 cm, base asymmetric, with one side narrowly cuneate and the other narrowly auriculate,
margin finely toothed, apex abruptly acuminate, acumen 26 cm long; midrib raised above;
lateral veins 68(10) pairs, distal ones with axillary glands, basal 14 pairs short, departing
from near the base of midrib; intercostal venation finely raised below; petiole 0.40.9
cm long, with subnodal gland. Syconia borne on stolons arising from the base of stem
and also on leafless twigs arising from branches and stems, ripening purplish, densely
yellowish or reddish brown hairy, depressed globose to turbinate, 1.62 cm diameter;
peduncle 0.150.4 cm long; lateral bracts lanceolate, up to 9 mm long; internal bristles
none. Perianth rudimentary or absent. Achenes lenticular, up to 0.9 mm long, smooth.
Vernacular names. Sarawakbuah abung (Kelabit), entimau, timopada (Iban).
Distribution. Endemic to Borneo (Sabah, Sarawak, Brunei, and Kalimantan). In Sabah
and Sarawak rare; recorded from Sabah by 1 collection SAN 2864 from Sandakan district
(Betotan), and from Sarawak by 4 collections (including the type) from Bau, Kuching,
Miri (G. Mulu NP), and Belaga (Upper Rajang) districts.
Ecology. In lowland mixed dipterocarp forest, including limestone habitat.
277
23. Ficus glandulifera (Wall. ex Miq.) King
Fig. 10.
(Latin, glanduliferus = gland-bearing; the petiole)

Ann. Roy. Bot. Gard. Calc. 1, 2 (1888) 143; Ridley l.c. (1924) 347; Backer & Bakhuizen f. l.c. 29;
Corner l.c. (1965) 47; Kochummen l.c. 147; Anderson l.c. 258; Primack l.c. 88; Whitmore, Tantra &
Sutisna l.c. 246; Coode et al. (eds.) l.c. 215. Basionym: Pogonotrophe glandulifera Wall. ex Miq. l.c.
(1848) 77. Type: Wall. Cat. 4481 (K). Synonyms: P. aurantiaca Miq. in Zollinger, Syst. Verz. 2 (1854)
93; P. sumatrana Miq., Fl. Ind. Bat. Suppl. (1861) 436; F. aurantiaca (Miq.) Miq. l.c. (1867) 293; F.
hasskarlii Merr., Philip. J. Sc. 11 (1916) Bot. 264; F. henschelii Merr. l.c. (1916) 264.
Dioecious tree, up to 30 m tall, 40 cm diameter; buttresses up to 1.5 m high. Bark

dark brown, smooth and lenticellate; inner bark pale yellow. Sapwood yellowish. Twigs
0.20.4 cm thick, reddish brown. Stipules ovate, c. 0.5 cm long, densely reddish brown
hairy. Leaves spirally arranged, membranous, densely or sparsely brownish hairy below,
very sparsely hairy on midrib and veins above; elliptic to narrowly obovate, (5)815.5
(2.5)3.58 cm, base cuneate, margin entire, plane, apex cuspidate or acuminate; midrib
flat above; lateral veins 46 pairs, raised below, faint above, basal pair reaching almost
1/2 the length of the blade, without glands in the axils; intercostal venation scalariform,
distinct below, faint above; petiole 1.57 cm long, sparsely reddish brown hairy, usually
with a subnodal gland. Syconia borne on twigs behind the leaves, ripening yellow to
red, subglobose or ellipsoid, c. 1 cm diameter, apex slightly convex and perforated, base
abruptly narrowed into a short stalk-like structure; peduncle up to 0.6 cm long; basal
bracts 3, 1.52.5 mm long; internal bristles numerous, 12 mm long, yellowish. Tepals
35, narrowly spathulate. Achenes 11.5 mm long, rugose, strongly keeled.
Distribution. Sumatra, Peninsular Malaysia, Java, Borneo, the Philippines, Sulawesi,
Maluku, and New Guinea. Widely distributed in Sabah and Sarawak. Also in Brunei
and Kalimantan.
Key to varieties
Leaves sparsely hairy below........................var. glandulifera
Leaves densely velvety hairy below...............
var. villosa Corner
(Latin, villosus = shaggy with long soft straight ascending hairs; the lower leaf surface)
Gard. Bull. Sing. 17 (1960) 433, l.c. (1965) 47; Whitmore, Tantra & Sutisna l.c. 246. Type: NGF
3753, New Guinea, Sepik district (holotype LAE).
Borneo, Morotai and Papua New Guinea. In Sabah very rare, known only by 1 collection,
SAN 2930. Not known in Sarawak, Brunei and Kalimantan.
24. Ficus grossularioides Burm. f.

(Latin grossularioides = like the gooseberry, Ribes grossularia; the shape of syconia)
Fl. Ind. (1768) 227; Backer & Bakhuizen f. l.c. 30; Corner l.c. (1965) 44, l.c. (1988) 547; Kochummen
l.c. 148; Anderson l.c. 259; Primack l.c. 89; Whitmore, Tantra & Sutisna l.c. 247; Coode et al. (eds.) l.c.
278
vi
215. Type: unknown. Synonyms: F. alba Reinw. ex Blume l.c. 467; F. nivea Blume l.c. 476; F. palmata
Roxb., Fl. Ind., ed. Carey, 3 (1832) 529; F. hunteri Miq. l.c. (1848) 225; F. chloroleuca Miq. l.c. (1859)
294; F. gossypina Wall. ex Miq. l.c. (1861) 425; F. mappan Miq. l.c. (1861) 425; F. alba Reinw. ex
Blume var. gossypina (Wall. ex Miq.) Kuntze, Rev. Gen. Pl. 1 (1891) 626; F. alba Reinw. ex Blume var.
nudinervis Kuntze l.c. (1891) 626; F. lobata Hunter ex Ridl., J. Str. Br. R. As. Soc. 53 (1909) 123.
Dioecious tree, up to 13 m tall, 15 cm diameter. Twigs 0.20.5 cm thick, reddish brown,

glabrous. Stipules pale hairy, lanceolate, acute, c. 0.5 cm long. Leaves spirally arranged,
thin-coriaceous, glaucous and velvety hairy below, very rough-hairy above; elliptic to
obovate, 5.519 3.511.5 cm, base cuneate, margin entire, lobed or distinctly toothed,
apex acute; midrib sunken above; lateral veins 46 pairs, sunken above, raised below,
basal pair reaching about 1/2 the length of the blade, with a gland in each axil below;
intercostal venation scalariform to reticulate, visible below, invisible above; petiole 1.5
11.5 cm long, pinkish on drying. Syconia axillary, sessile, in pairs, ripening yellowish
brown, ellipsoid to subglobose, 0.81 cm diameter, apex umbonate; basal bracts ovate,
12.5 mm long; internal bristles few, short, white. Tepals 3 or 5, glabrous. Achenes
c. 1 mm long, tuberculate.
Vernacular names. Sarawakkumpan, lenakan, lelingkan, lengkan (Iban), kukan (Bidayuh).
Distribution. Thailand, Sumatra, Peninsular Malaysia, Borneo, and Java. Common in
Sarawak, not yet reported for Sabah. Also in Brunei and Kalimantan.
Ecology. In primary and secondary lowland forests.
25. Ficus gul K. Schum. & Lauterb.

(local name of the plant in Papua New Guineagul)
Fl. Schutzgeb. (1901) 278; Corner l.c. 17 (1960) 452, l.c. (1965) 63; Whitmore, Tantra & Sutisna l.c.
247. Type: unknown. Synonyms: F. rudis Miq. l.c. (1867) 222; F. manilensis Warb. in Perkin, Fragm.
Fl. Philip. 3 (1905) 199.
Dioecious tree, up to 22 m tall, 30 cm diameter. Bark grey-brown, smooth; inner bark

pale. Sapwood pale. Twigs 0.20.35 cm thick, slender, initially rough-hairy. Stipules
lanceolate, acute, c. 0.5 cm long, hairy outside, caducous. Leaves spirally arranged, thincoriaceous, rough-hairy on both surfaces; elliptic to oblong or obovate, 817.5 3.8
8.5 cm, base symmetric, cuneate, margin toothed, apex acute; midrib flat above; lateral
veins 57 pairs, with short intermediate veins in between, faintly raised below, very faint
above, basal pair extending to nearly 1/2 the length of the blade, with a gland in each
axil below; intercostal venation scalariform, faint on both surfaces; petiole 1.55.5 cm
long, sparsely rough-hairy. Syconia hairy, in clusters, axillary or borne on twigs behind
the leaves, globose, 0.40.6 cm diameter, ripening dull red-purple, apex umbonate; peduncle
slender, c. 1 cm long; basal bracts absent, lateral bracts triangular, many; internal bristles
numerous, c. 0.3 mm long, white. Tepals (3)45, dark red, free, glabrous. Achenes c.
0.8 mm long, compressed, smooth, slightly keeled.
Distribution. Borneo, the Philippines, Sulawesi, Maluku, Tanimbar, Key Is., Flores, New
280
Guinea, Admiralty Islands, and New Britain. Widely distributed in Sabah, not yet recorded
for Sarawak. Not known in Brunei and Kalimantan.
Ecology. In lowland and hill forests.
26. Ficus hemsleyana King

(W.B. Hemsley, 18431924, sometime botanist and keeper of the Herbarium, Royal Botanic
Gardens Kew, U.K.)
Ann. Roy. Bot. Gard. Cal. 1, 2 (1888) 112; Merrill l.c. (1921) 224; Corner l.c. (1965) 79; Anderson l.c.
259; Primack l.c. 89; Whitmore, Tantra & Sutisna l.c. 247; Coode et al. (eds.) l.c. 214. Type: Beccari PB
2335, Borneo, Sarawak (FI).
Dioecious tree, up to 12 m tall, 20 cm diameter, or climber. Twigs c. 0.3 cm thick,

grey-white, slightly angled, covered with reddish brown hairs. Stipules lanceolate, acute,
1.51.8 cm long, brown hairy, subpersistent. Leaves spirally arranged, thin-coriaceous,
sessile, reddish brown hairy below to almost glabrous; oblong, 1935 513 cm, base
distinctly auriculate on one side, margin toothed, apex acuminate, acumen 45 cm long;
midrib raised above, keeled below; lateral veins 710 pairs, steeply ascending, raised
below, sunken above, basal pairs extending to almost 1/2 the length of the blade, with
a gland in each axil below; intercostal venation reticulate to scalariform, visible below,
distinctly sunken above. Syconia in clusters on tubercles arising from the stem, globose,
0.40.6 cm diameter, ripening orange, glabrous or hairy, apex depressed; peduncle 0.7
1.5 cm long; basal bracts 23, minute; internal bristles few, minute. Tepals 35, oblong
or ligulate, free or slightly joined at the base, white to red-brown. Achenes ovoid, c.
1.5 mm long, distinctly keeled at the apex.
Vernacular names. Sarawakkara (Iban), lunok (Kenyah).
Distribution. Endemic to Borneo. Common in Sarawak; rare in Sabah, known by a single
collection, SAN A 4417, from Kota Belud district. Also in Brunei and Kalimantan.
Ecology. In lowland mixed dipterocarp to lower montane forests, including limestone
habitat, up to 1600 m altitude, often by streams.
27. Ficus ixoroides Corner

(like Ixora, a genus in the Rubiaceae)
Gard. Bull. Sing. 18 (1960) 63, l.c. (1965) 94; Whitmore, Tantra & Sutisna l.c. 247. Type: Beccari PB
2781, Borneo, Sarawak (FI).
Dioecious shrub, without free hanging aerial roots. Twigs c. 0.2 cm thick, dark brown.
Stipules lanceolate, c. 2 cm long, glabrous, caducous. Leaves decussate or spirally
arranged, subcoriaceous, glabrous, drying greenish; lanceolate or narrowly oblong, 6
18 1.23.2 cm, base symmetric, cuneate, margin entire, plane, apex acute; midrib raised
above; lateral veins 816 pairs, well-spaced, curved, forming a broad angle with the
midrib, faintly raised on both surfaces, with short intermediate veins in between, joining
281
near the leaf margin, basal pair distinct; intercostal venation obsure; petiole 0.51.5 cm
long, with a subnodal gland. Syconia solitary, axillary, glabrous, subglobose, c. 1 cm
diameter, tapered at the base, apex flat; peduncle 0.20.4 cm long; basal bracts 3, c.
1 mm long, lateral bracts absent, apical bracts 5, small; internal bristles absent. Perianth
tubular, reddish brown. Achenes subtuberculate, not keeled.
Distribution. Endemic to Borneo. Rare in Sabah and Sarawak, known by 1 collection
each. Also in Kalimantan.
Ecology. In riparian forest.
28. Ficus lamponga Miq.

(of Lampong, Sumatra)
Fl. Ind. Bat., Suppl. (1861) 431; Corner l.c. (1965) 47; Kochummen l.c. 150; Anderson l.c. 259; Primack
l.c. 91; Whitmore, Tantra & Sutisna l.c. 248. Type: Teijsmann s.n., Sumatra, Lampong, Tarabangi Agong
(U?). Synonyms: F. lepidosa Wall. ex Kurz, J. As. Soc. Beng. 42 (1873) 107; F. balansae Gagnep., Not.
Syst. 4 (1927) 86.
Dioecious tree, up to 30 m tall, 50 cm diameter, without free hanging aerial roots. Bark
reddish brown. Twigs 0.20.4 cm thick, dark brown with prominent stipular scars. Stipules
ovate, 0.51.2 cm long, glabrous, caducous. Leaves spirally arranged, membranous,
glabrous on both surfaces; elliptic to oblong, 912.5 2.54.5 cm, base symmetric, cuneate,
margin entire, plane, apex acute; midrib raised above; lateral veins 713 pairs, wellspaced, curved, not steeply ascending, raised on both surfaces, basal pair distinct, without
axillary glands; intercostal venation reticulate, visible below, invisible above; petiole 1.5
2.5 cm long. Syconia in pairs on twigs behind leaves, ripening orange to red, globose
to ellipsoid, 0.81.3 cm diameter, base tapered, apex umbonate; peduncle up to 1.5 cm
long; basal bracts 2, 23 mm long; internal bristles numerous, c. 1 mm long, yellow.
Tepals 34, spathulate. Achenes oblong, c. 1 mm long, rugose, strongly keeled.
Vernacular names. Sarawakara, tempan (Iban).
Distribution. Assam, Burma, Indo-China, Thailand, Andaman Islands, Sumatra, Peninsular
Malaysia, Borneo, and Sulawesi. Very rare in Sabah, known by a single collection, SAN
31975, from Beaufort district. In Sarawak, recorded for Kapit, Lundu and Samarahan
(G. Buri) districts. Not known in Brunei and Kalimantan.
29. Ficus lepicarpa Blume

(Greek, lepis = scales, karpos = fruits; the occurrence of scale-like bracts on the surface of
syconia)
Bijdr. Fl. Ned. Ind. (1825) 459; Miquel l.c. (1867) 283; King l.c. (1888) 118; Merrill l.c. (1921) 224;
Ridley l.c. (1924) 343; Burkill l.c. 1028; Backer & Bakhuizen f. l.c. 33; Corner l.c. (1965) 90, l.c. (1988)
282
548; Kochummen l.c. 150; Anderson l.c. 259; Primack l.c. 91; Whitmore, Tantra & Sutisna l.c. 248;
Coode et al. (eds.) l.c. 215. Type: Blume s.n., Java (L?). Synonyms: Covellia volkameriaefolia Wall. ex
Miq. l.c. (1848) 464; C. didyma Miq. l.c. (1851) 65; C. lepicarpa (Blume) Miq. l.c. (1859) 328; F.
malaica Hunter ex Ridl., J. Str. Br. Roy. As. Soc. 53 (1909) 123; F. lepicarpa Blume var. brevipedunculata
Corner l.c. 18 (1960) 54; F. lepicarpa Blume var. brevibracteata Corner l.c. 18 (1960) 54; type: Haviland
s.n., Sarawak (K), syn. nov.
Dioecious tree, up to 10 m tall, 10 cm diameter, without free hanging aerial roots; buttresses
short. Bark greyish brown. Twigs c. 0.4 cm thick, grey-brown, angled, with prominent
grooves. Stipules lanceolate, acute, 12 cm long, persistent on twig ends. Leaves spirally
arranged, thin-coriaceous, glabrous on both surfaces; oblanceolate, 925 311.5 cm,
base symmetric, cuneate, margin entire, plane, apex acute; midrib raised above, rounded
below; lateral veins 810 pairs, well-spaced, curved, departing from the midrib at a broad
angle, raised below, faint above, basal pair distinct, without axillary glands; intercostal
venation scalariform, visible below, faint above; petiole 1.55 cm long. Syconia axillary,
sessile, ripening pale yellow to brownish, obovoid to subglobose, 1.21.5 11.2 cm;
basal bracts 3, 413 mm long, lateral bracts present; internal bristles absent. Tepals
fused into tubular perianth. Achenes c. 1 mm long, tuberculate, rough, keeled.
Vernacular names. Sarawakengkusuroh, kara (Iban).
Distribution. Burma, Thailand, Sumatra, Peninsular Malaysia, Borneo, Java, the Philippines,
Sulawesi, and Ambon. Common in Sabah and Sarawak. Also in Brunei and Kalimantan.
30. Ficus leptocalama Corner

(Greek, lepto = slender, kalamos = stalk; the slender-stalked syconia)
Gard. Bull. Sing. 17 (1960) 483, l.c. (1965) 79; Whitmore, Tantra & Sutisna l.c. 248; Coode et al. (eds.)
l.c. 215. Type: SFN 26829, Borneo, Sabah, Mt. Kinabalu, Menetendok R. (holotype SING).
Dioecious small tree, up to 5 m tall. Twigs slender, c. 0.1 cm thick, pale reddish yellow.
Stipules c. 0.8 cm long, subpersistent. Leaves distichous, thin-coriaceous, glabrous; elliptic,
815 2.54.5 cm, base symmetric, cuneate or auriculate, margin entire, plane, apex
caudate, acumen up to 3 cm long; midrib raised above, rounded below; lateral veins
710 pairs, well-spaced, curved, basal pair distinct, with a gland in one or both axils
below; petiole 0.10.3 cm long. Syconia solitary or in pairs, borne on twigs behind the
leaves or on stolons arising from the base of the stem, or on short drooping leafless
branches arising from near the base of the stem, ripening rose-pink to red, subglobose,
0.60.8 cm diameter, base tapered into a stalk 0.10.2 cm long; basal bracts 23, minute,
lateral bracts present; internal bristles absent. Tepals 4, white, free or slightly united
at the base, glabrous. Achene oblong, c. 1.2 mm long, slightly compressed, not keeled,
smooth.
Distribution. Endemic to Borneo. In Sabah very rare, known by the type only from
Mt. Kinabalu; not known in Sarawak. Also in Brunei and Kalimantan.
283
31. Ficus leptogramma Corner

(Greek, lepto = slender, gramma = streaks or lines; the striate stipules)
Gard. Bull. Sing. 17 (1960) 473, l.c. (1965) 73; Anderson l.c. 259; Primack l.c. 92; Whitmore, Tantra
& Sutisna l.c. 249. Type: SFN 27328, Borneo, Sabah, Mt. Kinabalu (holotype SING).
Dioecious tree, up to 12 m tall, 30 cm diameter. Bark dark grey, smooth. Twigs c. 0.3
cm thick, light greyish brown, sulcate or striate. Stipules paired, lanceolate, up to 1.5
cm long, glabrous, longitudinally striate, subpersistent. Leaves distichous, coriaceous,
glabrous; elliptic to subovate, 1035 2.714 cm, base rounded or cuneate, margin lobed
or faintly toothed, apex caudate, acumen up to 6 cm long; midrib raised above; lateral
veins 812 pairs, with short intermediate veins in between, sunken above, raised below,
basal pair short, with a gland in each axil below; intercostal venation scalariform or
reticulate, sunken above; petiole 0.32.5 cm long. Syconia borne in clusters on stem
and branches, ripening olive tinted purple to dark crimson, subglobose to ellipsoid, 1
1.5 11.2 cm, apex umbonate; peduncle 12.5 cm long; basal bracts absent, lateral
bracts many and scattered; internal bristles absent. Tepals 38, white or pinkish, free
or slightly joined at the base, glabrous. Achenes lenticular, c. 1 mm long, slightly
compressed, not keeled.
Distribution. Endemic to Borneo. Very rare in Sabah, more frequent in Sarawak. Not
known in Brunei and Kalimantan.
Ecology. In lowland to hill mixed dipterocarp forests, including limestone habitat, up
to 1200 m altitude.
32. Ficus macilenta King

(Latin, macilentus = thin; the leaves)
Ann. Roy. Bot. Gard. Calc. 1, 2 (1888) 155; Merrill l.c. (1921) 224; Corner l.c. (1965) 49; Primack
l.c. 92; Whitmore, Tantra & Sutisna l.c. 249. Type: Beccari PB 1696, Borneo, Sarawak (FI).
Dioecious tree, up to 8 m tall, 10 cm diameter. Twigs 0.30.5 cm thick, yellowish hairy,

glabrescent. Stipules lanceolate, c. 0.8 cm long, glabrous or hairy, caducous. Leaves
spirally arranged, thin-coriaceous, drying greenish yellow, glabrous or sparsely to densely
hairy below, rough-hairy above; elliptic to rhomboid, 3.519 1.510 cm, base cuneate,
margin entire and plane, or distantly shallowly lobed or toothed, apex acuminate, acumen
up to 1.5 cm long; midrib sunken above; lateral veins 5-7 pairs, raised below, faint above,
curving and joining near the leaf margin, basal pair short, sometimes with a gland in
each axil below; intercostal venation reticulate, distinct below, faint to invisible above;
petiole 0.52 cm long, glabrous or hairy. Syconia axillary, solitary or in pairs, sessile
or with short peduncle, densely hispid with stiff brown hairs, ripening red, ellipsoid or
subglobose, 0.52 cm diameter, glabrous or hairy; basal and apical bracts tiny; internal
bristles numerous, 12 mm long, white. Tepals 34, long-hairy, white. Achenes c. 1 mm
long, tuberculate, slightly keeled.
Vernacular names. Sarawaktalatad (Kelabit), tempan (Iban).
284
Distribution. Endemic to Borneo. Uncommon in Sabah and Sarawak. Also in Brunei.

Ecology. In hill mixed dipterocarp to upper montane forests, at 8002000 m altitude.
Key to varieties
1.
Leaves densely velvety hairy below. Syconia sessile................var. macilenta

Leaves sparsely hairy or glabrous below. Syconia short-peduncled...........2
2.
Upper half of leaves strongly lobed. Syconia solitary............

var. ilicifolia Corner
(Latin, ilicifolius = with leaves resembling those of holly treeIlex)
Gard. Bull. Sing. 17 (1960) 439, l.c. (1965) 49; Whitmore, Tantra and Sutisna l.c. 249. Type:
SFN 26952, Borneo, Sabah, Ranau district, Mt. Kinabalu (holotype SING).
Differs from the other varieties in the short-pubescent, lobed leaves and solitary
syconia.
Endemic to Borneo; rare and confined to Mt. Kinabalu in Sabah. In lower to
upper montane forests, at 15002000 m altitude.
Leaf margin entire, plane or faintly distantly toothed. Syconia in pairs............
var. gibbsiae (Ridl.) Corner
(L.S. Gibbs, 18701925, the British botanist)
Gard. Bull. Sing. 17 (1960) 439, l.c. (1965) 49; Whitmore, Tantra & Sutisna l.c. 249; Coode et
al. (eds.) l.c. 215. Basionym: F. gibbsiae Ridl., J. Linn. Soc. Bot. 42 (1915) 137. Type: unknown.
Differs from var. macilenta in the peduncled syconia and less hairy leaves and
syconia.
Endemic to Borneo. Rare in Sabah, more common in Sarawak. Also in Brunei.
In hill mixed dipterocarp to lower montane forests, at 8001300 m altitude.
33. Ficus macrostyla Corner

(Greek, makro = large or long, Latin, stylus = style; the long-styled female flowers)
Gard. Bull. Sing. 18 (1960) 43, l.c. (1965) 85; Whitmore, Tantra & Sutisna l.c. 249. Type: Haviland
3601, Borneo, Sarawak (K).
Dioecious shrub. Twigs c. 0.3 cm thick, dark brown, with large stipular scars, hairy.
Stipules narrowly lanceolate, up to 1.5 cm long, yellowish hairy, persistent. Leaves spirally
arranged, coriaceous, glabrous or yellowish brown hairy below; obovate or oblanceolate,
6.521 2.55.5 cm, base attenuate, margin entire, revolute, apex obtuse; midrib raised
above; lateral veins 811 pairs, with short intermediate veins in between, raised on both
surfaces, basal pair distinct, departing from the midrib at an acute angle, without gland
in the axils below; intercostal venation reticulate, distinct below, invisible above; petiole
1.24 cm long, yellowish brown hairy. Syconia axillary or borne on twigs behind the
leaves or on stolons, hairy, ovoid, c. 1.7 0.6 cm, surface completely covered with long,
flat, straight, appressed, brown-hairy bracts; internal bristles few, short, brown. Perianth
cup-shaped. Achenes c. 1 0.6 mm, subcompressed, smooth, strongly keeled.
285
Distribution. Endemic to Borneo, and confined to Sarawak, gregarious where it occurs.

Ecology. In lowland mixed dipterocarp forest by streams.
Use. Figs edible.
34. Ficus magnoliifolia Blume

(with leaves like that of Magnolia, Magnoliaceae)
Bijdr. Fl. Ned. Ind. (1825) 448; Miquel l.c. (1867) 263; Backer & Bakhuizen f. l.c. 27; Corner l.c. (1965)
31; Kochummen l.c. 151; Anderson l.c. 259; Primack l.c. 93; Whitmore, Tantra & Sutisna l.c. 249.
Type: Blume s.n., Java (L?). Synonyms: Urostigma modestum Miq. l.c. (1851) 5; U. euneuron Miq. l.c.
(1859) 599; F. nervosa (non Heyne ex Roth) Elmer, Leaft. Philip. Bot. 2 (1908) 537; F. apoensis Elmer,
Leafl. Philip. Bot. 4 (1911) 1249.
Monoecious tree, up to 45 m tall, 70 cm diameter; buttresses up to 2 m tall. Bark greybrown, smooth to scaly; inner bark brown; latex citron-yellow, acrid, poisonous. Sapwood
yellowish. Twigs c. 0.3 cm thick, dark brown, irregularly grooved on drying. Stipules
narrowly lanceolate, acute, c. 0.8 cm long, caducous. Leaves spirally arranged or
distichous, coriaceous, glabrous, brownish on drying; oblong, 920 2.58.5 cm, base
cuneate, margin entire, plane, apex acute to acuminate, acumen up to 2.5 cm long; midrib
sunken above; lateral veins 1012 pairs, flat to slightly raised above, prominent below,
distinctly curving and joining near the leaf margin, with shorter veins in between, basal
veins short, with a gland in each axil below; intercostal venation reticulate, faintly visible
on both surfaces; petiole 0.50.8 cm long. Syconia axillary, solitary or paired, globose,
1.51.8 cm diameter, apex with small aperture; peduncle up to 1.5 cm long; basal bracts
3, ovate 12 mm long; internal bristles absent. Tepals 24, red, free or somewhat fused,
glabrous. Achene 1.21.4 mm long, smooth, slightly keeled.
Vernacular names. Sarawaklumau (Iban), puro (Kayan).
Distribution. Nicobar and Andaman Islands, Sumatra, Peninsular Malaysia, Java, Borneo,
Sulawesi, and the Philippines. Very rare in Sabah and Sarawak, known by 1 collection,
SAN 33972, from Sabah and a few from Sarawak. Also in Kalimantan.
Ecology. In lowland forest.
35. Ficus megaleia Corner

(Greek, mega = large, leios = smooth to the touch; probably referring to the leaves)
l.c. 216. Type: Corner BRUN 5301, Borneo, Brunei, Ulu Belalong (holotype CGE).
Dioecious tree, up to 7 m tall, with drooping young twigs. Twigs, stipules, petioles and
leaf undersurface covered with 11.5 mm long, reddish brown or yellowish hairs. Twigs
0.40.7 cm thick. Stipules oblong, 37 cm long, caducous. Leaves distichous, pubescent
286
with rough short hairs above, subcoriaceous or chartaceous; elliptic to oblong, 40100
1435 cm, base asymmetric, cordate, margin finely toothed, apex caudate or acuminate,
acumen 2.54 cm long; midrib flat above; lateral veins 1024 pairs, raised below, visible
above, basal veins 67, short, mostly with a gland in each axil below; intercostal venation
scalariform, raised below, faint above; petiole 12.5 cm long. Syconia borne on long
stolons arising from the base of stem, globose, 34 cm diameter, densely stiff-hairy,
glabrescent, covered with many ovate-lanceolate bracts; internal bristles absent. Perianth
absent or rudimentary. Achenes c. 0.7 mm long, slightly keeled, slightly ridged.
Distribution. Endemic to Borneo. Rare in Sabah and Sarawak. Also in Brunei and
Kalimantan.
Ecology. In hill mixed dipterocarp to lower montane forests, at 8001400 m altitude.
Key to varieties
1.
Syconia glabrous..........
var. subuncinata Corner
(Latin, subuncinatus = somewhat barbed or hooked; the twigs and petioles)
Gard. Bull. Sing. 19 (1962) 400, l.c. (1965) 91; Whitmore, Tantra & Sutisna l.c. 249. Type:
RSNB 1576, Borneo, Sabah, Mt. Kinabalu (holotype K).
Very similar to var. megaleia differing in the glabrous syconia.

Endemic to Mt. Kinabalu in Sabah; in lower montane forest at c. 1500 m altitude.
Syconia hairy..................2
2.
Hairs yellowish, up to 1 mm long; lateral veins 1424 pairs..............

var. multinervia Corner
(Latin, multinervius = with many lateral veins; the leaves)
RSNB 970, Borneo, Sabah, Mt. Kinabalu (holotype K).
Leaves very much bigger than the typical variety. The hanging pink leaves are
conspicuous. Hairs yellowish, dense, up to 1 mm long; lateral veins 1424 pairs,
with 46 basal veins.
Endemic to Borneo, and confined to Mt. Kinabalu, Sabah; in hill mixed dipterocarp
forest up to 900 m altitude.
Hairs reddish brown, more than 1 mm long; lateral veins 1020 pairs....var. megaleia
36. Ficus melinocarpa Blume

(Greek, melinos = yellowish, karpos = fruit; the ripe syconia)
Bijdr. Fl. Ned. Ind. (1825) 460; King l.c. (1888) 94; Backer & Bakhuizen f. l.c. 29; Corner l.c. (1965)
68; Whitmore, Tantra & Sutisna l.c. 249. Type: Blume s.n. (= RHL Sheet No. 9081881059), Java, Mt.
Salak (L). Synonyms: F. daemonum Zoll. & Moritzi, Syst. Verz. (1846) 77; Pogonotrophe alnifolia
Miq. l.c. (1851) 51; F. obliqua (non Koen. ex Vahl) Miq. l.c. (1854) 92; F. alnifolia (Miq.) Miq. l.c.
(1867) 278; F. haggeri Merr., Philip. J. Sc. 18 (1921) 62; F. irosinensis Elmer, Leafl. Philip. J. Sc.
(1921) 62.
Dioecious tree, up to 35 m tall, deciduous; buttresses up to 4 m tall, 2 m wide. Bark grey287
brown; inner bark whitish. Twigs c. 0.3 cm thick, dark brown, rough-hairy at the tip, slightly
hollow. Stipules lanceolate, up to 1.2 cm long, rough-hairy, caducous. Leaves distichous,
thin-coriaceous, smooth and sparsely yellowish hairy on both surfaces; elliptic or subovate,
717.5 3.59 cm, base broadly cuneate, margin typically distantly toothed, rarely entire
and revolute, apex obtuse; midrib raised above; lateral veins 68 pairs, with shorter veins in
between, raised on both surfaces, basal pairs 13, extending up to 1/2 the length of the blade,
occasionally with a gland in each axil below; intercostal venation scalariform to reticulate,
distinct below, faint above; petiole 12 cm long, rough-hairy. Syconia axillary or borne on
twigs behind the leaves, short-hairy, glabrescent, ripening yellow, orange, red to purple black,
globose to pear-shaped, 0.71.2 cm diameter, apex umbonate, slightly perforate; peduncle
0.50.7 cm long; basal bracts 3, small; internal bristles numerous, minute. Tepals 36, reddish
pink, free, glabrous. Achenes lenticular, up to 1 mm long, almost smooth, scarcely keeled.
Distribution. Sumatra, Java, Borneo, Sulawesi, the Philippines, Maluku, New Guinea,
New Britain, and Solomon Islands. Common in Sabah, not yet recorded for Sarawak.
Ecology. In hill forest, up to 400 m altitude.
37. Ficus microcarpa L. f.

(Greek, mikro = small, karpos = fruit; with small syconia)
Sp. Pl., Suppl. (1781) 442; Backer & Bakhuizen f. l.c. 24; Corner l.c. (1965) 22, l.c. (1988) 542;
Kochummen l.c. 151; Anderson l.c. 259; Primack l.c. 93; Whitmore, Tantra & Sutisna l.c. 250; Coode et
al. (eds.) l.c. 216. Type: unknown. Synonyms: F. littoralis Blume l.c. 455; Urostigma microcarpum (L.
f.) Miq. l.c. (1847) 583; U. amblyphyllum Miq. l.c. (1847) 569; U. pisiferum Miq. l.c. (1847) 580; F.
amblyphylla (Miq.) Miq. l.c. (1867) 286; F. retusa L. var. pisifera (Miq.) Miq. l.c. (1867) 288.
Monoecious tree (or strangler), up to 30 m tall, 40 cm diameter, with abundant free hanging
aerial roots. Bark grey. Twigs c. 0.3 cm thick, dark brown. Stipules glabrous, lanceolate,
acuminate, up to 1.5 cm long. Leaves distichous, coriaceous, glabrous; elliptic to obovate,
49 24.5 cm, base symmetric, cuneate, margin entire, plane, apex acute; midrib flat
above, often pale coloured below when dry; lateral veins 46 pairs, well-spaced, curved,
steeply ascending, looping and joining near the leaf margin, with few equally prominent
shorter veins in between, visible below, invisible above, basal pair distinct, reaching
up to 1/31/2 the length of the blade, without axillary glands below; intercostal venation
invisible; petiole 0.41 cm long, grooved on the adaxial side. Syconia axillary, sessile,
in pairs, dark red when ripe, globose, 0.50.7 cm diameter, apex umbonate; basal bracts
ovate to lanceolate, 1.53.5 1.52.5 mm, apex rounded, persistent; internal bristles few
to numerous, up to 0.5 mm long. Tepals 34, spathulate or obovate, free. Achenes oblong,
c. 2 1 mm, smooth, with pink dots.
Vernacular name. Sarawaklunok (Murut).
Distribution. Cocos and Christmas Islands, Sri Lanka, India, S China, Ryu-Kyu, throughout
Malesia, New Britain, Australia, and Caroline Islands. Common in Sabah and Sarawak.
288
Ecology. Lowland forest by streams, coastal and swamp forest, often planted in urban
areas.
38. Ficus midotis Corner

(meaning obscure)
Gard. Bull. Sing. 17 (1960) 482, l.c. (1965) 79; Primack l.c. 94; Whitmore, Tantra & Sutisna l.c. 250.
Type: SFN 26324, Borneo, Sabah (holotype SING).
Dioecious tree, up to 10 m tall (or epiphyte). Twigs up to 0.25 cm thick, yellowish

brown. Stipules linear or lanceolate, up to 1.2 cm long, caducous. Leaves spirally arranged,
thin-coriaceous, glabrous, rough to the touch; ovate to elliptic or oblong, 8.518 2.5
8 cm, base symmetric and cuneate, or asymmetric and with tiny auricle on one side,
margin faintly lobed or toothed or entire and plane, apex caudate, acumen 14.5 cm
long; midrib flat or rarely raised above, sharply keeled below; lateral veins 57 pairs,
well-spaced, curved, with short veins in between, raised below, faint or rarely sunken
above, basal pair extending up to 1/3 the length of the blade, with a gland in the axils
of one or both of the veins; intercostal venation reticulate, faint on both surfaces; petiole
0.31 cm long. Syconia paired or in clusters, axillary or borne on twigs behind the
leaves or on stem, ripening yellow to red, subglobose, 0.161 cm diameter, apex umbonate;
peduncle 0.32.5 cm long; basal bracts 12, minute, lateral bracts absent; internal bristles
absent or few, minute. Tepals 57, whitish, oblong, puberulous, slightly united at the
base. Achenes oblong, c. 1.5 mm long, somewhat compressed, keeled at the apex.
Vernacular names. Sarawakandai (Bidayuh), kara (Iban).
Distribution. Endemic to Borneo. Uncommon in Sabah and Sarawak. Not known in Brunei
and Kalimantan.
Ecology. In lowland mixed dipterocarp to upper montane forests, up to 2800 m altitude.
39. Ficus minahassae (Teijsm. & de Vries) Miq.

(of Minahasa in Sulawesi)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 231; King l.c. (1888) 108; Merrill, Bull. Bur. For. Philip. 1 (1903)
18; Corner l.c. (1965) 63. Basionym: Bosscheria minahassae Teijsm. & de Vriese, Nat. Tijds. Ned. Ind.
23 (1861) 212. Type: unknown. Synonym: F. riedelii (non Teijsm. ex Miq.) Haberlandt, Bot. Trop.
(1893) 191.
Dioecious tree, up to 25 m tall, sometimes with stilt-roots. Twigs 0.40.6 cm thick,

pubescent with spreading hairs 24 mm long. Stipules ovate or lanceolate, 25 cm long,
hairy, caducous or persistent on the new shoots. Leaves spirally arranged, coriaceous,
rough-hairy above, yellowish soft-hairy below; ovate, (9)1330 (5)720 cm, base
rounded or cordate, margin entire, plane, apex acute or acuminate, acumen up to 1.2
cm long; midrib raised above; lateral veins 711 pairs, basal pairs 35, short, departing
from the midrib at a broad angle, with a gland in each axil of the lower most veins
below; intercostal venation scalariform slightly raised below; petiole 3.510 cm long.
289
Syconia borne in small globular clusters along unbranched leafless twigs hanging from
the trunk and main branches, glabrous or finely hairy, ripening red, obconical, 0.50.6
cm diameter; peduncle 0.150.2 cm long; basal bracts 3, ovate, c. 1.5 mm long, persistent;
internal bristles abundant, white. Tepals 34, spathulate, pinkish or pale reddish, glabrous,
free. Achenes c. 0.7 mm long, compressed, smooth, slightly keeled.
Vernacular name. Sabahtapian diwit (Dusun).
Distribution. Borneo, the Philippines and Sulawesi (Minahasa). There is only one record
of this species from Borneo, Agama 9464, from Timbun Mata Forest Reserve, Semporna
district, in Sabah.
Uses. Figs edible; bark used for making various utensils in the Philippines and Sulawesi.
40. Ficus moderata Corner

(Latin, moderatus = moderate; bridging the subser. Congestae and subser. Hispidae of the
ser. Tuberculifasciculatae)
Gard. Bull. Sing. 19 (1962) 396, l.c. (1965) 89; Whitmore, Tantra & Sutisna l.c. 250. Type: RSNB 2592,
Borneo, Sabah, Mt. Kinabalu (holotype K).
Dioecious tree, up to 10 m tall. Bark grey, smooth. Twigs c. 0.4 cm thick, solid, zigzag, brown, with ring-like stipular scars, hairy. Stipules lanceolate, c. 1.5 cm long, glabrous
or hairy at the base, caducous. Leaves spirally arranged, sometimes opposite, thincoriaceous, hairy below; elliptic to obovate, 821 511 cm, base broadly cuneate to
narrowly subcordate, margin entire or faintly lobed towards apex, apex acuminate, acumen
1.5 cm long; midrib raised above; lateral veins 69 pairs, with shorter veins in between,
finely raised below, basal veins short or extending up to 1/3 the length of the blade,
without axillary glands; intercostal venation scalariform, distinct below; petiole 0.92.5
cm long, powdery hairy. Syconia clustered on short branched compact woody processes
arising from stem and branches, pear-shaped, 1.52.3 cm diameter, surface rough with
incurved short lateral bracts; peduncle 0.41.4 cm long; basal bracts ovate, 23 cm long,
persistent; internal bristles absent. Perianth saccate or collar-like. Achenes c. 1 mm long,
keeled, obscurely tuberculate.
Distribution. Endemic to Borneo. Very rare, known only by 2 collections from Mt.
Kinabalu, Sabah.
Ecology. In hill mixed dipterocarp and lower montane forests, up to 1200 m altitude.
41. Ficus montana Burm. f.

(Latin, montanus = pertaining to mountains; the habitat)
Fl. Ind. (1768) 226; Backer & Bakhuizen f. l.c. 27; Corner l.c. (1965) 64; Whitmore, Tantra & Sutisna
l.c. 250. Type: unknown. Synonyms: F. humilis Roxb., Fl. Ind., ed. Carey, 3 (1832) 535; F. quercifolia
Roxb. l.c. 534; F. inconstans Miq. l.c. (1848) 232; F. sclerocoma Miq. l.c. (1851) 58; F. ampelas Burm.
f. var. bogoriensis (Koord. & Valeton) Hochr., Candollea 2 (1925) 328.
290
Dioecious shrub. Young parts rough-hairy. Twigs reddish brown, c. 0.3 cm thick, hollow.
Stipules paired, linear or lanceolate, c. 0.8 cm long, rough-hairy, caducous. Leaves spirally
arranged, coriaceous, rough-hairy below, sparsely hairy above; elliptic, 610 25 cm,
base cuneate, margin irregularly lobed, apex acuminate, acumen up to 1.5 cm long; midrib
raised above; lateral veins 57 pairs, distinct below, faint above, basal veins short, without
axillary glands; intercostal venation reticulate, distinct below, invisible above; petiole 0.5
1 cm long, rough-hairy. Syconia solitary or in pairs, axillary, rough-hairy, ripening red,
ovoid to subglobose, c. 0.5 0.3 cm, apex umbonate to flat; peduncle up to 0.3 cm
long; basal bracts 23, minute; internal bristles absent. Tepals 35, white, free, glabrous.
Achenes 11.3 mm long, slightly compressed, minutely tuberculate, hardly keeled.
Distribution. India, Burma, Thailand, Sumatra, Peninsular Malaysia, Java, and Borneo.
Very rare in Sabah and Sarawak; known by 2 collections, SAN A 1053 and SAN 117896,
from Sabah, and by S. 36002, S. 40985, and S. 44751 from Kapit district in Sarawak.
Ecology. In lowland to lower montane forests, usually by streams.
Uses. Young leaves are eaten as salad; mature leaves are used in opium smoking in the
past; the fruits edible.
42. Ficus nota (Blanco) Merr.

(Latin, notus = distinguishing mark; inference unknown)
Philip. Gov. Lab. Bur. Bull. 17 (1904) 10; Corner l.c. (1965) 88; Whitmore, Tantra & Sutisna l.c. 250.
Basionym: F. aspera G. Forst. var. nota Blanco., Fl. Filip. (1837) 677. Type: Merrill Spec. Blanc. No.
550, the Philippines, Luzon, Rizal Province, Antipolo (n.v).
Dioecious tree, up to 13 m tall, often shortly buttressed. Bark brown, smooth. Twigs
solid, 0.30.6 cm thick, brown, with large petiolar scars. Stipules appressed hairy,
lanceolate, c. 1.5 cm long, caducous. Leaves spirally arranged, coriaceous, sparsely hairy
below; obovate, 15.523 917 cm, base broadly cuneate, margin entire, plane, apex
obtuse; midrib raised above; lateral veins 67 pairs, raised on both surfaces, basal veins
34 pairs, reaching up to 1/2 the length of the blade, often with axillary glands below;
intercostal venation scalariform, visible on both surfaces; petiole 13.5 cm long, hairy.
Syconia borne on finger-like leafless processes arising from stem, ripening brown, pearshaped, 1.74.5 cm diameter, apex truncate with many bracts, surface rugose; peduncle
c. 0.5 cm long; basal bracts 3, 12 mm long; internal bristles few, white. Perianth cupshaped or collar-like. Achenes c. 1 mm long, smooth, keeled.
Distribution. Borneo and the Philippines. Very rare in Borneo, known only from Beaufort
district in Sabah.
Ecology. In lowland mixed dipterocarp forest.
43. Ficus obpyramidata King

(Latin, ob- = inverted, pyramidatus = pyramid-shaped; the shape of the syconia)
291
Ann. Roy. Bot. Gard. Cal. 1, 2 (1888) 116; Ridley l.c. (1924) 343; Burkill l.c. 1028; Corner l.c. (1965)
88, l.c. (1988) 551; Kochummen l.c. 152; Anderson l.c. 259; Primack l.c. 94; Whitmore, Tantra &
Sutisna l.c. 250; Turner l.c. 352. Type: Kunstler 1849, Peninsular Malaysia, Perak (K?).
Dioecious tree, up to 10 m tall, 20 cm diameter. Bark greyish brown. Twigs 0.30.6

cm thick, hollow, sparsely hairy with stiff dark brown hairs or glabrous. Stipules lanceolate,
22.8 cm long, dark brown hairy, caducous. Leaves spirally arranged, membranous,
sparsely dark brown hairy below; elliptic to obovate, 1420 4.58.5 cm, base cuneate,
symmetric, margin entire or faintly toothed, apex acute; midrib flat above; lateral veins
58 pairs, with shorter veins in between, raised on both surfaces, basal veins 23 pairs,
reaching up to 1/41/3 the length of the blade; intercostal venation reticulate, visible
below, faint above; petiole 1.55 cm long, hairy, glabrescent, with a subnodal gland.
Syconia borne on long finger-like projections arising from branches and stem, ripening
yellow to brownish ochre, obpyramidal or subconical with depressed apex, 22.5 cm
diameter, glabrous, with faint irregular lines and tiny projections; peduncle 13.6 cm
long; basal bracts 3, tiny, persistent, recurved; internal bristles few to abundant, whitish.
Perianth tubular, covering 1/3 to the whole ovary or stamen. Achenes ovoid, c. 1 mm
long, rather rough, slightly keeled.
Vernacular name. Sarawakengkunoh (Iban).
Distribution. Burma, Thailand, Peninsular Malaysia, and Borneo. Rare in Sabah and
Sarawak, known by SAN 91021 from Ranau district in Sabah, and by S. 18271 and S.
39842 from Belaga district, and S. 22036 from Kp. Segan in Bintulu district, Sarawak.
44. Ficus obscura Blume

(Latin, obscurus = indistinct; inference unknown)
Corner l.c. 17 (1960) 480, l.c. (1965) 78; Kochummen l.c. 152; Anderson l.c. 259; Primack l.c. 95;
Whitmore, Tantra & Sutisna l.c. 250; Turner l.c. 352; Coode et al. (eds.) l.c. 216. Type: Blume s.n. (=
RHL Sheet No. 908188590), Java (L). Synonyms: F. grewiaefolia Blume l.c. 473; F. scaberrima Blume
l.c. 474; F. pisifera Wall. ex Voight, Hort. Suburb. Calc. (1845) 285; F. acuminatissima Miq. l.c. (1848)
233; F. dasycaula Miq. l.c. (1848) 233; F. lobbii Miq. l.c. (1848) 233; F. remblas Miq. l.c. (1851) 60; F.
tajam Miq. l.c. (1851) 62; F. microtus Miq. l.c. (1861) 428; F. subdenticulata (Miq.) Miq. l.c. (1867)
296; F. grewiaefolia Blume var. angustata Miq. l.c. (1867) 273; F. microtus Miq. var. borneensis Miq.
l.c. (1867) 273; F. scaberrima Blume var. angustata Miq. l.c. (1859) 309; F. celebica Blume var.
kunstleri King, l.c.(1887) 12; F. obscura Blume var. scaberrima (Blume) Miq. l.c. (1867) 273, syn.
nov.; F. viridifolia Merr., Philip. J. Sc. (1913) Bot. 366; F. microsyce Ridl., Kew Bull. (1926) 82; F.
obscura Blume var. angustata (Miq.) Corner l.c. 17 (1960) 481, syn. nov.; F. obscura Blume var.
borneensis (Miq.) Corner l.c. 17 (1960) 481, syn. nov.; F. obscura Blume var. kunstleri (King) Corner
l.c. 17 (1960) 481, syn. nov.
Dioecious tree, up to 7 m tall, or epiphyte. Twigs c. 0.3 cm thick, terete, grooved, rough
or woolly hairy. Stipules lanceolate, c. 1.5 cm long, with long hairs along the margin,
292
persistent. Leaves distichous, membranous to subcoriaceous, almost sessile or with petiole

up to 0.7 cm long, rough-hairy to almost glabrous above, brownish soft-hairy below;
oblong, elliptic, oblanceolate or subtrapeziform, 3.528 2.59.5 cm, base strongly
asymmetric, cuneate, margin distantly toothed, lobed, or occasionally entire, apex acuminate
to caudate, acumen 36.5 cm long; midrib raised above; lateral veins 57 pairs, raised
below, faint above, basal pairs 23, short, often with orange-brown glands in the axils
below; intercostal venation reticulate, visible below, faint above. Syconia axillary or borne
on twigs behind the leaves, in pairs or in fascicles of 38, sessile or with short peduncle,
sparsely to densely hairy, ripening greyish white to pink, yellow, orange and scarlet,
subglobose, 0.51.2 cm diameter, apex umbonate, ostiole with 5 peripheral bracts; basal
bracts 23, minute; internal bristles absent or minute. Tepals 36, lanceolate, free or
united at base to various degree, white, sometimes with pink spots. Achenes c. 1.5
0.8 mm, strongly keeled at the apex.
Vernacular names. Sarawakdunduk (Bidayuh), kara (Iban).
Distribution. Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, the Philippines, and
Sulawesi. Common and widely distributed in Sabah and Sarawak. Also in Brunei and
Kalimantan.
Ecology. In lowland and hill forests, usually by streams, rarely on limestones.
Key to varieties
Twigs and leaves rough-hairy. Syconia sparsely hairy...........var. obscura
Twigs, leaves and syconia woolly soft-hairy............
var. lanata Kochummen
(Latin, lanatus = clothed with woolly hairs; the petioles and leaf undersurface)
Gard. Bull. Sing. 50 (1998) 215. Type: SAN 57361, Borneo, Sabah, Semporna district (holotype
KEP; isotype SAN).
Twigs reddish brown, woolly soft-hairy. Leaves woolly soft-hairy below, rough to
the touch above. Syconia axillary or borne on twigs behind the leaves, in pairs or
in clusters, sessile or peduncled, covered with woolly soft-hairs, subglobose, 0.8
1 cm diameter.
Endemic to Borneo. Frequent in Sabah; very rare in Sarawak, known by 1 collection
S. 28102 from Serian district. In lowland and hill mixed dipterocarp forests, often
by streams, occasionally on limestones. Not known in Brunei and Kalimantan.
45. Ficus oleifolia King

(with leaves like that of olive treeOlea, Oleaceae)
Ann. Roy. Bot. Gard. Calc. 1, 2 (1888) 160; Corner l.c. (1965) 38, Phil. Trans. Roy Soc. London Ser. B,
256 (1969) 313; Anderson l.c. 260; Primack l.c. 95; Whitmore, Tantra & Sutisna l.c. 251; Coode et al.
(eds.) l.c. 216. Type: Beccari PS 82, Sumatra, Mt. Singalang (holotype FI; isotype L). Synonym: F.
oleifolia King var. major King l.c. (1888) 160.
Dioecious shrub, up to 5 m tall. Twigs up to 0.2 cm thick, dark brown, irregularly swollen.
Stipules linear to lanceolate, 0.30.8 cm long, glabrous. Leaves spirally arranged,
293
subcoriaceous, glabrous, sessile or with petiole up to 1(1.3) cm long; elliptic, oblong,

lanceolate, obovate, narrowly elliptic to linear, 2.510.5 0.52.5 cm, base attenuate
to cuneate, margin entire, revolute or plane, apex rounded, obtuse or acute-acuminate;
midrib raised or sunken above; lateral veins 520 pairs, flat above, joining near the
leaf margin to form distinct intramarginal vein, basal veins 1 pair, short, with a gland
in each axil below; intercostal venation equally prominent as lateral veins or invisible.
Syconia in pairs, axillary, sessile or with peduncle up to 2.5 cm long, ripening dark
purple, subglobose to ellipsoid, 0.20.8 cm diameter, apex umbonate, sometimes basal
part slightly tapered; basal bracts 3, persistent; internal bristles absent. Tepals 3 or 4,
red, free. Achenes smooth or angled, kidney-shaped, c. 4 mm long.
Vernacular names. Sarawakara (Iban), kayu tanyit, lunok, sulok kondat (Kenyah).
Distribution. Sumatra, Borneo, the Philippines, and Sulawesi. Common in Sabah and
Ecology. In lowland to upper montane forests, up to 2400 m altitude, including riverbank
and limestone habitats.
Key to varieties
(The epiphytic varieties are annotated under the main key: lead 170)
1.
Leaf apex acute to acuminate....2

Leaf apex obtuse or rounded.........5
2.
Leaves linear or narrowly lanceolate, drying greenish, up to 1 cm wide...

var. linearifolia Corner
(Latin, linearifolius = linear-leaved)
Phil. Trans. Roy. Soc. London Ser. B, 256 (1969) 313; Coode et al. (eds.) l.c. 216. Type: Burt
& Martin B 5497, Borneo, Sarawak (holotype E).
Small tree. Twigs blackish. Leaves drying greenish, linear to narrowly lanceolate,
2.511 0.61 cm; base attenuate, often with 2 glands near petiole below, margin
revolute, apex acute to acuminate, acumen 1 cm long; midrib raised above; lateral
veins 710 pairs, very faint below, invisible above; petiole 0.20.9 cm long.
Syconia ripening red, globose, 0.70.8 cm diameter, apex umbonate; peduncle
0.10.7 cm long; basal bracts tiny, persistent.
Endemic to Borneo. Known in Sabah from Ranau and Kota Belud districts, and
in Sarawak from Marudi district (Mt. Dulit and Mt. Murud) and Kapit district
(Mt. Hose). In hill mixed dipterocarp to lower montane forests, at 10001800
m altitude.
Leaves of different shapes, very much wider........3
3.
Lateral veins 1315 pairs.............

var. riparia Corner
(Latin, riparius = frequenting riverbanks)
Gard. Bull. Sing. 17 (1960) 426, l.c. (1965) 38, l.c. (1969) 316; Coode et al. (eds.) l.c. 216.
Type: SFN 6871, Borneo, Sarawak, Marudi district, Batang Baram (holotype SING).
Small tree. Leaves elliptic to lanceolate, 312 0.54 cm; base cuneate, margin
294
entire, plane, apex obtuse or acute; midrib raised above; lateral veins 616 pairs,
visible below, invisible above; petiole 0.21.3 cm long. Syconia subglobose, 0.5
0.6 cm diameter; peduncle 1.52 cm long.
Endemic to Borneo. Common and widely distributed in Sarawak, not yet reported
for Sabah. Also in Brunei. In lowland mixed dipterocarp forest to lower montane
forest, up to 1800 m altitude.
Lateral veins 710 pairs, often indistinct..............4
4.
Leaves drying greenish yellow; lateral veins indistinct. Confined to limestone forest..
var. calcicola Kochummen
(Latin, calcicolus = inbabiting chalky soil)
Gard. Bull. Sing. 50 (1998) 215. Type: Lee S. 38626, Borneo, Sarawak, Bau district, G. Doya
(holotype KEP; isotypes SAN, SAR).
Small tree. Twigs grey-brown. Stipules linear, c. 0.4 cm long. Leaves drying
greenish yellow; elliptic, base cuneate, margin revolute, apex acute; midrib raised
above; lateral veins and intercostal venation very faint to inconspicuous; petiole
0.71 cm long. Syconia axillary or borne on twigs behind the leaves, ellipsoid
to subovoid, c. 0.3 cm diameter, apex umbonate; peduncle 0.30.4 cm long.
Endemic to Borneo. Recorded only from Sarawak, locally frequent. In limestone
forest.
Leaves drying brownish; lateral veins distinct. Not recorded from limestone forest.....
var. memecylifolia Corner
(Latin, memecylifolius = with leaves resembling those of Memecylon,
Melastomataceae)
Clemens 27939, Borneo, Sabah, Mt. Kinabalu (holotype BM).
Small tree. Leaves drying brownish; lanceolate to elliptic, 28 0.72.7 cm, base
cuneate, margin entire, plane, apex acute; midrib raised above; lateral veins 710
pairs, distinct below, joining near the leaf margin to form distinct intramarginal vein;
intercostal venation reticulate, faint below, invisible above; petiole up to 1 cm long.
Syconia axillary, subglobose to ellipsoid, 0.30.6 cm diameter; peduncle 0.31.5
cm long.
Borneo and Sulawesi. Common in Sabah and Sarawak. In hill forest to upper
montane forest, at 8002000 m altitude.
5.
Leaves obovate, apex rounded...........

var. myrsinoides Corner
(Greek, resembling Myrsine, Myrsinaceae; probably referring to the leaves)
SFN 27550, Borneo, Sabah, Mt. Kinabalu (holotype SING).
Tree up to 13 m tall. Leaves obovate, 2.59 0.83 cm, base cuneate, margin
entire, revolute, apex rounded; petiole 0.11 cm long. Syconia axillary or borne
on twigs behind the leaves, ellipsoid or subglobose, 0.50.6 cm diameter; peduncle
0.150.8 cm long.
Endemic to Borneo. Very rare, reported only from Mt. Kinabalu in Sabah. In
upper montane forest, up to 2100 m altitude.
Leaves elliptic or oblong, apex obtuse.............6
6.
Lateral veins 920 pairs. Syconia with peduncle 11.5 cm long.......
295
var. valida Corner

(Latin, validus = strong, robust-growing; the habit)
Phil. Trans. Roy Soc. London Ser. B, 256 (1969) 313; Coode et al. (eds.) l.c. 216. Type: S.
19068, Borneo, Sarawak (L).
Small tree. Leaves thick-coriaceous; oblong or lanceolate 4.513 1.54 cm,

base attenuate, margin distinctly revolute, apex usually obtuse, rarely acute; basal
glands distinct on the lower surface near petiole; midrib raised above; lateral
veins 920 pairs, with thin and shorter veins in between, looping and joining
near the leaf margin; intercostal venation invisible; petiole 0.31 cm long. Syconia
axillary, subglobose, 0.70.8 cm diameter; peduncle 11.5 cm long; basal bracts
persistent.
Endemic to Borneo; common in Sarawak, not yet reported for Sabah. Also in
Brunei. In hill mixed dipterocarp to upper montane forests, at 8002200 m altitude.
Lateral veins fewer than 9 pairs. Syconia with peduncle up to 0.6 cm long...
var. oleifolia
Sumatra and Borneo. In mossy lower to upper montane forests, at 12502400 m
altitude.
46. Ficus paramorpha Corner

(Greek, para- = similar to, -morphus = -shaped; inference unknown)
Gard. Bull. Sing. 17 (1960) 440, l.c. (1965) 49; Whitmore, Tantra & Sutisna l.c. 251. Type: Clemens
31777, Borneo, Sabah, Mt. Kinabalu, Upper Kinataki R. (holotype L).
Dioecious small tree. Twigs c. 0.3 cm thick. Stipules dark brown hairy, c. 0.7 cm long.
Leaves spirally arranged, somewhat chartaceous, lower surface dark brown hairy, upper
surface scabrous; elliptic, 614 35.5 cm, base symmetric, cuneate, margin toothed,
apex acute; midrib raised above; lateral veins 56 pairs, strongly raised below, basal
pair reaching up to 1/3 the length of the blade, with a gland in each axil below; intercostal
venation reticulate, raised below; petiole 1.22.5 cm long. Syconia axillary, in pairs,
sessile, ripening red, appressed dark brown hairy, subglobose, up to 1 cm diameter; basal
bracts ovate, up to 1.5 mm long; internal bristles numerous, golden yellow, up to 2.5
mm long. Tepals 34, white, hairy. Achenes shortly oblong, c. 1 mm long, somewhat
rough, slightly keeled.
Distribution. Endemic to Borneo. Very rare, known by 2 collections only from Mt. Kinabalu
in Sabah.
Ecology. In lower to upper montane forests, at 16002200 m altitude.
47. Ficus pseudotarennifolia Kochummen

(Greek, resembling but not equalling F. tarennifolia)
Gard. Bull. Sing. 50 (1998) 210. Type: S. 51027, Borneo, Sarawak, Marudi district, Bario (holotype
KEP; isotypes SAN, SAR).
Dioecious tree, up to 6 m tall. Twigs solid, c. 0.3 cm thick, reddish brown, angled.
Stipules lanceolate, acute, 1.51.7 cm long, caducous. Leaves opposite, thin-coriaceous,
296
glabrous; narrowly oblanceolate, oblong or elliptic, 9.52.5 13.5 cm, base cuneate,
margin entire, revolute, apex acute; midrib raised above; lateral veins 813 pairs, often
with glands in the axils below, curving and joining near the leaf margin, visible below,
very faint above; intercostal venation reticulate, visible below, almost invisible above;
petiole 12.5 cm long. Syconia borne on stem, in clusters, pear-shaped, c. 2 1.2 cm,
apex depressed; peduncle to 1.2 cm long; basal bracts tiny, persistent. Perianth whitish,
cup-shaped, irregularly lobed.
Distribution. Endemic to Borneo. Rare, recorded only for Sarawak by 2 collections, S.
11929 from Tatau district and the type.
Ecology. In lowland and hill mixed dipterocarp forests, up to 900 m altitude, by streams.
48. Ficus pubinervis Blume

(Latin, pubi- = softly hairy, nervus = vein; the soft-hairy midrib below)
Bijdr. Fl. Ned. Ind. (1825) 452; King l.c. (1887) 54; Merrill l.c. (1921) 226; Masamune l.c. 248; Backer
& Bakhuizen f. l.c. 16; Corner (1965) l.c. 31; Whitmore, Tantra & Sutisna l.c.252. Type: Blume s.n. (=
RHL Sheet No. 908188457), Java (holotype L). Synonyms: F. decaisanei Steud., Nom. Bot. ed. 2, 1
(1840) 636; F. similis Merr. Philip. J. Sc. 1 (1906) Suppl. 47; F. crassitora Elmer, Leafl. Philip. Bot. 2
(1908) 548.
Monoecious tree, up to 35 m; buttresses up to 2 m tall. Bark dark grey. Young parts

pubescent. Stipules lanceolate, 2.53.5 cm long, hairy outside, caducous. Leaves spirally
arranged, thin-coriaceous, glabrous except for the hairy midrib below; ovate, elliptic or
lanceolate, 9.513.5 45 cm, base attenuate, margin entire, plane, apex acute or acuminate;
midrib raised on both surfaces; lateral veins 57 pairs, looping and joining near the
leaf margin, with shorter veins in between, distinct below, basal pair short, mostly with
a gland in each axil below; intercostal venation reticulate; petiole 0.51.2 cm long, roughhairy when young. Syconia axillary, solitary or in pairs, sessile or short-peduncled, ripening
red, subglobose, c. 1.3 cm diameter, hairy, apex umbonate; basal bracts 3, ovate, acute,
11.5 mm long; internal bristles absent. Tepals 24, red, glabrous, free or basally united.
Achenes c. 1 mm long, slightly keeled.
Distribution. Sumatra, Java, Borneo, Philippines, Sulawesi, Maluku, Sumbawa, and Flores.
Very rare in Sabah, known by 1 collection, SAN 28950, from Mt. Kinabalu, not yet reported
for Sarawak.
Ecology. In lower montane forest.
49. Ficus racemosa L.

(Latin, racemosus = in a raceme-like bunch; the syconia)
Sp. Pl. (1753) 922; Corner l.c. (1965) 34; Kochummen l.c. 154; Primack l.c. 98; Turner l.c. 353. Type:
unknown. Synonyms: F. glomerata Roxb., Pl. Corom. 2 (1798) 13; Covellia glomerata (Roxb.) Miq.
l.c. (1848) 465; F. vesca F.v.M. ex Miq. l.c. (1861) 243.
297
Distribution. Asia and Australia. Only var. elongata is present in Sabah and Sarawak.
var. elongata (King) Barrett
(Latin, elongatus = elongate; inference unknown)
Bull. Torr. Bot. Club 73 (1946) 323; Corner l.c. (1965) 35; Backer & Bakhuizen f. l.c. 27; Kochummen
l.c. 154; Anderson l.c. 260; Whitmore, Tantra & Sutisna l.c. 252. Basionym: F. glomerata Roxb. var.
elongata King l.c. (1888) 174. Type: Kurz s.n., Burma, Chittagong (holotype CAL?). Synonyms: F.
lucescens Blume l.c. 444; F. lanceolata Buch.-Ham. ex Roxb., Fl. Ind., ed. Carey, 3 (1832) 557; Covellia
lanceolata (Buch.-Ham. ex Roxb.) Miq. l.c. (1848) 465; Sycomorus riparia Hochst. ex Miq. l.c. (1848)
114; Urostigma lucescens (Blume) Miq. l.c. (1859) 341; F. riparia (Hochst. ex Miq.) Miq. l.c. (1867)
295; F. acidula King l.c. (1888) 176; F. henrici King l.c. (1888) 176.
Monoecious tree, up to 18 m tall, 25 cm diameter. Bark greenish white, smooth; inner

bark yellowish. Sapwood whitish. Twigs slender, reddish brown, c. 0.2 cm thick, Stipules
narrowly lanceolate, up to 4 cm long, glabrous, often persistent. Leaves distichous,
membranous, glabrous, pale glaucous below; oblong, 511 1.53.5 cm, base broadly
cuneate, margin entire, plane, faintly undulate, apex obtuse; midrib raised above; lateral
veins 1015 pairs, curved, very faint on both surfaces, with shorter veins in between,
basal pair short, often with a gland in each axil below; intercostal venation reticulate,
faint; petiole slender, 14 cm long. Syconia borne on long leafless branched twigs arising
from stem and older branches, ripening red, pear-shaped, 11.5 cm diameter, apex
depressed; peduncle 0.31.2 cm long; basal bracts ovate, triangular, 12 mm long; internal
bristles absent. Tepals 34, red, glabrous, united at the base. Achenes lenticular, c. 1
mm long, smooth, not or scarcely keeled.
Distribution. Abyssinia, India, Burma, Sumatra, Peninsular Malaysia, Java, Borneo,
Sulawesi, and Lesser Sunda Islands. Uncommon in Sabah and Sarawak, mainly reported
for Sabah; from Sarawak known by a single collection, S. 50082, from Kapit district.
Not known in Brunei. Also in Kalimantan.
Ecology. In lowland forest, in open area by streams.
50. Ficus retusa L.

(Latin, retusus = with rounded or notched end; the leaf apex)
Mant. (1767) 129; King l.c. (1887) 50; Merrill l.c. (1921) 226; Burkill l.c. 1030; Backer & Bakhuizen f.
l.c. 35; Corner l.c. (1965) 20; Kochummen l.c. 154; Anderson l.c. 260; Primack l.c. 99; Whitmore,
Tantra & Sutisna l.c. 253; Turner l.c. 353; Coode et al. (eds.) l.c. 217. Type: unknown.
Monoecious tree, up to 15 m tall. Twigs 0.30.6 cm thick, pale yellowish or grey-brown,

distinctly grooved or with prominent stipular and petiolar scars (var. borneensis). Stipules
small and caducous or large and persistent (var. borneensis), ovate to oblong, 13
0.51.5 cm. Leaves spirally arranged, thin- to thick-coriaceous, glabrous, sometimes
slightly glaucous below; oblanceolate to narrowly obovate, 516 2.26 cm, base cuneate,
margin entire, plane (revolute in var. borneensis), apex acute, obtuse or acuminate, acumen
up to 1 cm long; midrib sunken above; lateral veins 58 pairs, sunken above, raised
below, curving and joining near the leaf margin to form distinct intramarginal vein, basal
298
pair reaching more than 1/2 the length of the blade, without glands in the axils below;
intercostal venation reticulate, sunken above; petiole 0.82.5 cm long, grooved on the
adaxial side. Syconia in pairs, axillary, sessile, ripening yellowish red, depressed globose,
0.50.6 cm diameter, apex umbonate; basal bracts ovate, 56 35 mm, or much smaller,
persistent; internal bristles absent. Tepals 24, spathulate or lanceolate. Achenes tuberculate.
Vernacular name. Sarawakara (Iban).
Distribution. Sumatra, Peninsular Malaysia, Java, and Borneo. Uncommon in Sabah and
Key to varieties
Leaves thin-coriaceous; stipules c. 1 0.5 cm, caducous. Basal bracts of syconia
indistinct.......var. retusa
Leaves thick-coriaceous; stipules c. 3 1.5 cm, persistent at the apex of twigs. Basal
bracts conspicuous, covering the lower half of syconium........
var. borneensis Corner
(of Borneo)
Gard. Bull. Sing. 17 (1960) 393, l.c. (1965) 20; Whitmore, Tantra & Sutisna l.c. 253; Coode et al.
(eds.) l.c. 217. Type: Hallier 1074, Borneo, Kalimantan, Sg. Sambas (holotype L).
Twig grey-brown, c. 1 cm thick, with prominent stipular and petiolar scars. Stipules
ovate, acute, c. 3 1.5 cm, persistent. Leaves thick-coriaceous, pale glaucous below.
Syconia with distinct basal bracts covering the lower half.
Endemic to Borneo. Rare in Sabah and Sarawak. Also in Brunei and Kalimantan.
In lowland mixed dipterocarp forest.
51. Ficus rubrocuspidata Corner

(Latin, rubrum = red, cuspidatus = sharp and narrowly pointed; the leaf apex)
Gard. Bull. Sing. 17 (1960) 483, l.c. (1965) 79; Primack l.c. 100; Whitmore, Tantra & Sutisna l.c. 253;
Coode et al. (eds.) l.c. 217. Type: SFN 27323, Borneo, Sabah, Mt. Kinabalu (holotype SING).
Dioecious small tree. Twigs c. 0.15 cm thick, brownish. Stipules linear to lanceolate,
up to 0.8 cm long, caducous. Leaves distichous, membranous, glabrous, drying greenish;
elliptic to obovate or narrowly oblong, 523 1.27 cm, base symmetric, cuneate, margin
entire, plane, apex cuspidate, acumen up to 4.5 cm long; midrib raised above; lateral
veins 510 pairs, well-spaced, curved, visible on both surfaces, looping and joining near
the leaf margin, basal pair short, inconspicuous, departing from the midrib at a broad
angle, with a gland in the axil of one of the pair; intercostal venation reticulate, visible
on both surfaces; petiole 0.20.6 cm long. Syconia paired or clustered, axillary or borne
on twigs behind the leaves, ripening orange to red, subglobose, c. 0.4 cm diameter, sessile
or short-peduncled, without basal bracts; internal bristles absent. Tepals 4, red, free,
glabrous. Achenes c. 0.8 mm long, scarcely keeled.
299
Vernacular name. Sarawakkara (Iban).

Distribution. Endemic to Borneo. Common in Sabah and Sarawak. Also in Brunei and
Kalimantan.
altitude, along rocky streams and on limestones.
52. Ficus rubromidotis Corner

(meaning obscure)
Gard. Bull. Sing. 17 (1960) 484, l.c. (1965) 79; Anderson l.c. 260; Primack l.c. 100; Whitmore, Tantra
& Sutisna l.c. 253. Type: Clemens 20699, Borneo, Sarawak, Bidi Cave Mountain (holotype K).
Dioecious small tree. Twigs c. 0.3 cm thick, yellowish, youngest angled. Stipules paired,
broadly lanceolate, 12 cm long, rough-hairy, persistent. Leaves distichous, coriaceous,
glabrous, drying yellowish green; oblong to elliptic, 1536 512.5 cm, base cuneate,
shortly auriculate on one side, margin entire, plane, apex acuminate or caudate, acumen
1.25 cm long; midrib raised above, sharply keeled below; lateral veins 1220 pairs,
curved, raised on both surfaces, looping and joining near the leaf margin, with few shorter
veins in between, basal pair short, inconspicuous, with a gland in the axil of one or
both veins below; intercostal venation scalariform, visible below, faint above; petiole 0.3
1 cm long. Syconia axillary or in clusters on the stem and branches, sessile or peduncled,
ripening orange to red, subglobose, 0.50.7 cm diameter, apex umbonate; basal bracts
23, minute, lateral bracts absent; internal bristles absent. Tepals 4, red or red-brown,
oblong, glabrous, free. Achenes oblong, c. 1 mm long, smooth, slightly keeled at apex.
Vernacular name. Sarawakkara (Iban).
Distribution. Endemic to Borneo. Common in Sarawak; very rare in Sabah and known
by a single collection, SAN 90622, from Ranau district. Not known in Brunei and
Kalimantan.
Ecology. In lowland and hill mixed dipterocarp forests, including limestone forest, by
streams.
53. Ficus ruficaulis Merr.

(Latin, rufus = reddish, caulis = stem or branch; with reddish branch)
Philip. Gov. Lab. Pub. 17 (1904) 13; Corner l.c. (1965) 47; Anderson l.c. 260; Primack l.c. 101; Whitmore,
Tantra & Sutisna l.c. 253. Type: unknown. Synonyms: F. gerontocarpa Warb. in Perkin, Fragm. Fl.
Philip. 3 (1905) 201; F. paloensis Elmer, Leafl. Philip. Bot. 2 (1908) 547; F. zambalensis Elmer, Leafl.
Philip. Bot. 9 (1937) 3207; F. ruficaulis Merr. var. paloensis (Elmer) Elmer, Leafl. Philip. Bot. 1 (1906)
203.
Dioecious tree, up to 18 m tall. Bark greyish brown, smooth. Twigs stout, 0.51 cm
thick, reddish brown, sparsely brownish rough-hairy. Stipules lanceolate, 1.22 cm long,
300
velvety brownish hairy. Leaves spirally arranged, thin-coriaceous, sparsely brownish roughhairy on both surfaces; ovate to oblong, 1828 715.5 cm, base cordate, rounded to
cuneate, margin entire, plane, apex acuminate, acumen up to 1.5 cm long; midrib raised
above; lateral veins 6 pairs or more, with shorter veins in between, raised on both surfaces,
basal veins reaching more than 1/2 the length of the blade, without axillary glands;
intercostal venation scalariform, visible below, faint above; petiole 3.56 cm long. Syconia
paired, axillary or borne on twigs behind the leaves, ripening green or yellow, ovoid
to subglobose, 0.70.8 cm diameter, apex umbonate; peduncle 0.70.8 cm long; basal
bracts c. 3 2 mm, persistent; internal bristles abundant, yellowish, 11.5 mm long.
Tepals 35, spathulate, free. Achenes c. 1.2 mm long, strongly keeled, tuberculate.
Distribution. Taiwan, Borneo, the Philippines, Sulawesi, and New Guinea. Very rare in
Sabah and Sarawak, known by 1 collection from Sabah (SAN 83127) and two collections
from Sarawak (S. 19191 and S. 37344). Not known in Brunei and Kalimantan.
Ecology. In hill to lower montane forests, at 7001200 m altitude.
54. Ficus schwarzii Koord.

(J.A.T. Schwarz, sometime teacher-missionary in Sulawesi)
Verl. Bot. Diens. Minahasa (1898) 607; Corner l.c. (1965) 89, l.c. (1988) 552; Kochummen l.c. 155;
Anderson l.c. 260; Primack l.c. 103; Whitmore, Tantra & Sutisna l.c. 254; Coode et al. (eds.) l.c. 217.
Type: Koorders s.n., Sulawesi, Minahasa, G. Lengkoan (BO?, L?).
Dioecious tree, up to 20 m tall, 20 cm diameter. Bark dark grey, smooth. Twigs 0.15
0.25 cm thick, grey-brown, young ones angled. Stipules lanceolate, up to 1.5 cm long,
caducous. Leaves distichous, membranous, glabrous; elliptic to oblanceoate, 820 2.5
6.5 cm, base cuneate, symmetric, margin entire, plane, apex caudate, acumen up to 2.5
cm long; midrib flat above; lateral veins 59 pairs, well-spaced, curved, raised below,
faint above, basal pair short, inconspicuous, without axillary glands; intercostal venation
scalariform to reticulate, visible on both surfaces; petiole 0.51 cm long. Syconia borne
in massive clusters on branched leafless twigs arising from the stem, pear-shaped, 2.5
3.75 cm diameter, ripening reddish brown, apex depressed with projecting scales; peduncle
13 cm long; basal bracts 3, 13 mm long; internal bristles absent or very few. Tepals
united, tubular or collar-like. Achenes c. 1 mm long, keeled, faintly tuberculate.
Vernacular names. Sabahperanak (Dusun). Sarawakara, engkabuloh, engkeruroh,
engkunoh (Iban), pipin (Bidayuh).
Distribution. Burma, Thailand, Sumatra, Peninsular Malaysia, Borneo, and Sulawesi. Very
common in Sarawak but rare in Sabah. Also in Brunei and Kalimantan.
Ecology. In lowland and hill forests by streams, up to 900 m altitude.
Use. In Sarawak, the latex is used in traditional medicine to treat ring-worms.
301
55. Ficus scortechinii King.

(Reverend Scortechini, 18451886, a Roman Catholic missionary; 18841886 government
botanist stationed in Taiping, Perak)
Ann. Roy. Bot. Gard. Calc. 1, 2 (1888) 112; Ridley l.c. (1924) 343; Corner l.c. (1965) 94; Kochummen
l.c. 156; Anderson l.c. 260; Turner l.c. 353. Type: Kunstler 934, Peninsular Malaysia, Perak (CAL,
K?).
Distribution. Burma, Thailand, Peninsular Malaysia, Singapore, and Borneo. Only var.
lanceata is recorded for Sabah and Sarawak.
var. lanceata Corner
(Latin, lanceatus = lance-shaped; the leaves)
Type: RSNB 2815, Borneo, Sabah, Mt. Kinabalu, Tenompok (holotype K).
Dioecious small tree, up to 6 m tall. Twigs c. 0.3 cm thick, solid, brown. Leaves spirally
arranged or opposite, thin-coriaceous, glabrous; lanceolate, 620 1.64 cm, base cuneate,
often slightly asymmetric, margin faintly toothed, apex acute, acuminate or obtuse; midrib
raised above; lateral veins 610 pairs, basal pairs short; intercostal venation reticulate;
petiole 0.51 cm long, often with a small subnodal gland. Syconia borne in dense clusters
on the stem, rarely on the main branches, or borne on twigs behind the leaves, ripening
red, obovoid, 0.71 cm diameter, surface ridged and irregularly tuberculate, apex depressed;
peduncle 12 cm long; basal bracts 0.51 mm long, lateral bracts absent; internal bristles
sparse, brownish. Perianth cup-shaped. Achenes up to 0.8 mm long, slightly keeled,
tuberculate.
Vernacular name. Sarawakengkunoh (Iban).
Distribution. Endemic to Borneo. Very rare, known only from Mt. Kinabalu in Sabah
(the type), and by S. 19388 from Sg. Anap, Tatau district, in Sarawak.
Ecology. In hill mixed dipterocarp forest to lower montane forest, up to 1500 m altitude.
56. Ficus septica Burm. f.
Fig. 11.
(Greek, septikos = putrefying; the sap)

Fl. Ind. (I768) 226; Miq. l.c. (1859) 311; Merrill l.c. (1921) 226; Burkill l.c. 1032; Masamune l.c.
250; Backer & Bakhuizen f. l.c. 32; Corner l.c. (1965) 92; Primack l.c. 104; Whitmore, Tantra &
Sutisna l.c. 254. Type: unknown. Synonyms: F. leucopleura Blume l.c. 443; F. laccifera Blanco, Fl.
Philip. (1837) 684; Covellia stictocarpa Miq. l.c. (1851) 65; F. stictocarpa (Miq.) Miq. l.c. (1867)
284; F. philippinensis Bonard ex Herincq., Hort. France (1869) 244; F. didymophylla Warb. in Perkin,
Fragm. Fl. Philip. 3 (1905) 200.
Dioecious tree, up to 25 m tall, 20 cm diameter. Bark pale grey. Twigs hollow, 0.2
0.4 cm thick, reddish brown. Stipules lanceolate, acute, 23 cm long, caducous. Leaves
spirally arranged, thin-coriaceous, glabrous; elliptic to oblong, 1028 3.711 cm, base
302
vi
303
broadly cuneate, margin entire, plane, apex acute; midrib pinkish on drying, above often
sunken on the lower half, raised or flat on the upper half; lateral veins 812 pairs, raised
on both surfaces, with few shorter veins in between, curving and joining near the leaf
margin, basal pairs inconspicuous, extending less than 1/3 the length of the leaf blade,
without axillary glands; intercostal venation finely reticulate, visible on both surfaces;
petiole 14.5 cm long. Syconia axillary or borne on twigs behind the leaves, subhemispherical
with depressed apex, 1.22 cm diameter, with pale dots and distinct ridges, ripening purplish;
peduncle up to 1.2 cm long; basal bracts 3, subacute, 12 mm long; internal bristles
short, few, white. Tepals 23, united. Achenes up to 1 mm long, slightly keeled, minutely
tuberculate.
Vernacular name. Sarawakuok (Kelabit).
Distribution. Ryu-Kyu, Taiwan, Sumatra, Java, Borneo, the Philippines, Sulawesi, Maluku,
the Lesser Sunda Islands, New Guinea, Admiralty Islands, New Britain, New Ireland,
Solomon Islands, New Hebrides, Santa Cruz Islands, and Queensland, Australia. Very
common in Sabah, not yet reported for Sarawak. Not known in Brunei.
Ecology. In lowland forest to lower montane forest, up to 1200 m altitude.
57. Ficus setiflora Stapf

(Latin, seta = bristles, florus = flower; the abundant bristles in between the flowers)
FMK (1894) 226; Merrill l.c. (1921) 226; Masamune l.c. 250; Corner l.c. (1965) 50; Anderson l.c. 260;
Primack l.c. 104; Whitmore, Tantra & Sutisna l.c. 254; Coode et al. (eds.) l.c. 217. Type: Haviland
1214, Borneo, Sabah, Mt. Kinabalu (K).
Dioecious shrub or small tree, up to 5 m tall. Twigs c. 0.2 cm thick, thinly appressed
pale brown hairy. Stipules lanceolate, 0.30.9 cm long, densely fawn-sericeous, caducous.
Leaves spirally arranged, membranous, sparsely to densely appressed pale brown hairy
on both surfaces; elliptic, oblong or lanceolate, 5.519 26 cm, base symmetric, cuneate,
margin coarsely serrate, apex acuminate, acumen up to 2 cm long; midrib raised above;
lateral veins 47 pairs, with shorter veins in between, faintly raised above, raised below,
basal pair reaching almost 1/2 the length of the blade, with a gland in each axil below;
intercostal venation reticulate, visible below, faint above; petiole 0.52 cm long, hairy.
Syconia axillary, in pairs, ripening rose-red, subglobose or oblong, 0.61 cm diameter,
surface sparsely hairy, apex umbonate; peduncle up to 0.1 cm long; basal bracts small,
hairy, persistent, lateral bracts absent; internal bristles numerous, up to 2 mm long. Tepals
34, white, hairy. Achenes 11.2 mm long, slightly keeled, rough.
Distribution. Endemic to Borneo. Uncommon in Sabah and Sarawak. Also in Brunei
and Kalimantan.
Ecology. In hill mixed dipterocarp forest to upper montane forest, up to 2400 m altitude.
Key to varieties
1.
Leaves sparsely hairy below...............var. setiflora

Leaves densely hairy below......2
304
2.
Indumentum comprising hairs 0.51 mm long. Styles in female flowers hairy..............

var. adelpha Corner
(Greek, adelphos = joined together; probably referring to the stamens)
al. (eds.) l.c. 217. Type: SFN 27876, Borneo, Sabah, Mt. Kinabalu (holotype SING).
Uncommon in Sabah and Sarawak; in montane forest.

Indumentum comprising hairs up to 0.5 mm long. Styles in female flowers glabrous..
var. puberula Corner
(Latin, peberulus = downy with very short soft hairs; the twigs and petioles)
al. (eds.) l.c. 217. Type: Endert 4275, Borneo, Kalimantan, W Kutai, Mt. Kemul (holotype L).
Very rare, recorded only for Sarawak; in montane forest.
58. Ficus sinuata Thunb.

(Latin, sinuatus = with wavy margin; the leaves)
Diss. Fic. (1786) 12; Corner l.c. (1965) 76, l.c. (1988) 554; Backer & Bakhuizen f. l.c. 26; Anderson l.c.
260; Kochummen l.c. 156; Primack l.c. 105; Whitmore, Tantra & Sutisna l.c. 254; Turner l.c. 354. Type:
Thunberg s.n., Java (UPS). Synonyms: F. raridens Miq. l.c. (1848) 430; F. obtusidens Miq. l.c. (1851)
59; F. angulidens Miq. l.c. (1859) 310; F. radicans Roxb. var. angulosa Miq. l.c. (1859) 306.
Dioecious small tree, up to 6 m tall, with drooping branches. Bark yellowish. Twigs
c. 0.2 cm thick, glabrous, yellowish. Stipules narrowly lanceolate, 0.50.7 cm long,
caducous. Leaves distichous, thin-coriaceous, glabrous; elliptic to oblong, 9.523 3.5
9 cm, base often slightly asymmetric, cuneate, margin sinuate, slightly crenate, or deeply
lobed, apex caudate, acumen up to 2 cm long; midrib raised above, sharply keeled below;
lateral veins 58 pairs, with shorter veins in between, curving and joining near the leaf
margin, distinct below, faint above, basal pair extending up to 1/3 the length of the blade,
with a gland in each axil below; intercostal venation reticulate, distinct below, visible
above; petiole 0.32 cm long. Syconia axillary, solitary or in pairs, or borne in clusters
on the twigs behind the leaves, ripening pinkish, orange to red, subglobose, c. 0.61
cm diameter, base narrowed into a slender stalk, without distinct peduncle; basal bracts
23, small; internal bristles absent. Tepals 25, shortly united at the base, glabrous, white
or red-speckled. Achenes 1.52 mm long, compressed, smooth, strongly keeled at the
apex.
Vernacular name. Sarawakterbudak (Kelabit).
Distribution. Assam, Thailand, Sumatra, Peninsular Malaysia, Java, and Borneo. Very
common in Sabah and Sarawak. Not known in Brunei.
Ecology. In hill forest.
59. Ficus spathulifolia Corner

(Latin, spathulatus = spathula-shaped, folium = leaf; with spathulate leaves)
305
Gard. Bull. Sing. 17 (1960) 391, l.c. (1965) 19; Kochummen l.c. 156; Anderson l.c. 261; Primack l.c.
105; Whitmore, Tantra & Sutisna l.c. 255; Coode et al. (eds.) l.c. 218. Type: SFN 33970, Peninsular
Malaysia, Selangor, Klang (holotype SING). Synonym: F. spathulifolia Corner var. stipitata Corner l.c.
17 (1960) 392, syn. nov.
Monoecious tree, up to 13 m tall, 15 cm diameter. Twigs 0.30.5 cm thick, grey-brown,

strongly ridged, with prominent stipular scars. Stipules hairy, oblong, acute, c. 1 cm long,
caducous. Leaves spirally arranged, coriaceous, glabrous; obovate to spathulate, 38
1.34 cm, base cuneate, margin entire, plane, apex rounded; midrib sunken above at
the lower half; lateral veins 35 pairs, raised below, flat above, looping and joining near
the leaf margin, basal pair reaching up to 1/2 the length of the blade, without axillary
glands; intercostal venation invisible or rarely faintly visible below; petiole 0.81.5 cm
long, flat on the adaxial side. Syconia in pairs, axillary, sessile, ripening yellow to red,
subglobose to subovoid, 0.60.7 cm diameter, apex umbonate; basal bracts up to 5 mm
long, persistent; lateral bracts absent; internal bristles absent. Tepals 23, free. Achenes
oblong, c. 1 mm long, slightly angled.
Vernacular name. Sarawakara (Iban).
Distribution. Indo-China, Peninsular Malaysia and Borneo. Widely distributed in Sabah
and Sarawak. Also in Brunei and Kalimantan.
Ecology. In Sabah and Sarawak mainly in peatswamp and kerangas forests.
60. Ficus stolonifera King

(Latin, stolon = underground runner, ferus = bearing; producing fig-bearing stolons)
Ann. Roy. Bot. Gard. Cal. 1, 2 (1888) 104; Merrill l.c. (1921) 226; Masamune l.c. 250; Corner l.c.
(1965) 91; Anderson l.c. 261; Primack l.c. 106; Whitmore, Tantra & Sutisna l.c. 255; Coode et al. (eds.)
l.c. 218. Type: Beccari PB 2799, Borneo, Sarawak (holotype FI).
Dioecious tree, up to 24 m tall, 30 cm diameter. Twigs up to 0.3 cm thick, yellowish

brown hairy. Stipules narrowly lanceolate, c. 1.2 cm long, appressed hairy, caducous.
Leaves distichous, rarely opposite, coriaceous, rough-hairy on both surfaces; elliptic to
oblong, 8.527 4.59 cm, base broadly cuneate, symmetric, margin denticulate to crenate,
apex acuminate, acumen 23 cm long; midrib flat above; lateral veins 47 pairs, with
shorter veins in between, raised below, visible above, basal pair short; intercostal venation
scalariform to reticulate, raised below, faint above; petiole, 0.30.6 cm long, yellowish
brown appressed hairy, with a small subnodal gland. Syconia borne on stolons, subsessile,
finely appressed hairy, ripening reddish pink, subglobose, c. 0.7 cm diameter; peduncle
0.10.3 cm long; basal bracts 3, lateral bracts few; internal bristles absent. Perianth
tubular or collar-like. Achenes c. 0.7 mm long, subcompressed, smooth, slightly keeled.
Vernacular name. Sarawakentimau (Iban).
Distribution. Endemic to Borneo. Uncommon in Sabah and Sarawak. Also in Brunei
and Kalimantan.
306

altitude.
61. Ficus subfulva Corner

(Latin, sub- = somewhat, fulvus = tawny; the indumentum on the twigs, petioles and veins of
leaves beneath)
Gard. Bull. Sing. 17 (1960) 432, l.c. (1965) 47; Primack l.c. 108; Whitmore, Tantra & Sutisna l.c.
255. Type: Hallier 2349, Borneo, Kalimantan, G. Klam (holotype L).
Dioecious tree, up to 20 m tall. Twigs 0.40.6 cm thick, yellowish brown hairy. Stipules
lanceolate, up to 1.5 cm long, yellowish brown appressed hairy, caducous. Leaves spirally
arranged, rigid-coriaceous, sparsely or densely yellowish brown velvety hairy on both
surfaces; elliptic, 1025 511.5 cm, base rounded to subcordate, or cuneate, margin
entire, revolute, apex subacuminate, acumen up to 1 cm long; midrib flat above; lateral
veins 46 pairs, with a number of shorter veins in between, raised below, basal veins
reaching up to 2/3 the length of the blade, with a gland in each axil below; intercostal
venation reticulate, distinct beneath; petiole 1.25 cm long, hairy. Syconia axillary, in
pairs, thinly appressed yellowish brown hairy, ripening orange-red, globose or ellipsoid,
11.2 cm diameter; peduncle 0.41 cm long; basal bracts ovate, acute, persistent, hairy,
lateral bracts absent; internal bristles abundant, 24 mm long, stiff, yellow. Tepals glabrous,
spathulate, red. Achenes c. 1.5 mm long, tuberculate.
Distribution. Endemic to Borneo. Very rare in Sabah and Sarawak. Also in Kalimantan.
Ecology. In lowland and hill mixed dipterocarp forests, at 3001000 m altitude, including
limestone habitat.
Key to varieties
Leaf undersurface densely velvety hairy...............
var. villosula Corner
(Latin, villosulus = finely shaggy hairy; the leaf under surface)
Gard. Bull. Sing. 17 (1960) 432, l.c. (1965) 475, Whitmore, Tantra & Sutisna l.c. 256. Type:
Hallier 2869, Borneo, Kalimantan, Lianggagang (holotype L).
Endemic to Borneo. Very rare in Sarawak; in lowland mixed dipterocarp forest; not
yet reported for Sabah. Also in Kalimantan.
Leaf undersurface sparsely velvety hairy................var. subfulva
62. Ficus subterranea Corner

(Latin, subterraneus = underground; the position of the syconia)
l.c. 218. Type: Corner BRUN 5327, Borneo, Brunei, Amo, Sg. Belalong (holotype CGE; isotypes K,
SAR).
307
Dioecious shrub or small tree, up to 5 m tall. Twigs up to 0.2 cm thick, appressed pubescent
with whitish to pale yellowish brown hairs. Stipules 1.53 cm long, appressed hairy,
caducous. Leaves distichous, membranous, upper surface glabrous, smooth or scabrous,
lower surface yellowish brown appressed hairy; narrowly elliptic to obovate, 1126
38 cm, base cuneate, margin entire, plane, apex acuminate, acumen 12.5 cm long;
midrib flat above; lateral veins 1115 pairs, with shorter veins in between, faint above,
raised beneath, distal ones with axillary glands, basal pair conspicuous, departing from
the midrib at an acute angle; intercostal venation finely scalariform, visible below, invisible
above; petiole 0.51.5 cm long, appressed yellowish brown hairy. Syconia borne on
subterranean, branched, slender stolons arising from the base of stem, glabrous, ripening
pale pink, subglobose, c. 1.4 cm diameter; peduncle 0.10.3 cm long; lateral bracts
abundant, uncinate, up to 0.6 mm long; internal bristles absent. Perianth absent. Achenes
c. 0.6 mm long, slightly keeled, sparsely ridged.
Distribution. Endemic to Borneo. Very rare in Sabah and Sarawak. Also in Brunei.
63. Ficus tarennifolia Corner

(with leaves like that of Tarenna, Rubiaceae)
Type: SFN 27510, Borneo, Sabah, Mt. Kinabalu (holotype SING).
Dioecious tree, up to 13 m tall. Bark brown, smooth, lenticellate; inner bark yellowbrown. Sapwood yellow-brown. Twigs slender, up to 0.3 cm thick. Stipules lanceolate,
up to 2 cm long, caducous. Leaves usually opposite, sometimes spirally arranged,
membranous, glabrous; lanceolate, elliptic or obovate, 721 28 cm, base cuneate, margin
distantly finely toothed, apex acute; midrib flat above; lateral veins 518 pairs, curving
and joining near the leaf margin, visible below, invisible above, basal pair inconspicuous,
without glands in the axils; intercostal venation reticulate, visible below, invisible above;
petiole 0.63 cm long, appressed hairy. Syconia borne on short, leafless twigs arising
from the stem, and also on long stolons arising from the base of stem, reddish brown
hairy, ripening to pinkish brown, pear-shaped to subglobose, 1.21.5 11.3 cm, surface
more or less smooth, apex rounded; peduncle 0.51.2 cm long; basal bracts 3, up to
2 mm long, lateral bracts absent; internal bristles absent. Perianth tubular or collarlike. Achenes c. 1 mm long, slightly keeled, tuberculate.
Vernacular name. Sarawakbuah tepur (Kelabit).
Distribution. Endemic to Borneo; confined to Mt. Kinabalu in Sabah.
Ecology. In hill mixed dipterocarp forest to upper montane forest, at 10002500 m altitude.
64. Ficus tinctoria G. Forst.

(Latin, tinctorius = used in dying; the latex)
308
Prod. Fl. Ins. Austral. (1786)76; Corner l.c. 17 (1960) 475, l.c. 21 (1965) 74; Primack l.c. 112. Type:
unknown.
Distribution. Andaman Islands, S China, Formosa, Indo-China, Thailand, Sumatra, Peninsular

Malaysia, Java, Borneo, the Philippines, Sulawesi, Maluku, Lesser Sunda Islands, New
Guinea, N Australia, Micronesia, and Polynesia. Only subsp. gibbosa is known in Sabah
and Sarawak.
subsp. gibbosa (Blume) Corner
(Latin, gibbosus = more swollen in one place than another; the syconia)
Gard. Bull. Sing. 17 (1960) 476, l.c. (1965) 74, l.c. (1988) 544; Backer & Bakhuizen f. l.c. 25; Kochummen
l.c. 159; Anderson l.c. 261; Primack l.c. 112; Whitmore, Tantra & Sutisna l.c. 257; Turner l.c. 355.
Basionym: F. gibbosa Blume l.c. 466. Type: Blume s.n., Java (L?) Synonyms: F. cuneata Blume l.c.
465; F. paradoa Blume, l.c. 467; F. irregularis Steud., Nom. Bot. ed. 2, 1 (1840) 636; F. angulata Miq.
l.c. (1848) 434; F. gibbosa Blume var. cuneata (Blume) Miq. l.c. (1851) 62; F. subobliqua Miq. l.c.
(1859) 225; F. latiuscula Miq. l.c. (1859) 225; F. gibbosa Blume var. unigibba Miq. l.c. (1867) 276; F.
gibbosa Blume var. dodonaefolia Miq. l.c. (1867) 276; F. latifolia Miq. l.c. (1867) 276; F. antoniana
Elmer, Leafl. Philip. Bot. 4 (1912) 1374.
Dioecious tree, up to 15 m tall, or strangler. Twigs up to 0.3 cm thick, pale yellowish.

Stipules up to 1.8 cm long. Leaves distichous, coriaceous, glabrous, drying brown; elliptic,
lanceolate, or rhomboid, 411 2.55 cm, base cuneate, asymmetric, margin entire, plane
or revolute, apex acute to acuminate; midrib raised above; lateral veins 45 pairs, wellspaced, curved, looping and joining near the leaf margin, distinct below, faint above,
with shorter veins in between, basal pair inconspicuous, with a gland in the axil of the
vein on the broader side of leaf base; intercostal venation reticulate, visible below, faint
to invisible above; petiole 0.51 cm long. Syconia axillary, sessile or subsessile, ripening
golden yellow or red, gibbous, 0.71 cm diameter, surface faintly ridged, with apical
pore; basal bracts 3, up to 2 mm long, caducous, lateral bracts absent; internal bristles
numerous, minute. Tepals 35, oblong to lanceolate, white, puberulous, free. Achenes
oblong, up to 1.3 mm long, subcompressed, not or only slightly keeled.
Distribution. Andaman Islands, S China, Indo-China, Thailand, Sumatra, Peninsular
Malaysia, Java, Borneo, the Philippines (Palawan), Sulawesi, Halmahera, Banda, and
Sumbawa. Widely distributed in Sabah and Sarawak.
Ecology. In lowland forest and limestone forest.
Key to varieties
Syconia with peduncle 0.30.4 cm long. Leaf margin plane...........var. gibbosa
Syconia sessile. Leaf margin revolute....................
var. rigida (Miq.) Corner
(Latin, rigidus = rigid; the leaves)
Gard. Bull. Sing. 18 (1960) 476, l.c. (1965) 75. Basionym: F. gibbosa Blume var. rigida Miq. l.c.
(1867) 276.
Sumatra, Borneo, Java, Sumbawa, and Alor. Rare in Borneo, known only by 2
309
collections each from Ranau district in Sabah (SAN 119802, SAN 121454), and from
Belaga and Tatau (Ulu Mayeng) districts in Sarawak (S. 19217, S. 39824).
65. Ficus treubii King

(Melchior Treub, 18511910, former Director of the Bogor Botanic Gardens, Indonesia)
Ann. Roy. Bot. Gard. Calc. 1, 2 (1888) 105; Merrill l.c. (1921) 228; Corner l.c. (1965) 91; Anderson l.c.
261; Whitmore, Tantra & Sutisna l.c. 257; Coode et al. (eds.) l.c. 218. Type: Beccari PB 2800, Borneo,
Sarawak (holotype FI).
Dioecious tree, up to 15 m tall, 20 cm diameter. Bark grey-brown. Twigs solid, c. 0.3

cm thick, sparsely hairy, with stipular scars. Stipules narrowly lanceolate, up to 1 cm
long, hairy, caducous or subpersistent. Leaves distichous, membranous, drying greenish,
rough-hairy below; elliptic to narrowly obovate, 1323 49.5 cm, base symmetric,
cuneate, margin entire or crenate, slightly revolute, apex acuminate or caudate, acumen
up to 3 cm long; midrib sunken above; lateral veins 79 pairs, with shorter veins in
between, often with submarginal glands, raised below, faint above; intercostal venation
scalariform to reticulate, distinct below, almost invisible above; petiole 0.71.2 cm long,
hairy. Syconia borne on long racemose twigs arising from the stem or on stolons arising
from the base of the stem, glabrous or thinly hairy, ripening yellowish brown, obovoid,
c. 1.5 cm diameter, surface pustulate, apex with a tiny ostiole; peduncle up to 0.7 cm
long; basal bracts 3, in a collar, up to 2 mm long, lateral and apical bracts often present;
internal bristles few to abundant, pale yellow to pale brown. Perianth cup-shaped or
saccate. Achenes c. 0.8 mm long, slightly tuberculate.
Distribution. Endemic to Borneo. Common in Sabah, especially on Mt. Kinabalu, rare
in Sarawak. Also in Brunei and Kalimantan.
altitude.
66. Ficus uncinata (King) Becc.

(Latin, uncinatus = hooked; the incurved lateral bracts on the syconium)
For. Born. (1902) 527; Merrill l.c. (1921) 228; Corner l.c. (1965) 91, l.c. (1988) 546; Kochummen l.c.
159; Anderson l.c. 261; Whitmore, Tantra & Sutisna l.c. 258. Basionym: F. geocarpa Teijsm. ex Miq.
var. uncinata King l.c. (1888) 127. Type: Beccari PB 2458, Borneo, Sarawak (holotype FI). Synonym:
F. hypogea (non King) Merr. l.c. (1921) 224.
Dioecious tree, up to 8 m tall, 15 cm diameter. Twigs solid, ridged, 0.30.6 cm thick,

young ones covered with spreading or appressed hairs. Stipules lanceolate, acuminate,
34 cm long, hairy outside, caducous or persistent. Leaves distichous, drooping, coriaceous,
thin-coriaceous or membranous, glabrous or rough-hairy above, whitish or reddish brown
soft-hairy below; oblong, obovate, lanceolate or elliptic, 645 216 cm, base prominently
asymmetric, one side rounded or cordate or subcordate, and the other sharply cuneate
or obliquely truncate, very rarely symmetric, margin finely dentate or entire and plane,
310
apex acuminate, acumen 27 cm long; midrib raised or sunken above; lateral veins 6
13 pairs, curved, with many shorter veins in between, raised below, faint above, basal
veins 58 pairs on the broader side of leaf base, short; intercostal venation reticulate,
scalariform, or subscalariform, distinct below, faint above; petiole 0.52 cm long, hairy,
with or without subnodal gland. Syconia borne on long runners arising from the base
of stem, ripening pink to dark red-brown or crimson, depressed globose, subglobose, oblong,
or pear-shaped, 1.73.5 24 cm, glabrous or hairy, with numerous incurved, lanceolate
lateral bracts crowded round the sunken ostiole; internal bristles absent. Perianth absent.
Achenes c. 1.2 mm long, compressed, slightly keeled, with a ridge on each side.
Distribution. Sumatra, Peninsular Malaysia and Borneo. Widely distributed in Sabah and
Sarawak.
altitude, often by streams.
Key to varieties
1.
Leaves whitish hairy on both surfaces............

var. C
Hairs whitish, spreading. Twigs ridged. Stipules ovate to lanceolate, c. 1.5 cm long,
persistent. Leaves membranous, whitish hairy on both surfaces, drying greenish;
oblong, 2632 8.511 cm, base asymmetric, one side rounded, the other sharply
cuneate, margin distinctly toothed, apex acuminate, acumen up to 3 cm long; midrib
raised above; lateral veins 57 pairs, faintly raised on both surfaces; intercostal
venation scalariform to reticulate; petiole 0.51 cm long, hairy. Syconia subglobose,
c. 0.7 cm diameter (young), hairy.
Endemic to Borneo. Known only from Sarawak by 2 collections, S. 34268 from
Sabal Tapong FR in Samarahan district, and S. 57178 from G. Putri in Lundu
district.
Leaves reddish brown hairy..................2
2.
Syconia glabrous.................................................3
Syconia hairy.................4
3.
Indumentum consisting of appressed hairs. Lateral veins 47 pairs...

.....var. uncinata (in part)
Indumentum consisting of spreading hairs. Lateral veins 910 pairs...........
var. A
Leaves thin-coriaceous, lower surface reddish brown hairy with spreading hairs;
elliptic or oblong, 2225 810 cm, base asymmetric, margin faintly toothed,
apex acuminate, acumen c. 2 cm long; midrib flat above; lateral veins 910
pairs; intercostal venation scalariform; petiole c. 1 cm long, hairy. Syconia
glabrous, 1.52 cm diameter.
Known by a single collection, SAN 94520, from Beluran, Labuk Sugut district,
Sabah.
4.
Stipules persistent...............
311
var. subbeccarii Corner

(somewhat similar to F. beccarii)
Gard. Bull. Sing. 19 (1962) 401, l.c. 92; Whitmore, Tantra & Sutisna l.c. 258. Type: RSNB
1619, Borneo, Sabah, Mt. Kinabalu (holotype K).
Indumentum comprising appressed, reddish brown hairs up to 1 mm long. Stipules

persistent, 12 cm long. Leaves coriaceous, appressed hairy on both surfaces;
obovate, 1030 4.510.5 cm, base symmetric or slightly asymmetric, margin
entire, plane, apex acuminate, acumen c. 1.5 cm long; lateral veins 47 pairs;
intercostal venation raised below. Syconia 1.42 cm diameter, rough-hairy.
Endemic to Borneo. Rare in Sabah and Sarawak.
Stipules caducous or subpersistent...............5
5.
Indumentum comprising hairs 24 mm long. Stipules subpersistent............

var. pilosior Corner
(Latin, pilosiorus = with thicker/longer hairs; the leaves)
al. (eds.) l.c. 219. Type: BRUN 5302, Borneo, Brunei, Sg. Belalong (holotype CGE).
Stipules lanceolate, up to 6 cm long, hairy, subpersistent. Leaves coriaceous,

densely hairy below; elliptic to obovate, 2045 716 cm, base cuneate on one
side and cordate to auriculate on the other, margin faintly toothed, apex acuminate,
acumen c. 2.5 cm long; lateral veins 1013 pairs; petiole 0.71.5 cm long. Syconia
subglobose, 22.5 cm diameter, sparsely hairy.
Endemic to Borneo. Known by a single collection, S. 24304, from Lambir Hill,
Miri district, Sarawak. Also in Brunei and Kalimantan.
Indumentum comprising shorter hairs. Stipules caducous..........6
6.
Syconia densely hairy.............7

Syconia sparsely hairy...................8
7.
Leaves 2.56 cm broad; hairs 0.51.5 mm long, appressed...............

var. gracilis Corner
(Latin, gracilis = thin, slender; inference unclear)
al. (eds.) l.c. 219. Type: BRUN 5303, Borneo, Brunei (holotype CGE).
Indumentum as in var. uncinata. Stipules caducous. Leaves thin-coriaceous; 9

24 2.56 cm, base asymmetric, margin faintly toothed, apex acuminate, acumen
1.52.5 cm long; midrib raised above; lateral veins 811 pairs; intercostal venation
scalariform to reticulate; petiole 0.31 cm long, hairy. Syconia subglobose, 1.2
1.5 cm diameter, villous.
Endemic to Borneo. Not known in Sabah; in Sarawak recorded only by a few
collections. Also in Brunei and Kalimantan.
Leaves 415 cm broad; hairs 12.5 mm long, erect...........
var. strigosa Corner
(Latin, strigosus = covered with rigid, closely pressed, rather short, bristle-like
hairs; the syconia)
Gard. Bull. Sing. 18 (1960) 59, l.c. (1965) 92; Kochummen l.c. 160; Whitmore, Tantra &
Sutisna l.c. 258, Coode et al. (eds.) l.c. 219. Type: SFN 25946, Peninsular Malaysia, Johore
(holotype SING).
312
Hairs 12.5 mm long, reddish, erect. Stipules caducous. Leaves thin-coriaceous,

soft-hairy below, rough-hairy above; 1216 46.5 cm, base cordate on one
side, cuneate on the other, margin faintly toothed, apex acuminate, acumen 1.2
2.4 cm long; midrib raised above; lateral veins c. 4 pairs; intercostal venation
reticulate; petiole 0.50.8 cm long, hairy. Syconia oblong, c. 1.5 1 cm, densely
hairy.
Sumatra, Peninsular Malaysia and Borneo. Not known in Sabah; very rare in
Sarawak, known only by 1 collection, S. 41983, from Betong district. Also in
8.
Leaves strongly asymmetric at the base, one side truncate, the other subcordate; lateral
veins 911 pairs.............................................9
Leaves not so; lateral veins 47 pairs.................
var. uncinata (in part)
Hairs 0.51.5 mm long, reddish, appressed or spreading. Stipules caducous. Leaves
coriaceous, reddish brown soft-hairy below, rough hairy above; ovate to lanceolate,
1421.5 45.5 cm, base cuneate on one side, cordate on the other, margin
faintly toothed, apex acuminate, acumen 37 cm long; midrib flat above; lateral
veins 47 pairs; intercostal venation scalariform to reticulate; petiole 0.51 cm
long, hairy. Syconia subglobose, 1.52.5 cm diameter, glabrous or sparsely hairy.
Common and widely distributed in Sarawak, not yet reported for Sabah.
9.
Leaves 59 cm broad..............
var. truncata Corner
(Latin, truncatus = truncate; the leaf base)
al. (eds.) l.c. 219. Type: BRUN 5329, Borneo, Brunei, Sg. Belalong (holotype CGE).
Hairs 0.51 mm long, appressed, reddish. Stipules caducous. Leaves coriaceous,

appressed short-hairy on both surfaces; oblong, elliptic, or obovate, 1629
59 cm, base asymmetric, one side obliquely truncate to cuneate the other
subcordate or subauriculate, margin entire, plane, apex acuminate, acumen 1.2
2 cm long; midrib flat above; lateral veins 914 pairs; intercostal venation
scalariform to reticulate, raised below; petiole 0.61.1 cm long, appressed hairy.
Syconia sparsely hairy.
Endemic to Borneo. Not known in Sabah; very rare in Sarawak, known by a
single collection, S. 34012, from the Lanjak-Entimau Wildlife Sanctuary. Also
Leaves 714 cm broad.............
var. B
Hairs erect, brown, sparse. Stipules caducous, lanceolate, c. 2 cm long, hairy.
Leaves coriaceous, sparsely hairy on both surfaces; oblong or elliptic, 15.531
714 cm, base strongly asymmetric, rounded on one side, sharply cuneate on
the other, margin faintly toothed, apex acuminate, acumen 12 cm long; midrib
flat above; lateral veins 811 pairs, raised below, faint above; intercostal venation
scalariform, sunken above, distinct below; petiole 0.81.5 cm long, hairy. Syconia
sparsely hairy.
Endemic to Borneo. Known by 3 collections, S. 40005 and S. 66404 from Sarawak,
and Veldkamp 8379 from Bukit Raya, Kalimantan.
313
67. Ficus variegata Blume
Fig. 12.
(Latin, variegatus = blotched; the leaves)

Ridley, l.c. (1924) 335; Burkill l.c. 1032; Backer & Bakhuizen f. l.c. 28; Corner l.c. (1965) 82; Kochummen
l.c. 160; Anderson l.c. 262; Whitmore, Tantra & Sutisna l.c. 259; Coode et al. (eds.) l.c. 219. Type:
Blume s.n. (= RHL Sheet No. 9081882067, Java (L). Synonyms: F. subracemosa Blume l.c. 469; F.
cordifolia Blume l.c. 438; F. amboinensis Kostel., Allg. Med. Pharm. Pl. 2 (1831) 408; F. racemifera
Roxb., Fl. Ind., ed. Carey, 3 (1832) 560; F. laevigata Blanco, Fl. Filip. (1837) 682; F. cerifera Blume ex
Bleekrode, Ann. Sc. Nat. IV, 3 (1855) 333; F. subopaca Miq. l.c. (1859) 320; Urostigma javanicum
Miq. l.c. (1859) 334; F. variegata Blume var. pilosior Miq. l.c. (1867) 295; F. latsoni Elmer, Leafl.
Philip. Bot. 1 (1906) 204; F. polysyce Ridl., J. Roy. As. Soc. Str. Br. 82 (1920) 195.
Dioecious tree, up to 40 m tall, 70 cm diameter, without free hanging aerial roots; buttresses
up to 2 m high. Bark grey-brown, smooth. Twigs 0.20.4 cm thick, reddish brown. Stipules
ovate, up to 0.6 cm long, silky hairy outside, caducous. Leaves spirally arranged,
coriaceous, glabrous; narrowly oblong, or ovate to lanceolate, 920 4.512 cm, base
symmetric, cuneate to rounded or subcordate, margin entire (toothed in saplings), plane,
apex acuminate, acumen up to 1.5 cm long; midrib flat above; lateral veins 57 pairs,
with shorter veins in between, well-spaced, curved, not steeply ascending, raised on both
surfaces, basal pair conspicuous, reaching more than 1/2 the length of the blade, often
with a small gland in the axils; intercostal venation scalariform, distinct to faint on both
surfaces; petiole 38 cm long. Syconia borne in dense clusters on short twigs or tubercles
arising from stem and branches, ripening to rose-red, often streaked with white dots,
ellipsoid or pear-shaped, 22.5 cm diameter, apex depressed; peduncle 13 cm long;
basal bracts 3, persistent; internal bristles absent. Tepals 34, lanceolate or spathulate,
free or shortly united at the base, glabrous, red to brownish. Achene 11.5 mm long,
lenticular, slightly keeled, smooth.
Distribution. India, Burma, Indo-China, Andaman Islands, Malesia, Solomon Islands, and
Queensland, Australia. Common in Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. Open areas in lowland forest.
Key to varieties
Leaves c. 12 5.5 cm; intercostal veins few, lax. Syconia ellipsoid or pyriform.........
var. ilangoides (Elmer) Corner.
(meaning unclear)
Gard. Bull. Sing. 18 (1960) 33. Basionym: Ficus ilangoides Elmer, Leafl. Philip. Bot. 9 (1937)
3445. Type: unknown.
Borneo and the Philippines. In Borneo, very rare, known only from Kuala Penyu
and Semporna (Palabag Is.) districts in Sabah.
Leaves larger; intercostal veins numerous, regular. Syconia ellipsoid...........var. variegata
314
vi
68. Ficus virens Ait.

(Latin, virens = green; the drying leaves)
Hort. Kew (1789) 451; Corner l.c. 17 (1959) 376. Type: unknown.
Distribution. India, Sri Lanka, Andaman Islands, Burma, S China, Hainan, Indo-China,
Thailand, Malesia, Caroline Islands, Solomon Islands, and N Australia. Only var. glabella
occurs in Sabah and Sarawak.
var. glabella (Blume) Corner
(Latin, glabellus = somewhat glabrous; the leaves)
Gard. Bull. Sing. 17 (1960) 377, l.c. (1965) 9; Backer & Bakhuizen f. l.c. 34; Kochummen l.c. 161;
Anderson l.c. 262; Whitmore, Tantra & Sutisna l.c. 259. Basionym: F. glabella Blume l.c. 452. Type:
Blume s.n., Java (L?). Synonym: Urostigma caulocarpum Miq. l.c. (1848) 568; F. caulocarpa (Miq.)
Miq. l.c. (1867) 265, syn. nov.
Monoecious small tree, without or with a few to numerous tufted free hanging aerial
roots. Twigs pale reddish brown, 0.30.5 cm thick, irregularly grooved. Stipules ovate,
up to 1.5 cm long, caducous. Leaves spirally arranged, thin-coriaceous, glabrous; elliptic
or obovate, 5.514 2.56 cm, base symmetric, cuneate, margin entire, plane, apex
acute or acuminate, acumen obtuse, c. 1.2 cm long; midrib raised above, keeled below,
lateral veins 812 pairs, with a number of shorter veins in between, well-spaced, curved,
departing from the midrib at a broad angle, faint on both surfaces, curving and joining
near the leaf margin, basal pair conspicuous; intercostal venation scalariform, faintly
visible on both surfaces; petiole 1.53.5 cm long, channelled on the adaxial side. Syconia
axillary or borne on twigs behind the leaves, in pairs or in clusters, ripening pink purple
to black, globose, 0.40.7 cm or 13 cm diameter, apex umbonate, sessile or peduncled,
not covered by irritant hairs; basal bracts broad, persistent, lateral bracts absent; internal
bristles abundant, white. Tepals 24, ovate, spathulate or narrowly lanceolate, free or
shortly united at the base. Achenes ovoid, smooth.
Distribution. Sri Lanka, Burma, Thailand, Sumatra, Peninsular Malaysia, Java, Borneo,
the Philippines, Sulawesi, Maluku, and New Guinea. Rare in Sabah and Sarawak. Not
known in Brunei.
Ecology. In lowland and hill forests, also in rocky coastal habitats.
69. Ficus virescens Corner

(Latin, virescens = becoming green; the drying leaves)
l.c. 219. Type: RSNB 965, Borneo, Sabah, Mt. Kinabalu (holotype K).
Dioecious tree, up to 13 m tall. Young twigs, petioles, stipules, and veins on the leaf
undersurface appressed rough-hairy. Twigs solid, 0.20.4 cm thick, yellowish, striate.
Stipules lanceolate, up to 1.5 cm long, caducous. Leaves spirally arranged or opposite,
316
thin-coriaceous, sparsely hairy below, drying to pale greenish; elliptic to narrowly obovate,
1230 412 cm, base symmetric, cuneate, margin faintly lobed, apex acuminate to
caudate, acumen 1.23 cm long; midrib flat above, slightly sunken towards lower half,
sharply keeled below; lateral veins 57 pairs, with shorter veins in between, flat above,
raised below, basal pair short or extending up to 1/4 the length of the blade; intercostal
venation scalariform, raised below; petiole 17 cm long, with a subnodal gland. Syconia
borne in clusters on leafless branched twigs arising on the stem, from ground-level up
to about 3 m above the ground, ripening yellowish brown, brown short-hairy, subellipsoid,
11.5 0.71 cm, surface with distinct ridges, apex depressed with distinct ostiole; peduncle
0.52.2 cm long; basal bracts 3, in a collar, ovate, 2.54 mm long, glabrous; internal
bristles many, yellowish, minute. Perianth collar-like or saccate. Achenes c. 1 mm long,
tuberculate.
Distribution. Endemic to Borneo. In Sabah very rare, known by 2 collections only (SAN
116842 from Sandakan district and the type). Also in Brunei and Kalimantan.
altitude.
70. Ficus xylophylla Wall. ex Miq.

(Greek, xylon = wood; phullon = leaf; the thick cardboard-like leaves)
Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 286; King l.c. (1887) 29; Merrill l.c. (1921) 228; Ridley l.c. (1924)
332; Burkill l.c. 1033; Corner l.c. (1965) 16; Kochummen l.c. 162; Anderson l.c. 262; Whitmore, Tantra
& Sutisna l.c. 260; Coode et al. (eds.) l.c. 219. Type: unknown.
Monoecious small to medium-sized tree, or more frequently strangler. Bark pale pinkish
grey, smooth. Twigs stout, 11.2 cm thick, strongly ridged. Stipules ovate to lanceolate,
c. 3.5 cm long, hairy outside, caducous. Leaves spirally arranged, very thick-coriaceous,
cardboard-like, glabrous, drying yellowish brown above; usually obovate, 13.527 6.5
14 cm, base cuneate, margin entire, revolute, apex obtuse or rounded; midrib sunken
above; lateral vein 47 pairs, looping and joining near the leaf margin, prominently raised
below, raised above except basal pair which is sunken, basal pair extending up to 1/
2 the length of the blade; intercostal venation reticulate, lax, visible below, invisible above;
petiole 46 cm long, wrinkled when dry. Syconia axillary, sessile, solitary or in pairs,
ripening orange and then purplish red, oblong, 34.5 22.8 cm, apex flat, surface rugose
on drying; basal bracts persistent, 1014 mm long, ovate, triangular, acute, keeled, concealed
by the syconia; internal bristles absent. Tepals 24, lanceolate to subspathulate. Achenes
tuberculate.
Distribution. Laos, Thailand, Sumatra, Peninsular Malaysia, and Borneo. Uncommon in
Ecology. In lowland forest, including swamp forest.
Uses. The brightly coloured figs and the leaves are attractive enough to introduce this
plant in landscaping.
317
4. PARARTOCARPUS Baill.
(resembling Artocarpus)
minggi or pinggi (Iban), terap (Malay)
Rusea Go
Adansonia 11 (1875) 294; Beccari, For. Born. (1902) 632; Merrill, EB (1921) 218; Masamune, EPB
(1942) 251; Jarrett, J. Arn. Arb. 41 (1960) 320; Corner, Gard. Bull. Sing. 19 (1962) 241, ibid. 28 (1976)
183; Backer & Bakhuizen f., FJ 2 (1965) 17; Kochummen TFM 3 (1978) 162; Anderson, CLTS (1980)
262; Wong, DMT (1982) 8; Primack, Foresters Guide to the Moraceae of Sarawak (1983) 118; Whitmore,
Tantra & Sutisna, CLK 2, 1 (1990) 260; Rohwer in Kubitzki et al. (eds.), Fam. & Gen. Vasc. Pl. 2 (1993)
447; Coode et al. (eds.), CLBD (1996) 220; Argent et al. (eds.), MNDT-CK 2 (1997) 439, PROSEA 5,
3 (1998) 425. Synonym: Gymnartocarpus Boerl., Icon. Bog. 1 (1897) 73.
Medium-sized to large trees; buttresses sometimes present. Bark smooth or scaly, covered
with large pustular lenticels; inner bark mottled with cream and orange patches; latex
abundant, milky white. Sapwood pale yellow to brown. Twigs lenticellate or with prominent
leaf scars. Stipules non-amplexicaul, caducous, scars intrapetiolar. Leaves simple, spirally
arranged, thin- to thick-coriaceous, entire, glabrous or variously pubescent or scabrous.
Inflorescences unisexual, rarely bisexual, capitate; males and females indistinguishable
externally before anthesis, solitary or paired in leaf axils, peduncled; basal involucre,
if present, comprising 34(8) deltoid or ovate persistent bracts. Male, female, or bisexual
flowers sunken in numerous cavities over the entire surface of the receptacle; receptacle
armoured with closely set spine-like, conical, obtuse or truncate processes comprising
perianth segments and interfloral bracts; at anthesis, anthers and stigma exserted between
the processes. Male flowers without well-developed perianth, stamens 13 in each
receptacle cavity, filaments free or united, anthers extrorse, rarely introrse, shortly exserted,
pistillode absent. Female or bisexual flowers with solitary ovaries in each receptacle
cavity; ovary unilocular, style apical, with short exserted lanceolate or fimbriate stigmatic
arms; ovule subapical. Syncarp formed by the enlargement of the entire female inflorescence,
of various sizes, subglobose to globose, covered with closely set spine-like connate
processes. Fruits drupaceous, with firm exocarp. Seeds large, testa membranous with
a thickened apical cap; embryo curved, cotyledons unequal, radicle large, straight; germination
epigeal.
Distribution. Four species, distributed in Peninsular Thailand and Malesia (except Maluku
and Lesser Sunda Islands). All four species occur in Sabah and Sarawak.
Uses. Timber is traded as terap, a light hardwood. Seeds and latex are reported to be
poisonous and the latter is used in the preparation of arrow poison.
Notes. The inflorescence is somewhat similar to that of Artocarpus but differs in having
a usually well-developed involucre consisting of 34(8) basal bracts. In Artocarpus the
male and female inflorescences differ in shape, where as in Parartocarpus they are
indistinguishable before floral anthesis. In Parartocarpus the anthers are extrorse or rarely
318
introrse, while in Artocarpus they are consistently introrse. The bark of Parartocarpus
trees is always covered with large lenticels.
Key to Parartocarpus species

(for sterile specimens)
1.
Upper and lower leaf surfaces evenly scabrous; leaf margin conspicuously revolute......
........3. P. spinulosus
Upper and lower leaf surfaces glabrous or sparsely or densely velvety hairy; leaf
margin plane or only very slightly revolute.....2
2.
Leaves 1.64 cm wide; lateral veins forking to form irregular, rather jagged veinloops near the leaf margin.......2. P. microcarpus
Leaves 310(12) cm wide; lateral veins forming smooth vein-loops near the leaf
margin..........................3
3.
Lower leaf surface densely velvety hairy all over..1. P. bracteatus

Lower leaf surface glabrous or at most with scattered hairs on the veins....
............4. P. venenosus
Key to Parartocarpus species

(for fertile specimens)
1.
Inflorescences bisexual; lateral veins of leaf forking to form irregular, rather jagged
vein-loops near the leaf margin............2. P. microcarpus
Inflorescences unisexual; lateral veins of leaf forming smooth vein-loops near the
leaf margin.........2
2.
Syncarp up to 3 cm diameter. Leaf surface evenly scabrous......3. P. spinulosus

Syncarp 1018 cm diameter. Leaf surface glabrous or sparsely or densely velvety
hairy below..........3
3.
Peduncle densely short patent-hairy; basal bracts 68 mm long. Syncarp processes

spine-like, 35 23 mm. Lower leaf surface densely velvety hairy all
over.............1. P. bracteatus
Peduncle sparsely appressed short-hairy; basal bracts 35 mm long. Syncarp processes
truncate to flattened or spine-like, mostly 47 35 mm. Lower leaf surface glabrous
or at most with scattered hairs on the veins.....4. P. venenosus
1. Parartocarpus bracteatus (King ex Hook.) Becc.

(Latin, bracteatus = with bracts; the well-developed involucre of bracts at the base of the
inflorescence and syncarp)
For. Born. (1902) 632; Jarrett, J. Arn. Arb. 40 (1959) 12, l.c. (1960) 324; Kochummen l.c. 164; Anderson
l.c. 264; Primack l.c. 120; Whitmore, Tantra & Sutisna l.c. 260; Turner, Gard. Bull. Sing. 47 (1995) 356;
319
Argent et al. (eds.) l.c. 440. Basionym: Artocarpus bracteata King in Hooker f., Fl. Brit. Ind. 5 (1888)
540, Ridley l.c. (1924) 352. Lectotype (Jarrett, 1960): Griffith 4663, Peninsular Malaysia, Malacca
(holotype CAL; isotype K).
Tree up to 50 m tall, 80 cm diameter; bole straight, with many branches, without buttresses.
Bark whitish to brownish grey. Sapwood cream to pale white. Twigs brown, glabrous,
with prominent petiolar scars. Stipules broadly lanceolate, 0.51 cm long, rufous pubescent.
Leaves thick-coriaceous, glabrous above, densely velvety rufous hairy below; obovate
to oblong, (14)1620(24) (6)810(12) cm, base obtuse to cuneate, margin entire,
plane or slightly revolute, apex rounded to acute; midrib sunken above, raised below;
lateral veins 1012 pairs, sunken or flat above, prominent beneath, forming smooth veinloops near the leaf margin; intercostal venation scalariform, raised below; petiole 24
cm long, pubescent. Inflorescences unisexual; peduncle 24 cm long, densely patent shorthairy; basal bracts 3, 68 mm long. Male heads globose, 2.53.5 cm diameter, spinulous;
stamens 2 in each cavity, up to 8 mm long, filaments free or united at the base, anthers
oblong, apiculate, 2.53 mm long. Female heads spinulous as the males, most of the
processes basally connate in groups of 24; styles lanceolate, up to 3 mm long, exserted
in between the free apices of the processes. Syncarps subglobose, up to 10 cm diameter,
shallowly lobed; surface with fine spine-like processes 35 23 mm, those on lobes
bulbous at the base; peduncle 45 cm long, densely short patent-hairy; core c. 4 cm
across. Fruits numerous, ellipsoid, c. 1.5 1.2 cm.
Vernacular names. Sabahterap (Malay). Sarawakkaher (Kayan), kateh, minggi, pinggi
(Iban), selangking (Malay).
Distribution. Peninsular Malaysia, Sumatra and Borneo. Rare and uncommon in Sabah
and Sarawak. Also in Kalimantan.
Ecology. In mixed dipterocarp forest on hillsides, on clayey soils, up to 500 m altitude.
Flowering and fruiting from AprilOctober.
2. Parartocarpus microcarpus Corner

(Greek, mikros = small, karpos = fruit; with small syncarps)
Gard. Bull. Sing. 28 (1976) 184; Primack l.c. 121; Whitmore, Tantra & Sutisna l.c. 260. Type: S. 28676,
Borneo, Sarawak, Kapit district, Ulu Baleh, Bt. Tibang (holotype SAR; isotypes K, SAN).
Tree up to 30 m high, 45 cm diameter; buttresses up to 2 m tall. Bark whitish to grey;

inner bark yellowish pink. Twigs 0.30.5 cm thick, greyish, sparsely covered by stiff
whitish short hairs. Stipules 0.20.35 cm long, appressed hairy, caducous. Leaves thickcoriaceous, glabrous above, sparsely velvety hairy below; oblanceolate to elliptic, 310
1.64 cm, base attenuate, acute, margin entire, plane, apex rounded with mucronate
tip; midrib flat above, raised below; lateral veins 68 pairs, conspicuous and prominent
on both surfaces, forking to form irregular, rather jagged vein-loops near the leaf margin;
intercostal venation fine, somewhat reticulate, indistinct on both surfaces; petiole 1.5
3.5 cm long, sparsely covered with short white hairs. Inflorescences bisexual, globose,
c. 1.75 cm across; peduncle 13 cm long; basal bracts 4, short-hairy, persistent. Male
320
flowers numerous, in hairy receptacle cavities 45 34 mm; tepals 2, minute; stamens

2, rarely 1 or 3, anthers 34 1.5 mm, introrse, not exserted, filaments very short, free,
with conspicuous hairs around the base; pistillode absent. Female/bisexual flowers solitary
in sparsely hairy receptacle cavities, few; tepals 23(4); ovary glabrous, style terminal,
embedded in the receptacle wall, exserted between the tepals, stigma short, simple, hardly
projecting. Syncarps subglobose, 2.53 cm diameter, covered with flat, scarcely enlarged
truncate processes. Fruits 11.5 1 cm, dark brown; exocarp thin, endocarp woody,
c. 0.5 mm thick; seed-coat membranous.
Vernacular names. Sarawakkateh, kati (Iban), minggi (Malay).
Distribution. Endemic to Borneo. Recorded only for Sabah and Sarawak, uncommon.
In Sabah known from Tambunan district, and in Sarawak recorded for Miri and Limbang
districts.
Ecology. In lower montane forest, at 13001800 m altitude. Flowering and fruiting from
MarchOctober.
3. Parartocarpus spinulosus R. Go
Fig. 13.
(Latin, spinulosus = minutely spiny; the rough-hairy leaf surface)

Sandakania 12 (1998) 1. Type: Soepadmo, Rena & Kalkausar FRI 41312, Borneo, Sabah, Kinabatangan
district, Telupid, Bt. Tawai FR (holotype KEP; isotypes K, L, SAN, SAR).
Tree up to 10 m, 15 cm diameter. Bark scaly to shallowly fissured, brown; inner bark

yellow. Sapwood pale yellow. Twigs with prominent petiolar scars, sparsely appressed
hairy when young, glabrescent. Stipules c. 0.15 0.1 cm, connate, caducous. Leaves
thick-coriaceous, evenly scabrous; obovate, 814(16) (4)57 cm, base cuneate with
small auriculate extension, margin entire, conspicuously revolute, apex rounded to strongly
emarginate; midrib and lateral veins flattish above, raised beneath; lateral veins 810
pairs, forming smooth vein-loops near the leaf margin, hairy on the veins below; petiole
23 cm long, strongly wrinkled, sparsely stiff-hairy when young, glabrescent. Inflorescences
unisexual; males globose, 13 cm diameter; peduncle 24 cm long, glabrous; basal bracts
4, persistent. Male flowers: stamens two per receptacle cavity, anthers lanceolate, 0.5
1 mm long, exserted at anthesis. Female flowers not seen. Syncarps globose, up to
3 cm diameter (immature), with obtuse or truncate processes. Fruits numerous, embedded
in the club-shaped receptacle.
Distribution. Endemic to Borneo. Rare, only known by 2 collections, the type and Leopold
SAN 43885 from Mile 87, Telupid Road, Sandakan district, Sabah.
Ecology. In lowland mixed dipterocarp forest, up to 150 m altitude, on ultrabasic soils,
along stony river banks.
Notes. Close to Parartocarpus venenosus subsp. borneensis but differs by its strongly
scabrous leaf surface.
321
vi
4. Parartocarpus venenosus (Zoll. & Moritzi) Becc.
Fig. 14.
(Latin, venenosus = very poisonous; the latex)

For. Born. (1902) 632; Jarrett l.c. (1960) 326; Kochummen l.c. 165; Anderson l.c. 262; Primack l.c. 121;
Whitmore, Tantra & Sutisna l.c. 260; Turner l.c. 356; Coode et al. (eds.) l.c. 220; Argent et al. (eds.) l.c.
440. Basionym: Artocarpus venenosa Zoll. & Moritzi, Nat. Gen. Arch. Neerl. Ind. 2 (1845) 213. Type:
Beccari PB 2557, Borneo, Sarawak (K, P). Synonyms: A. tylophylla Miq., Fl. Ind. Bat. 1 (1859) 289; A.
cerifera Miq., Ann. Mus. Lugd.-Bat. 3 (1867) 212; A. riedelii Miq. l.c. (1867) 213; P. beccarianus
Baill., Adansonia 11 (1875) 294; Gymnartocarpus venenosus (Zoll. & Moritzi) Boerl., Handl. Fl. Ned.
Ind. 3 (1900) 73; A. woodii Merr., Philip. J. Sc. 3 (1908) Bot. 221.
Tree up to 35 m tall, 50 cm diameter; buttresses up to 1.5 m high, or absent. Bark

yellowish to greyish brown. Twigs greyish, rugose, appressed pubescent. Leaves thinto thick-coriaceous, glabrous or sparsely hairy on the veins beneath, drying grey-green
to purplish brown above, yellow-green to red-brown below; obovate, oblong, oblanceolate
or elliptic, 423(30) 310 cm, base rounded or cuneate, sometimes auriculate, margin
entire, plane to slightly revolute, apex rounded, obtuse to short-acuminate; midrib prominent,
flattish or sunken above, raised beneath; lateral veins 615 pairs, forming smooth veinloops near the leaf margin; intercostal venation reticulate, obscure above; petiole 1.5
5 cm long. Inflorescences unisexual; males globose, 1.53 cm diameter, covered with
closely set spine-like, conical, obtuse, umbonate or truncate processes; females similar
to the males but the processes mostly connate at the base in groups of 24; peduncle
1.26 cm long, sparsely appressed short-hairy; bracts 34 (8), deltoid, short-pubescent
to glabrous, 35 mm long. Male flowers: stamens 13 per receptacle cavity, filaments
up to 4 mm long, united, anthers oblong, obtuse, 22.5 mm long. Female flowers: style
lanceolate, fimbriate, exserted. Syncarps up to 18 cm diameter, subglobose, covered with
truncate to flat or spine-like processes 47 35 mm; peduncle 310 cm long, sparsely
appressed short-hairy. Fruits ellipsoid, with thick stony endocarp.
Vernacular names. Sabahbangkil, tanggajun (Malay). Sarawakbangkil (Kenyah),
buah klidang (Murut), kateh, kati, minggi, pinggi (Iban).
Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, Java, Borneo, the
Philippines, Sulawesi, Aru Is., Papua New Guinea, Bismarck Archipelago, and Solomon
Islands. P. venenosus subsp. borneensis is very common in Sabah and Sarawak, while
the other 2 subspecies are uncommon. Also in Brunei and Kalimantan.
Ecology. In lowland to lower montane forests, including peatswamp forest, at 3001500
m altitude, on yellow clayey soils. Flowering and fruiting from FebruaryNovember.
Uses. The latex is used for preparing arrow poison. Ripe fruit is edible but the unripe
seeds are poisonous.
Key to subspecies
1.
Leaves thin-coriaceous; intercostal veins visible on both surfaces. Syncarp with truncate
or obtuse depressed processes.........................
323
vi
subsp. venenosus
In Sabah very rare, known only by 2 collections, SAN 37723 and SAN 57831.
Leaves thick-coriaceous; intercostal veins invisible on both surfaces. Syncarp with
spine-like processes..............................2
2.
Leaf base cuneate; midrib not sunken above. Syncarp peduncle 34 cm long. Confined
to peatswamp forest.................
subsp. forbesii (King ex Hook. f.) Jarrett
(H.O. Forbes, 18511932, Scottish botanist)
J. Arn. Arb. 41 (1960) 137; Kochummen l.c.165; Primack l.c. 122; Whitmore, Tantra & Sutisna
l.c. 260; Turner l.c. 356; Coode et al. (eds.) l.c. 220. Basionym: Artocarpus forbesii King in
Hooker f., Fl. Brit. Ind. 5 (1888) 539; Ridley l.c. (1924) 352. Type: King 10829, Peninsular
Malaysia (CAL, K, P). Synonym: P. excelsa Becc. l.c. 634; Gymnartocarpus triandra J.J.
Smith, Bull. Jard. Bot. Buitenz. 3, 4 (1922) 233; P. triandra (J.J. Smith) J.J. Smith, Bull. Jard.
Bot. Buitenz. 3, 6 (1924) 80, Browne, FTS (1955) 357.
Leaves thick-coriaceous; obovate, elliptic, oblanceolate to oblong, 813 46

cm, base cuneate; midrib prominent above and raised beneath; lateral veins 6
10 pairs, appressed short-hairy below, glabrescent. Male inflorescences globose;
peduncle 1.23.5 cm long. Syncarps with spine-like or umbonate processes;
peduncle 34 cm long.
Sumatra, Malay Peninsular and Borneo. In Sarawak known only from peatswamp
forest. Also in Brunei and Kalimantan.
Leaf base rounded to cuneate; midrib sunken above. Syncarp peduncle 48 cm long.
Not found in peatswamp forest.................
subsp. borneensis (Becc.) Jarrett
(of Borneo)
J. Arn. Arb. 41 (1960) 332; Anderson l.c. 262; Primack l.c. 123; Whitmore, Tantra & Sutisna
l.c. 260; Coode et al. (eds.) l.c. 220. Basionym: P. borneensis Becc. l.c. 634. Type: Beccari PB
2005, Borneo, Sarawak (FI, K).
Leaves thick-coriaceous, glabrous and dark brown above, sparsely hairy with
rusty minute stiff hairs below; obovate to elliptic, (6)810(12) 57 cm, base
rounded to cuneate, margin entire, slightly revolute, apex rounded, notched or
emarginate to acute; midrib sunken above, raised beneath; lateral veins 811
pairs, prominent below; petiole 47 cm long, sparsely covered by stiff short hairs.
Male inflorescences globose, c. 2 cm diameter; peduncle 35 cm long, sparsely
short-hairy. Female inflorescences globose, about the same size as the males,
sparsely short-hairy. Syncarps up to 8 cm diameter, spinulous, yellowish green
when mature; peduncle 48 cm long; bracts 4, persistent.
Endemic to Borneo. Very common in Sabah and Sarawak. Also in Brunei and
Kalimantan. In lowland and hill mixed dipterocarp forests, on ridges and hill
slopes, on yellow clayey soils, at 3001000 m altitude. Flowering and fruiting
from FebruaryNovember. Fruits edible.
325
5. PRAINEA King
(David Prain, 18571944, former curator of the Herbarium,
Royal Botanic Gardens, Culcutta, India)
selangking (Iban, Malay)
Rusea Go
In Hooker f., Fl. Brit. Ind. 5 (1888) 546; Beccari, For. Born. (1902) 635; Ridley, FMP 3 (1924) 358;
Jarrett, J. Arn. Arb. 40 (1959) 30; Burgess, TBS (1966) 399; Kochummen, TFM 3 (1978) 165;
Anderson, CLTS (1980) 262; Primack, Foresters Guide to the Moraceae of Sarawak (1983) 124;
Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 260; Rohwer in Kubitzki et al. (eds.), Fam. & Gen.
Vasc. Pl. 2 (1993) 448; Coode et al. (eds.), CLBD (1996) 220; PROSEA 5, 3 (1998) 469.
Trees or woody climbers (not in Sabah and Sarawak), unarmed, monoecious or dioecious.
Bark reddish brown; slash with milky white latex. Stipules paired, lateral or intrapetiolar,
non-amplexicaul. Leaves simple, spirally arranged or distichous, penninerved; lateral veins
distinctly looping near the leaf margin. Inflorescences unisexual, capitate, peduncled,
solitary or paired in the leaf axils; flowers among numerous peltate, clavate, or spathulate
interfloral bracts; perianth free, enclosing a single stamen or ovary; involucral bracts
absent. Male inflorescences with numerous flowers; male flowers: perianth tubular, bilobed
or perforated above, stamens shortly exserted. Female inflorescences with fewer flowers;
female flowers: perianth tubular, clavate, fleshy above, the mouth contracted, very shortly
4-lobed or merely perforated, ovary unilocular, style apical, bifid, the arms exserted, ovules
subapical. Syncarps subglobose to ellipsoid; perianth strongly enlarged, projecting from
the syncarp surface, fleshy, globose to ellipsoid. Fruits drupaceous, large, attached laterally
near the base of the syncarp, interfloral bracts persistent. Seeds without endosperm; embryo
straight, cotyledons equal fleshy; radicle and plumule minute, apical.
Distribution. Four species, distributed in Sumatra, Peninsular Malaysia, Borneo, Maluku,
and Papua New Guinea. In Sabah and Sarawak two species are found, of which one
is endemic to Borneo (Prainea frutescens).
Ecology. Mostly found in lowland and hill forests, below 1000 m altitude.
Uses. Source of medium hardwood for local use.
Key to Prainea species

Leaves 516 37 cm, thin-coriaceous to chartaceous. Syncarp with a peduncle 1.3
1.6 cm long............1. P. frutescens
Leaves 1035 415 cm, thick-coriaceous. Syncarp with a peduncle 7.511.5 cm
long........2. P. limpato
326
1. Prainea frutescens Becc.

(Latin, frutescens = shrubby; the habit)
For. Born. (1902) 635; Corner l.c. (1962) 242; Anderson l.c. 262; Primack l.c. 125; Whitmore, Tantra &
Sutisna l.c. 261. Type: Beccari PB 667, Borneo, Sarawak (holotype FI; isotypes K, P). Synonym:
Artocarpus frutescens (Becc.) Renner, Engl. Bot. Jahrb. 39 (1907) 367.
Tree up to 30 m tall, 40 cm diameter; with short, rounded buttresses or without buttresses.

Bark grey, becoming brownish or reddish near the buttresses, smooth; inner bark reddish.
Sapwood yellowish. Young twigs sparsely hairy or puberulent, older ones glabrescent
to glabrous. Leaves thin-coriaceous to chartaceous, glabrous on both surfaces; obovate,
elliptic to oblong, 516 37 cm, base cuneate, rarely asymmetric, margin entire, plane,
apex abruptly contracted into narrow acumen c. 2 cm long; midrib prominent below;
lateral veins obscure, 610 pairs, looping near the leaf margin; intercostal venation
reticulate, obscure on both surfaces; petiole slender, up to 1.2 cm long. Inflorescences
solitary. Male inflorescences subglobose, 0.40.6 cm diameter; peduncle 0.51.2 cm long.
Male flowers: perianth 0.81 mm long, minutely pubescent; stamens slender, c. 1.2 mm
long, filament cylindrical, anthers oblong, up to 0.2 mm long; bracts with minute stalk.
Female inflorescences globose, up to 1 cm diameter, each bearing at least 2030 flowers
and numerous bracts. Syncarps globose, up to 1.5 cm diameter, compactly covered by
2030 developing female flowers and numerous bracts; fruiting perianth ovoid, c. 6 mm
diameter, projecting up to 7 mm long, apices obtuse, mouth shortly bilobed; bracts connate
in groups; peduncle 1.31.6 cm long. Fruits ripening red.
Vernacular names. Sarawakselangking (Malay), selangking batu (Iban).
Distribution. Endemic to Borneo. In Sabah uncommon; in Sarawak only known from
the west and northern parts. Not known in Brunei.
Ecology. In primary lowland mixed dipterocarp forest, up to 300 m altitude, including
heath forest (kerangas) on yellow sandy or brownish soils. Flowering either in February
March or from OctoberNovember; fruiting in JuneAugust or NovemberDecember,
February and April.
2. Prainea limpato (Miq.) Beumee ex Heyne
Fig. 15.
(Sumatran plant namelimpatu)

Nutt. Pl. Ned. Ind., ed. 2, 1 (1927) 579; Jarrett l.c. (1959) 34; Burgess l.c. 399; Kochummen l.c. 673;
Anderson l.c. 262; Primack l.c.126; Whitmore, Tantra, Sutisna l.c. 261; Coode et al. (eds.) l.c. 220.
Basionym: Artocarpus limpato Miq. Fl. Ind. Bat., Suppl. (1861) 421. Type: Diepenhorst HB 2500,
Sumatra, Priaman (holotype U; isotype L), Synonyms: Urostigma diepenhorstii Miq. l.c. (1861) 439;
Ficus diepenhorstii (Miq.) King, Ann. Roy. Bot. Gard. Calc. 1, 2 (1888) 181; P. cuspidata Becc. l.c.
636; P. multinervia Merr., Philip. Journ. Sci 29 (1926) 364.
Deciduous tree, up to 30 m tall, 40 cm diameter; with or without buttresses. Bark greyish

to reddish brown, smooth to somewhat scaly and sometimes flaky; inner bark greyish
white or pale grey. Sapwood pale greenish. Twigs rounded or flat, 0.20.7 cm thick,
appressed puberulent. Leaves thick-coriaceous; oblong to obovate or elliptic, 1035
327
vi
415 cm, base asymmetric, cuneate to rounded, margin revolute, apex abruptly contracted
into narrow acumen up to 2 cm long; lateral veins 1214 pairs, prominently raised beneath,
looping near the leaf margin; intercostal venation reticulate, faint to distinct above; petiole
up to 3 cm long. Inflorescences solitary or paired. Male inflorescences globose, up to
1.5 cm diameter; peduncle 34 cm long, glabrous or short-hairy. Male flowers: perianth
tubular, perforated, minutely pubescent; stamens c. 1.8 mm long, filament cylindrical,
anthers oblong, c. 0.5 mm long; bracts with slender stalk, ciliate. Female inflorescences
globose; interfloral bracts abundant, spathulate or with peltate upper parts, densely or
sparsely hairy. Syncarps up to 5.5 cm diameter, very irregularly shaped, loosely covered
by numerous bracts, yellow-green, drying brown; fruiting perianth 820 per head, projecting
to 1520 mm, ellipsoid, c. 10 mm diameter, obtuse or broadly attenuate; peduncle 7.5
11.5 cm long, puberulent.
Vernacular names. Sabahkasusur aboi, kesusu (Dusun), kayu susu (Malay, Sandakan),
tampang (Malay), usu (Dusun, Banggi). Sarawakempatak (Iban), karen or karon (Bidayuh),
tampang (Malay).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah scattered, uncommon;
in Sarawak mostly found in Kuching district, uncommon. Also in Brunei and Kalimantan.
Ecology. In lowland and hill mixed dipterocarp forests on loamy, sandstone-derived or
sandy soils, on limestone ridges and riverbanks, up to 700 m altitude. Flowering May
August; fruiting in AugustJanuary.
Use. Source of light hardwood known as beruni.
6. STREBLUS Lour.
(Greek, streblos = crooked or distorted; referring to the twigs)
gili-gili (Bassap-Malbuh, Sabah); tempinis (Malay)
Rusea Go
Fl. Cochinch. (1790) 615; Ridley, FMP 3 (1924) 323; Corner, Gard. Bull. Sing. 19 (1962) 215, WSTM
3rd. edition 2 (1988) 555; Kochummen, TFM 3 (1978) 167; Anderson, CLTS (1980) 262; Primack,
Foresters Guide to the Moraceae of Sarawak (1983) 125; Berg, Proc. Kon. Ned. Akad. Wetensch. C
91 (1988) 356; Rohwer in Kubitzki et al. (eds.), Fam. & Gen. Vasc. Pl. 2 (1993) 446; Coode et al.
(eds.), CLBD (1996) 220; PROSEA 5, 3 (1998) 537. Synonyms: Bleekrodea Blume, Mus. Bot. Lugd.Bat. 2 (1856) 87; Paratrophis Blume l.c. (1856) 81; Taxotrophis Blume l.c. (1856) 77; Sloetia Teijsm.
& Binnen. ex Kurz, J. Lin. Soc. Bot. 8 (1864) 168; Pseudostreblus Bureau in A. DC., Prodr. 17 (1873)
219; Pseudotrophis Warb., Bot. Jahrb. 13 (1891) 294.
Monoecious or dioecious, small to medium-sized trees or climbers (not in Sabah and

Sarawak), unarmed or with spines. Bark of various shades from pale to black, smooth,
sometimes lenticellate or finely flaky; inner bark creamy white, yellowish to greenish;
latex white to yellowish. Leaves distichous, simple, mostly oblong, elliptic to lanceolate.
329
Inflorescences axillary, bisexual or unisexual, cymose, racemose, spicate or capitate.

Female flowers solitary or paired, 4-merous; tepals free or joined; ovary superior. Male
flower with pistillode, 35-merous, mostly 4-merous; tepals free or joined, stamen filaments
inflexed in bud, anthers small, kidney-shaped, extrorse. Fruits simple, drupaceous, sometimes
with fleshy base, with persistent tepals which are enlarged or not.
Distribution. About twenty species in Asia, Australia, Madagascar, and Africa; with 15
species in Malesia, and 4 species in Sabah and Sarawak.
Ecology. Mostly found in lowland and hill forests, up to 900 m altitude.
Key to Streblus species

1.
Twigs and stem spiny.....3. S. ilicifolius

Twigs and stem not spiny........2
2.
Leaf margin serrate to crenate; lateral veins 56 pairs....4. S. macrophyllus

Leaf margin entire or undulate; lateral veins 618 pairs..3
3.
Lateral veins 1315 pairs. Inflorescences bisexual spikes or catkins, 56 cm long.

Fruits with fleshy base, covered by enlarged tepals.........1. S. elongatus
Lateral veins 918 pairs. Inflorescences unisexual; males in cluster of 210, females
either solitary or paired in leaf axils. Fruits without fleshy base, tepals not
enlarged.....2. S. glaber
1. Streblus elongatus (Miq.) Corner

(Latin, elongatus = elongate; the elongated inflorescences)
Gard. Bull. Sing. 19 (1962) 227, l.c. (1988) 556; Kochummen l.c. 168; Primack l.c. 128; Turner l.c. 356.
Basionym: Artocarpus elongatus Miq., Fl. Ind. Bat., Suppl. (1861) 419. Type: Teijsmann s.n., N Sumatra,
Lumut, W Siboga (L?, U?). Synonyms: Sloetia sideroxylon Teijsm. & Binnen. ex Kurz l.c. 168; S.
pinangeana D. Oliver in Hooker, Ic. Pl. (1886) t.1531; S. wallichii King in Hooker f., Fl. Br. Ind. 5
(1888) 493; S. elongata (Miq.) Koord., Exk. Fl. Java 2 (1912) 90.
Monoecious tree, up to 35 m tall, 50 cm diameter, unarmed. Bark rough, grey-brown;

inner bark soft, reddish; latex white. Sapwood yellowish white. Twigs glabrous, 0.15
0.25 cm thick, greyish brown, lenticellate. Terminal buds conical, c. 0.1 0.15 cm. Stipules
paired, 0.51.5 cm long, caducous, scars incompletely ring-like. Leaves thin-coriaceous,
glabrous on both surfaces; elliptic or obovate, (7)1020(26) (3)58(10) cm, base
asymmetric, cuneate, margin entire or undulate, revolute, apex acuminate, acumen up
to 1.5 cm long; midrib prominently raised above; lateral veins prominent on both surfaces,
1315 pairs; intercostal venation reticulate, prominent below; petiole 0.51.5 cm long,
glabrous. Inflorescences bisexual or male, axillary pendent spikes, 56 cm long, solitary
or in clusters of 24; male flowers numerous; female flowers 15 per inflorescence. Male
flowers sessile; tepals 3(4); stamens 3(4), 22.5 mm long, white; pistillode conical.
Female flowers mostly at the base of the spike; tepals 4; style up to 5 mm long, stigmatic
330
arms 1220 mm long. Fruits round, 0.81.2 cm diameter, with fleshy base, enclosed
by the enlarged tepals.
Vernacular name. Sarawaktempinis (Malay).
Distribution. Sumatra, Peninsular Malaysia, Singapore, Borneo, and Sulawesi. Uncommon
in Sabah and Sarawak. Not known in Brunei.
Ecology. In lowland and hill forests on yellow loam soils, up to 800 m altitude.
Use. The standard Malaysian name for the timber is tempinis. It is very hard and very
heavy, suitable for all forms of heavy construction.
2. Streblus glaber (Merr.) Corner
Fig. 16.
(Latin, glaber = without hairs; the plant parts)

Gard. Bull. Sing. 19 (1962) 221; Kochummen l.c. 168; Anderson l.c. 263; Primack l.c. 129; Turner l.c.
356; Coode et al. (eds.) l.c. 220. Basionym: Gironneira glabra Merr., Philip. J. Sci. 1 (1906) Suppl. 42.
Type: Whitford 1205, the Philippines, Luzon, Mt. Mariveles, Lamao R. (holotype K). Synonyms:
Aphananthe negrosensis Elmer, Leafl. Phil. Bot. 2 (1909) 575; Pseudostreblus caudatus Ridl., J. Fed.
Mal. Str. Mus. 6 (1915) 54; Paratrophis glabra (Merr.) Steenis, J. Bot. 72 (1934) 8; Streblus laevifolius
Diels, Bot. Jahrb. 67 (1935) 171; Chevalierodendron glabrum (Merr.) Leroy, C.R. Acad. Sc. Paris 227
(1948) 146.
Dioecious tree, up to 40 m tall, 60 cm diameter, unarmed. Bark smooth, dark grey to

brown; latex white. Twigs up to 0.2 cm thick, often somewhat flattened, greyish brown
to blackish, smooth. Stipules paired, 0.20.5 cm long, caducous. Leaves thin-coriaceous,
glabrous; lanceolate, (5)710(12) 23 cm, base cuneate, margin entire, plane, apex
sharply acuminate, acumen up to 2 cm long; midrib prominent on both surfaces; lateral
veins 918 pairs, prominent on both surfaces; intercostal venation reticulate, often slightly
raised beneath; petiole less than 1 cm long. Inflorescences unisexual; bracts ovate,
lanceolate or reniform. Male inflorescences unbranched, 330-flowered, clustered in
groups of 210, 12.5 cm long; peduncle up to 1 cm long, red when fresh. Male flowers
sessile, 4-merous; tepals reddish, stamens whitish, inflexed, pistillode columnar, 4-angled.
Female inflorescences solitary or paired in the leaf axils, 19-flowered; peduncle 1
2 cm long. Female flowers: tepals ovate, obtuse, 13 mm across; style c. 1 mm long,
stigmatic arms 1.54 mm long. Fruits pyriform, up to 1 cm diameter, without fleshy
base, glabrous, ripening dark purple, turning to black on drying; tepals persistent, not
enlarged.
Vernacular name. Sarawakselangking (Iban).
Distribution. Peninsular Malaysia, Borneo, the Philippines, Sulawesi, New Guinea, and
Solomon Islands. In Sabah and Sarawak uncommon. Also in Brunei and Kalimantan.
Ecology. In hill to upper montane forests, mostly on ridges with brown to blackish soils,
up to 2500 m altitude. Flowering from JanuaryMay; fruiting from MarchAugust.
331
vi
3. Streblus ilicifolius (Vidal) Corner

(having leaves resembling those of the holly treeIlex)
Gard. Bull. Sing. 19 (1962) 227, l.c. (1988) 557; Kochummen l.c. 167; Turner l.c. 356. Basionym:
Taxotrophis ilicifolia Vidal, Rev. Pl. Vasc. Filip. (1886) 249; Ridley l.c. (1924) 322. Lectotype (Chew,
ined. 1959): Vidal 1795, the Philippines, Isle of Marinduque (holotype P?). Synonyms: Pseudotropis
laxifolia Warb., Bot. Jahrb. 13 (1891) 294; Taxotropis obtusa Elmer, Leafl. Philip. Bot. 5 (1913) 1813;
T. triapiculata Gamble, Kew Bull. (1913) 188.
Dioecious tree, up to 10 m tall, 15 cm diameter, spiny. Bark greyish black, smooth and
lenticellate; inner bark yellowish with white to yellowish latex. Twigs 0.10.3 cm thick,
grey-brown. Stipules triangular, 0.30.5 cm long, caducous, on falling leaving complete
ring-like scars. Leaves thick-coriaceous, glabrous on both surfaces; elliptic, oblong, or
obovate, (4)612(15) (2)48(10) cm, base rounded, margin with sharp, spine-like
serration or undulate, apex acuminate; midrib raised on both sides; lateral veins 812
pairs, joining near the leaf margin to form looped intramarginal veins, flattish to slightly
sunken above, raised below; intercostal venation reticulate, faint on both surfaces; petiole
0.21 cm long. Inflorescences axillary, unisexual. Male inflorescences spicate, unbranched,
15 cm long, with many flowers; bracts reniform, 23 mm wide. Male flowers 4-merous,
tepals up to 1.5 mm long, pistillode 4-angled. Female inflorescences racemose with 3
6 flowers, 14 cm long; peduncle 0.31.5 cm long; bracts reniform. Female flowers
pedicelled; tepals 24 mm long; style up to 1 mm long, stigmatic arms 23.5 mm long.
Fruits ovoid to cylindrical, 0.60.8 cm diameter, with acute tip, seated on a very thick
fleshy base, more or less covered by the enlarged tepals.
Vernacular name. Sabahnunuk daun berduri (Malay).
Distribution. Chittagong, Burma, Indo-China, Thailand, Hainan, Peninsular Malaysia,
Borneo, the Philippines, Sulawesi, Maluku, Key Is., and Timor. In Sabah and Sarawak
uncommon and scattered. So far no record for Brunei and Kalimantan.
Ecology. In primary and secondary forests on sandy, black, or rocky ultrabasic soils,
frequently on hillsides or rocky ridges. Flowering in JuneOctober; fruiting from November
February.
4. Streblus macrophyllus Blume

(Greek, makro = large, phullon = leaf; large-leaved)
Mus. Bot. Lugd.-Bat. 2 (1856) 80; Corner l.c. (1962) 227; Kochummen l.c. 168; Turner l.c. 357. Type:
Teijsmann s.n. (= RHL Sheet No. 908189706), Sulawesi, near Likupang (holotype L). Synonyms:
Taxotrophis macrophylla (Blume) Boerl., Handl. 3 (1900) 359; Pseudotrophis mindanaensis Warb. in
Perkin, Fragm. Fl. Filip. 1 (1904) 165; Paratrophis caudata Merr., Philip. J. Sci. 1 (1906) Suppl. 183;
Taxotrophis balansae Hutch., Kew Bull. (1918) 151.
Dioecious or sometimes monoecious tree, up to 10 m tall, 15 cm diameter, unarmed.

Bark finely flaky, whitish brown or yellowish to greenish. Twigs 0.10.2 cm thick,
yellowish grey. Stipules 0.21 cm long, subpersistent or caducous. Leaves thin-coriaceous,
glabrous on both surfaces; ovate to oblanceolate, (4)712(15) (1.5)34(6) cm, base
333
cuneate to rounded, margin serrate to crenate, apex cuspidate, acumen up to 2.5 cm

long; midrib flat above, distinct and raised below; lateral veins 56 pairs, slightly raised
beneath; intercostal venation reticulate, obscure on both surfaces; petiole up to 0.3 cm
long. Inflorescences axillary, solitary, unisexual. Male inflorescences spicate, 414 cm
long, puberulous; peduncle 0.20.4 cm long; bracts subreniform to semicircular, 0.30.5
mm long. Male flowers sessile; tepals c. 1 mm long; pistillode 4-angled. Female
inflorescences racemose, 23 cm long, 210-flowered, glabrous; peduncle 0.21 cm long.
Female flowers with pedicel up to 1 cm long; style 23 mm long, stigmatic arms 1.5
5 mm long. Fruits ripening to reddish yellow, up to 1.5 cm long, thinly fleshy at the
base.
Vernacular names. Sabahkamuning (Malay, Sandakan), lampasih (Malay, Keningau).
Distribution. Indo-China, Peninsular Malaysia, Borneo, Sulawesi, and the Philippines. In
Borneo, so far only known from Sandakan and Keningau districts, Sabah.
Ecology. In lowland forest near riverbanks, on hill tops and slopes on brown soils. Flowering
and fruiting all year round.
334
MYRISTICACEAE (DE WILDE)
MYRISTICACEAE
W.J.J.O. de Wilde
Rijksherbarium/Hortus Botanicus,
Leiden, the Netherlands
Gamble, J. As. Soc. Beng. 75, 2 (1912) 205; Merrill, EB (1921) 268; Ridley, FMP 3 (1924) 53; Masamune,
EPB (1942) 300; J. Sinclair, Gard. Bull. Sing. 16 (1958) 205; Backer & Bakhuizen f., FJ 1 (1964) 137;
Burgess, TBS (1966) 407; Whitmore, TFM 1 (1972) 315; Anderson, CLTS (1980) 263; Whitmore,
Tantra & Sutisna, CLK 2, 1 (1990) 262; Turner, Gard. Bull. Sing. 47, 2 (1995) 357; Coode et al. (eds.),
CLBD (1996) 220; Argent et al. (eds.), MNDT-CK 2 (1997) 445.
Dioecious, or monoecious (Endocomia), or rarely paroecious (in a few species of Myristica),

monopodial trees occasionally with stilt roots or buttresses; crown often pyramidal.
Branches more or less horizontal and whorled. Bark brown or black (most species of
Myristica), smooth, fissured or flaky; inner bark fibrous, reddish brown; sap red. Sapwood
white, soon brownish on exposure, soft; rays narrow, tanniferous tubes present; growth
rings usually distinct and visible to the unaided eye. Twigs striate or smooth, often
lenticellate, their apical portions often tomentose. Leaves simple, entire, alternate, often
appearing distichous, exstipulate, often tomentose on both surfaces when young and later
glabrous or sometimes remaining tomentose on the lower surface; lateral veins distinct,
generally anastomosing near the leaf margin. Inflorescences axillary or borne on older
branches and stems, branched or unbranched panicles of subumbels, or fascicles of flowers
borne on scar-covered woody brachyblasts (in Knema and partly in Myristica); bracts
present, often caducous; bracteole absent or present; pedicels usually present. Flowers
radially symmetrical; buds mostly more or less campanulate, or subglobose, that of female
flowers usually larger than those of male ones, inside cream or red, often hairy outside,
sometimes fragrant; perianth gamophyllous, sometimes splits to the base, 25-lobed, lobes
valvate. Male flowers: stamens united into an elongated, globose, or pyramidal column,
stalked or sessile; anthers 245, extrorse, 2-loculed, adnate to the column, the latter
sometimes (Knema) expanded into a disc. Female flowers: ovary superior, unilocular,
subglobose, style almost absent, stigmas 2, with few to many laciniations; ovule 1,
anatropous, placentation sub-basal. Fruits subglobose to oblong, dehiscing by a circumferential
suture into 2 valves; pericarp coriaceous or fleshy, glabrous or hairy, sometimes lanose,
ripening yellow or red. Seeds single, large, hard, conform to the fruits; endosperm hard,
mostly ruminate; aril red or orange-red, waxy or fleshy, covering the seed, variously laciniate
or entire.
Vernacular names. Preferred trade namepenarahan. Sabahdarah-darah (preferred
name). Sarawakbinarah (Murut), bindara (Kelabit), jela bala (Kenyah), kayo bela
(Kayan), kayo raha (Berawan), kumpang (preferred name), pang (Bidayuh), pendarahan,
pumpu (Bidayuh Sadong), raha meban (Punan Tutoh).
335
Distribution. A pantropical family comprising 20 genera and c. 500 species. In Sabah

and Sarawak the family are represented by 5 genera with c. 110 species, of which several
species are local endemics.
Ecology. The Myristicaceae is a family of rain forest of mainly low and medium altitudes.
Sometimes they constitute a considerable component of the forest, especially of the middle
storey of the lowland rain forest, but not gregarious. Few species are confined to the
mountains. Flowering and fruiting generally throughout the year. Pollination presumably
is by insects (e.g., small beetles). The blackish seed, contrasting with the red aril and
the yellow or brown or pink opened pericarp, attracts birds (e.g., forest pigeons), which
disperse the seeds.
Uses. The timber, traded under the preferred name penarahan, is mostly soft or moderately
hard and moderately heavy, not durable, but easily treated with preservatives. It is easy
to work with, but sometimes easily splitting. The sapwood is pale, not well defined,
and the heartwood is often dark reddish brown. The wood is mainly used for light
construction work, partitioning, flooring, wall-panelling, moulding and other types of
interior finishing, and for making tool-handles, woodcrafts, match boxes and splints, packing
cases, crates, and household utensils. It is also suitable for manifacturing plywood, rotary
veneer and wrapping and writing papers. The seed (with endosperm) of Myristica fragrans
is the nutmeg of commerce (pala), while the aril around the seeds supplies the mace.
Seeds of some Knema species contain oil that is locally used to light lamps.
Taxonomic notes. All members of the Myristicaceae can be recognized in the field by
their general habit, i.e. slender bole, with monopodial crown, with the branches more
or less verticillate and tiered, the longish exstipulate distichous leaves, like in the
Annonaceae. These latter differ in flower structure, fruit, and lack the red exudate of
the cut bark. Further differences are obvious in the transversely cut twig where the bark
has radiating parenchyma in the Annonaceae, and also the pollen is different. The genera
of Myristicaceae can be told apart on vegetative characters only with special experience,
and flower, inflorescence and fruit characters are necessary for a definite identification.
Useful characters are the non-striate, usually finely lenticellate and granulate bark of the
twigs in Gymnacranthera, or the dry leaves not readily breaking up into pieces in most
of Knema and Gymnacranthera. Certain leaf and wood anatomical characters are diagnostic
of a particular genus. Since myristicaceous specimens are either male (staminate) flowering,
or female (pistillate) flowering and/or fruiting, keys have been constructed for both sexes,
using also certain vegetative characters. However, specimens with only pistillate flowers
may be problematic to assign to a particular genus, and one should search and attempt
within both keys, using especially the type of inflorescence.
Key to genera
(for male flowering specimens)
1.
Inflorescences sessile or short-peduncled fascicles of flowers borne on brachyblasts

(= short, tubercle-like, usually woody shoots) bearing scars of fallen pedicels and
bracts; bracteole present.......2
Inflorescences branched, short- or long-peduncled panicles, with distal parts of the
branches woody and bearing scars or not; bracteole present or absent..........3
336
2.
Androecium comprising a staminal disc to which the anthers are radially attached
by their bases, and a cylindrical or tapering stalk or staminal column. Bracteole minute,
situated at the base or at about the middle part of the pedicel.......................4. Knema
Androecium consisting of an elongated synandrium made up of a thickish central
column to which anthers are adnate by their backsides, a sterile apex and a stalk
or androphore. Bracteole embracing the flower bud, inserted at or near the top of
the pedicel........5. Myristica (in part)
3.
Bracteole present, inserted at or near the top of the pedicel. Inflorescences with scarcovered perennial distal portions present or absent........................5. Myristica (in part)
Bracteole absent. Inflorescences not as above.......4
4.
Synandrium (comprising a swollen, solid or excavated column to which elongated

anthers are completely or partially adnate by their backsides) subsessile or shortstalked, broadly ellipsoid or almost globose, usually considerably broader than the
stalk or androphore. Perianth inside glabrous, lobes not spreading at anthesis.
Inflorescences peduncled, not branched from the base.........3. Horsfieldia
Synandrium narrowly elongated or almost globose; the central column at apex excavated
or not, narrow, about as wide or narrower than the androphore. Perianth inside or
on both sides hairy or papillose-hairy, lobes erect or reflexed at anthesis. Inflorescences
usually branched at or near the base.................................5
5.
Synandrium elongated, sessile or very short-stalked; anthers laterally tightly connate

and for most of their backsides adnate to the excavated column, with apical free
portions. Perianth lobes (in male flowers) erect at anthesis.............2. Gymnacranthera
Synandrium short, depressed globose or short-ellipsoid, distinctly stalked (androphore
well-developed); anthers short, completely sessile, connate, and adnate to the slender,
non-excavated column by their backsides, without free apical portions. Perianth lobes
(in male flowers) spreading or reflexed at anthesis..............1. Endocomia
Key to genera
(for fruiting specimens)
1.
Aril divided into segments to or almost to the base....................2

Aril entire or laciniate (incised) in the apical portion only, for up to halfway from
the top................3
2.
Infructescences paniculate, usually branched from near the base and with scars of
cataphylls at base. Bark of twigs smooth, never distinctly striate. Fruits 1.53 cm
long. Bracteole absent..........................2. Gymnacranthera
Infructescences either simple or furcate, short, subsessile, or panicle-like with peduncle,
without scars of cataphylls at the base. Bark of twigs usually striate, or longitudinally
cracked, or flaky. Fruits generally larger. Bracteole (scar) on fruiting pedicel
present.......................5. Myristica
3.
Aril coarsely laciniate from the upper 1/41/2. Seeds usually variegated, often bluntly
pointed at the apex. Bracteole absent. Stigma narrowly 2few-lobed. Inflorescences
337
paniculate. Trees monoecious............1. Endocomia

Aril at apex convoluted or shallowly laciniate. Seeds not variegated, not pointed at
one end. Bracteole absent or present. Stigma 2many-lobed. Inflorescences paniculate
or flowers borne on subsessile brachyblasts. Trees dioecious.........................4
4.
Bracteole absent. Inflorescences paniculate. Stigma usually minutely 2-lobed. Leaves

when dry usually easily breaking up into pieces; lower surface usually not
glaucous................3. Horsfieldia
Bracteole present. Inflorescences short, simple or forked, scar-covered, subsessile
brachyblasts. Stigma fewmany-lobed. Leaves when dry generally not easily breaking
up into pieces; lower surface usually more or less glaucous...........4. Knema
1. ENDOCOMIA W.J. de Wilde

(Latin, endo = inside, within, coma = hair tuft; the presence of
hair tuft inside the perianth)
Blumea 30 (1984) 173; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 262; Coode et al. (eds.), CLBD
(1996) 220; PROSEA 5, 3 (1998) 214.
Monoecious trees. Twigs subterete, without lines from petiole to petiole; bark usually
not striate, when older longitudinally cracked and sometimes flaky; without lenticels.
Terminal leaf buds densely, minutely dull grey-brown pubescent. Leaves distichous,
distinctly petioled, blades not particularly brittle when dry, lower surface not glaucous,
without brownish or blackish dots; lateral veins flat to raised above. Inflorescences
paniculate, usually several times branched at or near the base, pubescent or glabrescent,
with either male or mixed male and female flowers, sometimes the female flowers dominant;
at base of peduncle with or without a few minute cataphylls. Flowers pedicelled, pubescent
or glabrescent, always in umbellate clusters or fascicles with the flowers specifically either
all in the same or in different stages of development; pedicel at base not articulated;
bract caducous; bracteole absent; perianth membranous to coriaceous, 35-lobed, at
anthesis clefts to c. 3/4 or nearly to the base, greenish or yellowish, inside papillosehairy to various degree, the lobes in male flowers at anthesis horizontally spreading or
more or less reflexed. Male flowers: buds broadly ellipsoid or subglobose, sometimes
faintly angular; synandrium depressed globose or short-ellipsoid, central staminal column
narrow, not wider than the androphore, at apex not excavated; androphore short to long,
cylindrical, distinct; anthers short, completely sessile, connate, and adnate to the staminal
column by their backsides, without free apical portion, 26, vertical, up to 0.5 mm long,
opening extrorsely. Female flowers: buds resembling the males; ovary ovoid, glabrous,
stigma sessile, narrowly 2few-lobed, 0.20.3 mm high, the lobes narrow or minutely
25-lobulate. Infructescences paniculate, up to 30 cm long, bearing a few to many fruits.
Fruits obovoid or ellipsoid, glabrous; perianth caducous. Seeds ellipsoid, often bluntly
pointed at the apex; testa usually variegated; aril either completely covering or shorter
than the seed, usually conspicuously deeply laciniated to 1/41/2 from the top.
Distribution. Four species, ranging from S China through SE Asia to Malesia but absent
338
from C and E Java and the Lesser Sunda Islands. In Sabah and Sarawak represented
by 3 species.
Ecology. Little is known, possibly not much diverging from that of the family.
Key to Endocomia species

1.
Male flowers in one cluster or semi-cluster all in about the same stage of development;
synandrium depressed globose; androphore shorter than the synandrium, and hence
the latter situated close to the bottom of the perianth. Anthers 4. Perianth of male
flowers (3)45-lobed. Leaf buds, twig apices and inflorescences rusty pubescent with
hairs (0.3)0.5(0.7) mm long, sometimes late-glabrescent......2. E. rufirachis
Male flowers in one cluster usually in different stages of development, i.e. flower
buds of different sizes; synandrium depressed globose or short-ellipsoid; androphore
about as long as or longer than the synandrium, hence the latter usually situated
well above the bottom of the perianth. Anthers mostly 3 or 4. Perianth of male flowers
34-lobed. Leaf buds, twig apices and inflorescences initially greyish or grey-brown
pubescent with minute hairs c. 0.1 mm long, early glabrescent............2
2.
Leaves 820 cm long; upper surface drying olivaceous; lateral veins 712 pairs.
Inflorescences 26-flowered. Anthers 34. Fruits 4.57 cm long; pericarp 58 mm
thick, drying brown..............3. E. virella
Leaves 1535 cm long; upper surface usually drying dark brown; lateral veins 11
24 pairs. Inflorescences 410-flowered. Anthers 23. Fruits 3.54(5.5) cm long;
pericarp 13 mm thick, drying blackish.............1. E. macrocoma
1. Endocomia macrocoma (Miq.) W.J. de Wilde
Fig. 1FH.
(Greek, makro = large; Latin, coma = hair tuft; hairy with long tufted hairs, the twings,
leaves and inflorescences)
Blumea 30 (1984) 182, ibid. 41 (1996) 375; Whitmore, Tantra & Sutisna l.c. 262. Basionym: Myristica
macrocoma Miq., Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 207, ibid. 2 (1865) 49, p.p. (excl. spec. Celeb. =
Horsfieldia irya). Lectotype (W.J. de Wilde, 1984): Teijsmann 5553, Maluku, Halmahera (U). Synonym:
Horsfieldia macrocoma (Miq.) Warb., Monogr. Myrist. (1897) 299, J. Sinclair, Gard. Bull. Sing. 16
(1958) 392, ibid 23 (1975) 75, p.p., Anderson l.c. 265.
subsp. longipes W.J. de Wilde

(Latin, longus = long, pes = foot; the long-stalked synandrium)
Blumea 30 (1984) 185. Type: De Vogel 888, Borneo, SE Kalimantan (holotype L; isotype K).
Tree up to 20 m tall. Bark greyish or chocolate-brown, somewhat fissured or not and

little to profusely scaling off in tiny pieces; inner bark 717 mm thick, cream or light
brown or light brown-red, or yellowish; sap pale orange or reddish, watery. Sapwood
pale yellow or pale brown. Twigs towards apex 26 mm diameter, initially grey-brown
to rusty tomentose with hairs c. 0.1 mm long, early glabrescent; bark of lower part tends
339
to be longitudinally cracked, sometimes flaky, lenticels absent or inconspicuous. Terminal

leaf buds 825 mm long, minutely tomentose with hairs c. 0.1 mm long. Leaves membranous
or chartaceous, upper surface drying dark brown, glabrous, lower surface glabrescent;
elliptic to oblanceolate, (12)1535 (4)510 cm, base rounded or attenuate, apex acuteacuminate; midrib flat or slightly raised above; lateral veins 1124 pairs, flat or slightly
raised above, interarching connection generally indistinct; intercostal venation sometimes
trabeculate, distinct on both surfaces or not; petiole 12.5 cm long, glabrescent. Inflorescences
630 cm long, early glabrescent or persistently pubescent with hairs c. 0.1 mm long;
peduncle up to 5 cm long; bract 1.53 mm long, caducous. Flowers arranged in lax
or dense umbel-like, widely spaced clusters of 410; flowers in one cluster usually in
different stages of development; perianth 34-lobed, outside glabrous, early glabrescent
or tomentose with thin hairs c. 0.1 mm long, lobes inside towards the apex with few
to many coarse pale brownish hairs 0.20.4 mm long, more or less arranged in rows
corresponding with the spaces between the anthers, or sometimes the hairs present only
near the lobe-sutures. Male flowers: pedicel slender, (2)47 mm long; buds broadly
ellipsoid or ovoid, 1.52.3 1.32 mm, apex rounded or subacute, base rounded or attenuate;
perianth 34-lobed, at anthesis clefts to 4/55/6, lobes 0.20.3(0.5) mm thick, inside
greenish or yellowish when fresh; synandrium depressed globose, 0.20.3 0.20.4 mm;
anthers 2 or 3; androphore longer than the synandrium, (0.6)0.71 mm long. Female
flowers: pedicel 26 mm long; buds ovoid-ellipsoid, 2.22.6 mm long; ovary narrowly
ovoid, 1.31.8 mm long, glabrous, stigma 0.20.3 mm long, narrowly 2-lipped. Infructescence
a pendent, loose panicle, (10)1530 cm long, bearing 312 fruits. Fruits narrowly ovoidellipsoid, 3.54(5.5) 1.42 cm, apex blunt or subacute, base rounded or narrowed,
glabrous; pericarp 13 mm thick, drying blackish, finely granulate, not or sparingly warted;
stalk c. 1.5 cm long. Seeds c. 3 cm long, apex pointed or with a beak up to c. 2 mm
long; aril at apex laciniate to 1/5.
Vernacular name. Sarawakkumpang limau (Iban).
Distribution. Sumatra and Borneo (Sabah, Sarawak, E and S Kalimantan).
Ecology. In primary mixed dipterocarp forest along stream banks with alluvial or deep
clay soils, and on low ridges with sandy, sedimentary soils, up to 1000 m altitude. Flowering
2. Endocomia rufirachis (J. Sinclair) W.J. de Wilde
Fig. 1AE.
(Latin, rufus = reddish; rachis = axis; the reddish tomentum on the inflorescence rachis)
Blumea 30 (1984) 192; Whitmore, Tantra & Sutisna l.c. 262. Basionym: Horsfieldia macrocoma (Miq.)
Warb. var. rufirachis J. Sinclair l.c. (1958) 393. Type: Wood SAN A 4770, Borneo, Sabah (holotype
SING, n.v.; isotypes K, L, SAN).
Tree up to 40 m tall; buttresses short and rounded. Bark usually blackish, brittle, smooth
with superficially longitudinal cracks or thinly flaky; inner bark 1015 mm thick, pale
yellow or brown to reddish brown or orange. Sapwood pale, whitish to brown-yellow,
soft; little pale red exudate from inner sapwood. Twigs towards apex 3.55 mm diameter,
apex initially rusty tomentose with hairs c. 0.5 mm long, late-glabrescent; bark lower
340
vi
341
part striate, tending to be flaky, lenticels inconspicuous. Terminal leaf buds 812 mm
long, rusty pubescent with hairs 0.30.5 mm long. Leaves membranous or thinly chartaceous,
upper surface drying brown, glabrous, lower surface early glabrescent; elliptic to lanceolate,
1838 611 cm, base subcordate or rounded or short-cuneate, apex acute-acuminate;
midrib glabrescent on both surfaces, flattish or moderately raised above; lateral veins
1725(30) pairs, partly raised above, interarching connection often distinct and regular;
intercostal venation distinct or faint above; petiole 0.82.5 cm long, glabrescent.
Inflorescences rusty pubescent with hairs 0.50.7 mm long, 35-times branched, manyflowered, 830 525 cm; peduncle 1.55 cm long; bract 26 mm long, caducous. Male
flowers in dense clusters of 510, sometimes in lax clusters of 2 or 3 only; all flowers
of one cluster in the same stage of development; pedicel slender, 23.5 mm long; buds
ovoid to more or less obconical, 1.51.8 1.21.5 mm, apex rounded to subtruncate;
perianth (3)45-lobed, pubescent with hairs 0.20.5 mm long outside, late-glabrescent,
lobes inside rusty pubescent with similar hairs arranged in few longitudinal rows, at anthesis
clefts to 3/44/5, the lobes spreading or more or less recurved, c. 0.2 mm thick; synandrium
depressed globose, (0.2)0.3 0.50.6 mm; anthers 4; androphore much shorter than
the synandrium, c. 0.3 0.20.3 mm. Female flowers not seen. Infructescences large
panicles up to 30 cm long, bearing 618 fruits. Fruits ellipsoid-oblong, glabrous, apex
narrowly rounded, base subacute or often tapering into a 410 mm long narrow portion,
4.56.5 1.82.5 cm, glabrous, drying blackish, finely granulate, without lenticels, not
warted; dry pericarp 1.53(4) mm thick; stalk 0.81.6 cm long. Seeds 3.54.5 cm long,
apex with a beak 0.24 mm long; aril laciniate at apex to 1/51/3.
Vernacular name. Sabahsikut nabalu (Dusun).
Distribution. Endemic to Borneo (Sabah, Sarawak and E Kalimantan).
Ecology. In primary and logged-over lowland rain forests on flat lands and hill slopes,
up to 400 m altitude; on leached clays, loam soils, black soils, also on sandstone and
limestone or on periodically inundated ground. Flowering and fruiting throughout the
year.
3. Endocomia virella W.J. de Wilde

(Latin, virellus = greenish; the drying leaves)
Blumea 30 (1984) 194; Whitmore, Tantra & Sutisna l.c. 262; Coode et al. (eds.) l.c. 220. Type: Sadau
SAN 49546, Borneo, Sabah (holotype L; isotypes K, SAN).
Tree up to 20 m tall. Bark greenish, brown-yellow, yellowish green or black, smooth

or scaly; inner bark either reddish, orange-yellow, or brownish; sap clear or red. Sapwood
white, soft or medium-hard. Twigs towards apex 1.54 mm diameter, apex initially greybrown tomentose with hairs up to 0.1 mm long, early glabrescent; lower part faintly
longitudinally cracked, not flaky, lenticels absent. Terminal leaf buds 610 0.61 mm,
sparsely pubescent with hairs up to 0.1 mm long, early glabrescent. Leaves membranous
or thinly chartaceous, upper surface drying olivaceous, glabrous, lower surface pale green,
glabrous; elliptic to oblong, 820 3.58 cm, base cuneate, apex acute-acuminate; midrib
flat above; lateral veins 712 pairs, flat or slightly raised above, interarching connection
342
distinct or not; intercostal venation indistinct on both surfaces; petiole slender, glabrous,
11.8 cm long. Inflorescences lax, initially sparsely tomentose with minute hairs less
than 0.1 mm long, early glabrescent or glabrous, 515 1.58(10) cm; peduncle 0.5
2.5 cm long; bract caducous. Male flowers in widely spaced, lax, umbel-like clusters
of 26, those in one cluster usually in different stages of development; pedicel slender,
46 mm long; buds broadly ellipsoid or broadly ovoid, 1.82 1.51.6 mm, apex rounded
or subacute, base broadly rounded, in cross-section more or less 3(4)-angular; perianth
34-lobed, outside at first with sparse tomentum of hairs less than 0.1 mm long, early
glabrescent; lobes inside towards the apex pubescent with pale hairs 0.20.4 mm long
arranged in a few rows corresponding with the spaces in between the anthers, at anthesis
clefts to c. 5/6, lobes spreading or more or less recurved, c. 0.2 mm thick; synandrium
depressed globose, 0.20.3 0.30.4 mm; anthers 3 or 4; androphore slender, 0.81
0.2 mm. Female flowers not seen. Infructescences little-branched panicles, 1225 cm
long, bearing 14 fruits. Fruits ellipsoid-obovoid to ellipsoid-oblong, (4.5)57 2.5
3.5 cm, apex rounded, base subattenuate or with a narrowed portion up to 7 mm long,
glabrous, finely granulate; pericarp more or less woody, 58 mm thick, drying bright
dark brown; stalk 2.53 cm long. Seeds c. 4 cm long, apex subacute; aril laciniate to
1/41/3.
Distribution. Endemic to Borneo (Sabah and Sarawak).
Ecology. In primary forest on hill sides and ridges, up to 400 m altitude, on brown
or blackish soils. Flowering in January and May; fruiting in January, August and October.
2. GYMNACRANTHERA (A. DC.) Warb.

(Greek, gymnos = naked; acron = top; anthera = anther)
penarahan (preferred name), darah-darah (Malay), kumpang (Iban)
Ber. Pharm. Ges. 2 (1892) 227; Monogr. Myrist. (1897) 131; Gamble, J. As. Soc. Beng. 75, 2 (1912)
222; Ridley, FMP 3 (1924) 61; J. Sinclair, Gard. Bull. Sing. 16 (1958) 434, ibid. 17 (1958) 96; Whitmore,
TFM 1 (1972) 318; Anderson, CLTS (1980) 263; Schouten, Blumea 31(1986) 451; Whitmore, Tantra &
Sutisna, CLK 2, 1 (1990) 262; PROSEA 5, 2 (1995) 255; Coode et al. (eds.), CLBD (1996) 221; Argent
et al. (eds.), MNDT-CK 2 (1997) 445. Basionym: Myristica sect. Gymnacranthera A. DC., Ann. Sc.
Nat. 4, 4 (1855) 31.
Dioecious trees or shrubs. Twigs terete or bluntly angular, appressed hairy, usually
glabrescent; bark smooth, when older finely cracked, never distinctly striate, always
lenticellate but sometimes inconspicuously so. Leaves distichous, membranous, chartaceous
or coriaceous, rather tough when dry, lower surface often glaucous, glabrescent or with
persistent tomentum of usually rather remote, appressed, inconspicuous hairs (tomentum
more or less persistent in G. bancana); lateral veins more or less flat above, flat to prominent
beneath; intercostal venation usually more or less indistinct on both surfaces. Inflorescences
axillary, rarely on the older branches, paniculate, glabrescent or pubescent to various
degree; at base of peduncle with few scars of minute cataphylls; bract small, caducous;
males usually several times branched at or near the base; females condensed, few-flowered.
343
Flowers pedicelled, pubescent to various degree, more or less solitary (in females) or
in loose clusters; all flowers in one inflorescence usually in about the same stage of
development; pedicel at base not articulated; bracteole absent; perianth thinly coriaceous,
(2)34-lobed, thinly appressed pubescent on both sides, outside yellow, inside never
red, lobes of various depth, in females deeper than in males, at anthesis in males erect
or slightly spreading, in females spreading or more or less reflexed. Male flowers: buds
ellipsoid or ellipsoid-oblong; androecium markedly uniform in all species, about as long
as the perianth tube (shorter in G. bancana); synandrium more or less cylindrical-ellipsoid,
sessile or on a short narrow stalk (androphore); anthers 512, laterally tightly connate
and for most of their backsides adnate to the central column, separated by deep clefts,
apices free, more or less erect or somewhat curved inward; central column with excavation
at the top, narrow, about as wide or narrower than the androphore. Female flowers:
buds smaller and usually wider than the males, ovoid or pyriform; pedicel shorter; ovary
subglobose or broadly ovoid, short-pubescent, stigma small, obliquely 2-lobed and each
lobe usually minutely 36-lobulate. Infructescences paniculate, few-branched from near
the base, with scars of cataphylls at the base. Fruits subglobose to ellipsoid, 1.53.5
cm long; pericarp fleshy but hard, drying brownish, short-pubescent or glabrescent; perianth
not persistent. Seeds ellipsoid; aril completely covering the seed, deeply laciniate to near
the base (as in Myristica); endosperm containing a fixed oil but no starch.
Distribution. Seven species, ranging from S India to S Thailand, Sumatra, Peninsular
Malaysia, Borneo, and eastwards to New Guinea and Bismarck Archipelago; not known
from Palawan, Java and the Lesser Sunda Islands. In Sabah and Sarawak represented
by 5 species.
Ecology. Scattered and frequent understorey trees in well-drained, primary and secondary
mixed dipterocarp forests, at altitude up to 1300 m. In peatswamp and kerangas forest
they constitute elements of the main canopy layer.
Key to Gymnacranthera species

1.
Male flowers 46 mm long; androecium much smaller than the perianth, and shorter
than the perianth tube. Tomentum of young twig and inflorescence composed of woolly
hairs (0.5)1(2) mm long. Lower leaf surface pubescent with woolly hairs c. 0.5
mm long. Leaves 1842 cm long; petiole 0.30.5 cm thick. Fruits 2.53.5 cm long,
pubescent with hairs 0.3(0.5) mm long.........................1. G. bancana
Male flowers 25 mm long; androecium relatively longer, about as long as the perianth
tube. Tomentum of young twig and inflorescence less conspicuous, glabrescent or
composed of hairs up to 0.5 mm long. Lower leaf surface glabrescent or with appressed
hairs less than 0.5 mm long. Leaves of various sizes; petiole 0.10.3 cm thick. Fruits
generally smaller, glabrescent.............................2
2.
Young twig, including the apical portion, conspicuously densely set with distinct
lenticels. Lower leaf surface pubescent with distinct, usually brown hairs; midrib flat
above. Lateral veins forming an angle of c. 45 with the midrib. Fruits ovoid-ellipsoid,
short-pubescent, c. 2 cm long, with truncate base........5. G. ocellata
Twig towards apex without or with fewer lenticels. Lower leaf surface glabrescent
344
or with scattered, inconspicuous, usually greyish or pale brown hairs; midrib flat
or sunken above. Lateral veins generally forming an angle of more than 45 with
the midrib. Fruits globose or ellipsoid-oblong, glabrescent or pubescent, of various
sizes, base not truncate................................3
3.
Twig usually slender, towards apex 12 mm diameter, about 10 cm lower down 2

3.5 mm diameter. Leaves 520 1.55.5(6) cm; midrib flat or sunken above.........
........3. G. farquhariana
Twig stout, towards apex (2)2.54(5) mm diameter, about 10 cm lower down (3)
3.55.5 mm diameter. Leaves 1433 (5.5)613 cm; midrib usually sunken
above................4
4.
Lateral veins on lower leaf surface distinct but only slightly prominent, at an angle
of 6070 with the midrib in the middle of the leaf. Leaves drying flat, not undulate.
Anthers 6(8), straight..................2. G. contracta
Lateral veins on lower leaf surface very distinct and prominent, at an angle of
(35) 4050(55) with the midrib in the middle of the leaf. Leaves generally drying
irregularly undulate, not flat. Anthers 610, sometimes twisted..............4. G. forbesii
1. Gymnacranthera bancana (Miq.) J. Sinclair
Fig. 2AF.
(of Bangka, Sumatra)

Gard. Bull. Sing. 16 (1958) 436, l.c. 17 (1958) 99; Schouten l.c. 463; Whitmore l.c. 319; Anderson l.c.
263; Whitmore, Tantra & Sutisna l.c. 262; Turner, Gard. Bull. Sing. 47 (1995) 357; Coode et al. (eds.)
l.c. 221. Basionym: Myristica bancana Miq., Fl. Ind. Bat., Suppl. 1 (1861) 383. Type: Teijsmann 3279,
Sumatra, Bangka (U). Synonyms: M. murtonii Hook. f., Fl. Brit. Ind. 5 (1886) 105; G. murtonii (Hook.
f.) Warb. l.c. (1897) 357; M. ferruginea Wall. ex King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 298; M.
amplifolia Warb. l.c. (1897) 517; G. murtonii (Hook. f.) Warb. var. borneensis Warb. l.c. (1897) 359; M.
murtonii Hook. f. var borneensis (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 88; G. bancana (Miq.)
J. Sinclair var. borneensis (Warb.) J. Sinclair l.c. 16 (1958) 439.
Tree up to 40 m tall; crown dense or spreading; bole smooth; buttresses absent. Bark
brown to grey, slightly fissured, finely or thickly flaky. Twigs stout, faintly angular,
towards apex 36 mm diameter, when young rusty brown tomentose with woolly hairs
(0.5)1(2) mm long; bark of lower part smooth or finely cracked, densely set with lenticels.
Terminal leaf buds 1020 mm long, pubescent with rusty hairs 0.51 mm long. Leaves
coriaceous, upper surface drying olivaceous, often shining, lower surface densely rusty
tomentose with woolly hairs c. 0.5 mm long, glabrescent, grey-brown; elliptic to lanceolate,
1842 7.519 cm, base attenuate to rounded or subcordate, margin often revolute, apex
acute-acuminate, acumen up to 1.5 cm long; midrib flat above; lateral veins (13)15
23 pairs, flat or slightly raised above, very prominent beneath; intercostal venation indistinct
on both surfaces; petiole late-glabrescent, 12 cm long, 0.30.5 cm thick. Inflorescences
pubescent with rusty woolly hairs 12 mm long; bract 24 mm long, caducous; males
610 cm long, many-flowered; females 25 cm long, few-flowered. Flowers rusty pubescent
on both sides with hairs 0.20.3 mm long. Male flowers: buds ellipsoid-oblong, 46
mm long; perianth 3(4)-lobed, at anthesis clefts to c. 1/4 to nearly halfway, tube 2.5
3.5 mm long; androecium truncately ellipsoid, subsessile, shorter than the perianth tube,
345
vi
1.52 mm long; anthers (7)910, free apices minute, more or less erect. Female flowers:
pedicel (known from immature flowers) c. 1 mm long; buds ovoid, 46 mm long; perianth
coriaceous, 3-lobed; ovary subglobose, densely pubescent. Fruits 28 per infructescence,
ellipsoid, 2.53.5 1.52.2 cm, rusty pubescent with hairs c. 0.3(0.5) mm long; pericarp
c. 3 mm thick; stalk 0.30.5 cm long.
Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo (Sarawak and Brunei).
Ecology. In primary and secondary well-drained forests, on hill sides and ridges, up to
250 m altitude, also in swamp forest, on granite rock, sand and sandy loam soils. Flowering
in SeptemberOctober; fruiting throughout the year.
2. Gymnacranthera contracta Warb.

(Latin, contractus = contracted; the condensed inflorescences)
Monogr. Myrist. (1897) 360; Merrill, EB (1921) 269; Masamune, EPB (1942) 300; J. Sinclair l.c. 16
(1958) 439, l.c. 17 (1958) 100, p.p.; Whitmore l.c. 320 (excl. Malayan specimens); Anderson l.c. 263;
Schouten l.c. 471; Whitmore, Tantra & Sutisna l.c. 262; Coode et al. (eds.) l.c. 221; Argent et al. (eds.)
l.c. 446. Synonyms: Myristica contracta (Warb.) Boerl. l.c. 88. Type: Beccari 321, Borneo, Sarawak
(holotype K; isotypes BO, C, FI, G, M, NY, P, S).
Tree up to 26 m tall. Bark reddish brown, nearly smooth, very finely scaly; inner bark
hard, pale reddish brown. Twigs stout, subterete to angular or more or less ridged, towards
apex 34(5) mm diameter, at first tomentose with very minute hairs up to 0.5 mm long,
glabrescent; bark chocolate to greyish brown, smooth, finely longitudinally cracked, with
lenticels when older. Terminal leaf buds 48 mm long, sometimes with additional cataphylls
(scales), minutely appressed pubescent. Leaves chartaceous to subcoriaceous, drying flat
and not undulate, upper surface drying brownish, often shining, lower surface pubescent
with scattered appressed hairs less than 0.5 mm long, glabrescent, greyish to pale brown;
oblong to lanceolate, more or less parallel-sided, (16)2029 69.5 cm, base shortattenuate to rounded, margin plane, apex acute-acuminate, acumen up to 2 cm long; midrib
slightly sunken above, narrow; lateral veins (11)1318 pairs, flat above, distinct but
not much prominent beneath, at an angle of 6070 with the midrib in the middle of
the leaf; intercostal venation indistinct on both surfaces; petiole 12(2.5) cm long, 0.1
0.3 cm thick. Inflorescences pubescent with rusty hairs 0.30.5 mm long; bract c. 2.5
3 mm, pubescent, caducous; males condensed panicles, 3.54.5 cm long; females
contracted, 1.52 cm long, fewer but more densely flowered. Flowers rusty pubescent
on both sides, hairs 0.10.2 mm long. Male flowers: pedicel 22.5 mm long; buds ellipsoidoblong, 2.52.8 mm long; perianth 34-lobed, at anthesis clefts to 1/3 to nearly 2/3, tube
11.8 mm long; androecium more or less truncately ellipsoid-oblong, c. 1.7 mm long
(i.e. about as long as the perianth tube), subsessile; anthers 6(8), the free apices minute,
more or less erect. Female flowers: pedicel 12 mm long; buds narrowly obovoid, 3
3.5 mm long; ovary subglobose, pubescent, c. 1.2 mm diameter. Fruits c. 7 per infructescence,
ellipsoid-oblong, 22.2 11.4 cm, pubescent with hairs c. 0.1 mm long or early
glabrescent, base not truncate; pericarp 1.21.4 mm thick; stalk 0.30.7 cm long.
Vernacular name. Sabahlanau (preferred name).
347
Distribution. Endemic to Borneo (Sabah, Sarawak, Brunei, and possibly W Kalimantan).

Ecology. In primary lowland forest, up to 600 m altitude. Flowering in JulySeptember;
fruiting in OctoberDecember.
3. Gymnacranthera farquhariana (Hook. f. & Thomson) Warb.

(W. Farquhar, 17701839, major in the British Army, the first colonial resident of Singapore)
Monogr. Myrist. (1897) 365; Gamble l.c. 225; Ridley l.c. 62; Schouten l.c. 476; Whitmore, Tantra &
Sutisna l.c. 262; Turner l.c. 357; Coode et al. (eds.) l.c. 221; Argent et al. (eds.) l.c. 446. Basionym:
Myristica farquhariana Wall. ex Hook. f. & Thomson, Fl. Ind. (1855) 161, p.p. Lectotype (Schouten,
1986): Wallich Cat. 6795, Singapore (K).
Tree up to 45 m tall; crown dense. Bark brown or grey, slightly fissured and finely
flaky or scaly; inner bark dark brown, 415 mm thick. Sapwood white to yellow; heartwood
yellow to brown, hard. Twigs subterete, slender, towards apex 12 mm diameter, at first
tomentose with minute hairs c. 0.1 mm long, early glabrescent; bark later on grey-brown,
smooth, lenticellate. Terminal leaf buds 510 mm long, pubescent with appressed greyish
hairs c. 0.1 mm long. Leaves chartaceous to coriaceous, upper surface drying olivaceous
or chocolate, sometimes shining, lower surface at first thinly tomentose with appressed
hairs c. 0.1 mm long, glabrescent, pale brown to grey-purplish; elliptic to lanceolate,
520 1.56 cm, base more or less attenuate, margin conspicuously revolute or not,
apex acute-acuminate, acumen up to 2 cm long; midrib flat or sunken above; lateral
veins 711(15) pairs, at an angle of more than 45 with the midrib, flat or raised beneath,
distinct and contrasting in colour with the lower leaf surface or not; intercostal venation
generally indistinct; petiole 0.61.5 cm long, 0.10.3 cm thick. Inflorescences paniculate,
pubescent with hairs c. 0.2 mm long; bract c. 1 mm long, caducous; males 2.512 cm
long, many-flowered; females up to 4 cm, few-flowered. Flowers rusty pubescent with
hairs c. 0.1 mm long on both sides. Male flowers: pedicel 1.54 mm long; buds ellipsoid
to ellipsoid-oblong, 2.54 mm long; perianth 34-lobed, at anthesis clefts to 1/32/3,
tube 12.3 mm long; androecium more or less truncately ellipsoid-oblong, 12.5 mm
long, subsessile; anthers (6)711(13), free apices up to 0.5 mm long, more or less
erect or somewhat incurved. Female flowers: pedicel 1.53 mm long; buds ovoid or
obpyriform, 23 mm long; ovary minutely pubescent, c. 1 mm diameter, stigma minute.
Fruits up to 13 per infructescence, subglobose or ellipsoid-oblong, 1.82.8 1.11.9
cm, base not truncate, glabrescent (glabrous) or inconspicuously pubescent with hairs
c. 0.1 mm long; pericarp c. 1 mm thick; stalk 0.41.5 cm long.
Vernacular name. Sarawakkumpang putih (Iban).
Distribution. A widespread and variable species, ranging from Peninsular Malaysia through
Malesia to the Bismarck Archipelago. Three closely related varieties occur in Sabah and
Sarawak.
Key to varieties
1.
Leaves oblong-lanceolate, 513.5 1.54.5 cm; lateral veins not or hardly raised
beneath....................................
348
var. eugeniifolia (A. DC.) R.T.A. Schouten

(with leaves resembling that of Eugenia, Myrtaceae)
l.c. 480; Turner l.c. 357; Coode et al. (eds.) l.c. 221. Basionym: Myristica eugeniifolia A. DC.,
Ann. Sc. Nat. 4, 4 (1855) 29. Type: Gaudichaud 116, Peninsular Malaysia, Penang (FI, G, P).
Synonyms: G. apiculata Warb. l.c. (1897) 359; M. apiculata (Warb.) Boerl. l.c. 88; G.
eugeniifolia (A. DC.) J. Sinclair l.c. 16 (1958) 444, p.p.; Whitmore l.c. 320; G. eugeniifolia A.
DC. var. eugeniifolia: J. Sinclair l.c. 16 (1958) 444, p.p., l.c. 17 (1958) 112, p.p.
Tree up to 30 m tall. Leaves chartaceous to coriaceous, widest at or below the

middle, margin little or conspicuously revolute or not; lateral veins 610 pairs;
petiole 0.71.4 cm long, 0.10.15 cm thick. Fruits 13(5) per infructescence,
1.82.2 1.31.8 cm; stalk 0.41.3 cm long.
Sumatra, Peninsular Malaysia, Singapore, and Borneo (Sabah, Sarawak). In primary
and degraded forests, on hill sides and ridges as well as in periodically inundated
places, near streams and rivers, and in kerangas forest, also found on limestone
and sandy soils, up to 1300 m altitude. Flowering mainly in MarchOctober;
fruiting in JulyFebruary.
Leaves elliptic, elliptic-oblong to lanceolate, 620 28.5 cm; lateral veins usually
clearly visible and contrasting, usually distinctly raised beneath..........2
2. Leaves coriaceous, elliptic to elliptic-oblong, 615(17) 35.5(6) cm, margin
conspicuously revolute. Fruits short-ellipsoid to globose.................
var. farquhariana
Turner l.c. 358; Coode et al. (eds.) l.c. 221. Synonyms: G. farquhariana (Hook. f. & Thomson)
Warb. var. griffithii (Hook. f.) Warb. l.c. (1897) 368; G. eugeniifolia (A. DC.) J. Sinclair var.
griffithii (Hook. f.) J. Sinclair l.c. 17 (1958) 113, Whitmore l.c. 320; Myristica farquhariana
Hook. f. & Thomson var. major King, l.c. 306; G. farquhariana (Hook. f. & Thomson) Warb.
var. major (King) Gamble l.c. 226.
Tree up to 30 m tall, occasionally with stilt roots. Leaves widest at or above

the middle; lateral veins 711 pairs; petiole 0.81.8 cm long, 0.150.2 cm thick.
Fruits 16 per infructescence, globose to short-ellipsoid; stalk 0.61.5 cm long.
Peninsular Thailand, Sumatra, Peninsular Malaysia, Singapore, and Borneo (Sabah
and Sarawak). In primary and degraded forests, mostly in peatswamp forest but
occasionally also found on hill sides, up to 1000 m altitude. Flowering mainly
in AprilJuly; fruiting in SeptemberMarch.
Leaves membranous, chartaceous or sometimes coriaceous, elliptic-oblong to lanceolate,
820 26 cm, margin plane, rarely revolute. Fruits ellipsoid to oblong...............................
var. zippeliana (Miq.) R.T.A. Schouten
(A. Zippelius, 17971828, horticulturist, sometime assistant curator of the Bogor
Botanic Gardens)
l.c. 482; Turner l.c. 358. Basionym: Myristica zippeliana Miq. l.c. 2 (1865) 50. Type: Zippelius
s.n., New Guinea, Vogelkop (L). Synonyms: G. zippeliana (Miq.) Warb. l.c. (1897) 373; G.
paniculata (A. DC.) Warb. var. zippeliana (Miq.) J. Sinclair l.c. 17 (1958) 108; G. suluensis
Warb. l.c. (1897) 373.
Tree up to 45 m tall. Leaves widest at or above the middle; lateral veins 8

11(15) pairs, pale yellowish and little contrasting in colour; petiole 0.71.5 cm
long, 0.10.2 cm thick. Fruits (1)213 per infructescence, (1.5)1.82.5 1.1
1.5 cm; stalk 0.40.8 cm long.
Sumatra, Peninsular Malaysia, Singapore, Borneo (Sabah and Sarawak), the
Philippines, Sulawesi, Maluku, and New Guinea. In primary and degraded forests,
349
mostly on hill sides and ridges, on sandstone, clay, and loam, and granite rock,
up to 900 m altitude. In Borneo, all records from 4001200 m. Locally abundant,
frequently mentioned as part of regenerating forest. Flowering mainly in July
November; fruiting in MarchMay and OctoberNovember.
4. Gymnacranthera forbesii (King) Warb.

(H.O. Forbes, 18511932, Scottish botanist)
Monogr. Myrist. (1897) 363; Gamble l.c. 224; Ridley l.c. 61; J. Sinclair l.c. 16 (1958) 441, l.c. 17 (1958)
101; Whitmore l.c. 320; Anderson l.c. 263; Schouten l.c. 472; Turner l.c. 358; Argent et al. (eds.) l.c.
446. Basionym: Myristica forbesii King l.c. 306. Lectotype (Schouten, 1986): Forbes 2976, Sumatra
(hololectotype K; isolectotypes BM, L).
Tree up to 35 m tall. Bark grey-brown, smooth or finely fissured or sometimes flaky;

inner bark pink to red-brown, laminated, sometimes fibrous. Sapwood white, pale yellow
to orange-brown. Twigs stout, subterete, towards apex (2)2.44 mm diameter, without
or with a few lenticels, early glabrescent, rarely with sparse pubescence comprising hairs
up to 0.1 mm long on young twigs; bark later on brown or grey, finely cracked, with
many lenticels. Terminal leaf buds 510 mm long, greyish tomentose with hairs c. 0.1
mm long. Leaves chartaceous or coriaceous, upper surface drying often coarsely undulate
(not flat), olivaceous brown, sometimes shining, lower surface at first tomentose with
scattered appressed minute hairs less than 0.5 mm long, glabrescent, grey-brown; elliptic
to lanceolate, 1433 (5.5)613 cm, base attenuate to broadly rounded, margin plane,
apex acute-acuminate, acumen up to 1 cm long; midrib sunken above; lateral veins 10
18 pairs, flat above, yellowish, very prominent beneath, at an angle of (35)4050
(55) with the midrib in the middle of the leaf; intercostal venation sometimes distinct
on the lower surface; petiole 0.82 cm long, 0.10.3 cm thick. Inflorescences paniculate,
pubescent with grey-rusty woolly hairs 0.20.4 mm long; bract c. 3 mm long, caducous;
males 412 cm long, many-flowered; females 14 cm long, generally few-flowered. Flowers
grey-rusty pubescent on both sides with hairs 0.10.2 mm long. Male flowers: pedicel
23 mm long; buds ellipsoid-oblong, 2.54 mm long; perianth 3(4)-lobed, at anthesis
clefts to c. 1/3 to nearly halfway, tube 1.52.3 mm long; androecium more or less truncately
ellipsoid-oblong, subsessile, 1.52.3 mm long; anthers 610, often somewhat twisted, free
apices 0.30.5 mm long, more or less erect. Female flowers: pedicel c. 2 mm long;
buds ovoid or pyriform, 22.5 mm long; ovary 11.3 mm diameter, minutely pubescent;
stigma minutely 2-lobed. Fruits 425 per infructescence, ellipsoid-oblong, 1.82.4 1
1.4 cm, base not truncate, glabrescent or short-pubescent with hairs c. 0.1 mm long;
pericarp c. 1 mm thick; stalk 0.41 cm long.
Distribution. S Thailand, Sumatra, Peninsular Malaysia, Singapore, and Borneo.
Key to varieties
Leaves thickly coriaceous; lateral veins on lower surface prominent and thick, on the
average 0.50.7 mm wide. Infructescences conspicuously branched from the base, manyfruited...........................
var. crassinervis (Warb.) J. Sinclair
350
(Latin, crassinervis = prominently veined; the leaf undersurface)

l.c. 17 (1958), 102; Schouten l.c. 475. Basionym: G. crassinervis Warb. l.c. (1897) 362. Lectotype:
Beccari 1119, Borneo, Sarawak (K). Synonym: Myristica crassinervis (Warb.) Boerl. l.c. 88.
Leaves elliptic-oblong to oblong-lanceolate, 1633 612 cm. Infructescence bearing

825 fruits.
Endemic to Borneo (Sabah, Sarawak, Brunei, and Kalimantan). In primary and degraded
well-drained as well as swampy forests, on sandy, sandy-clay, and loamy soils, up
to 1250 m altitude. Flowering mainly in April and August; fruiting in April and August
December.
Leaves chartaceous to coriaceous; lateral veins on lower surface moderately prominent,
0.30.5 mm wide. Infructescences usually not much branched at the base, few-fruited.............
var. forbesii
J. Sinclair l.c. 17 (1958), 101; Schouten l.c. 474; Coode et al. (eds.) l.c. 221.
Leaves elliptic to oblong, 1428 (5.5)613 cm. Infructescences bearing 410 fruits.
S Thailand (Pattani), Sumatra, Peninsular Malaysia, Singapore, and Borneo (several
collections from Sabah and Sarawak). In primary and secondary forests on hill sides
and riverine alluvial forest, on sandy and limestone- as well as sandstone-derived
soils, up to 600 m altitude. Flowering in FebruaryApril and AugustSeptember;
fruiting in MayAugust and DecemberJanuary.
Vernacular names. Sarawakdarah-darah (Malay), kumpang (Bidayuh), lahu (Kayan).
5. Gymnacranthera ocellata R.T.A. Schouten
Fig. 2GH.
(Latin, ocellus = a little eye; the many lenticels on the twigs)

Blumea 31 (1986) 469; Coode et al. (eds.) l.c. 221; Argent et al. (eds.) l.c. 446. Type: Saikeh SAN
72177, Borneo, Sabah, Beaufort Hill (holotype L; isotypes K, KEP, SAN, SAR, SING).
Tree up to 25 m tall. Bark smooth or fissured or regularly cracked, sometimes flaky,

grey to dark red-brown; inner bark 1015 mm thick, brown. Sapwood whitish streaked
with pale red; heartwood light brown to blackish brown. Twigs usually angular or
compressed, stout, towards apex 23.5(5) mm diameter, tomentose with appressed hairs
c. 0.2 mm long, glabrescent; bark usually grey, smooth or finely cracked, including that
of apical portion of the twig densely set with conspicuous lenticels of mixed sizes; at
base of each seasonal shoot with distinct scars of cataphylls of bud scales. Terminal leaf
buds rather stout, c. 10 3 mm, densely appressed pubescent. Leaves chartaceous or
thinly coriaceous, upper surface drying olivaceous, often shining, lower surface with distinct
more or less spaced appressed rusty brown hairs less than 0.5 mm long, glabrescent,
grey-brown; ovate-elliptic to lanceolate, 1025 49.5 cm, base short-attenuate to broadly
rounded, margin plane, apex acute; midrib flat above; lateral veins 1118 pairs, flat to
slightly raised above, distinct but not prominent beneath, at an angle of c. 45 with the
midrib in the middle of the leaf; intercostal venation indistinct on both surfaces; petiole
0.91.8 cm long, 0.20.3 cm thick. Inflorescences broadly paniculate, pubescent with
rusty hairs 0.30.5 mm long; bract c. 2.5 3 mm, caducous; males 38.5 cm long, manyflowered; females up to 2 cm long, few-flowered. Flowers rusty pubescent on both sides
with hairs 0.1(0.2) mm long. Male flowers: pedicel 1.53.5 mm long; buds ellipsoidoblong, 34(5) mm long; perianth 34-lobed, at anthesis clefts to 1/31/2, tube 22.5
mm long; androecium more or less truncately ellipsoid, subsessile, 1.52.3 mm long; anthers
351
710, the free apices 0.40.5 mm long, more or less erect. Female flowers: pedicel 1.5
2.5 mm long; buds narrowly ovoid, 2.73 mm long; ovary 1.21.5 mm diameter, minutely
pubescent, stigma shallowly 2-lobed. Fruits 410 per infructescence, ovoid-ellipsoid with
truncate base, 1.82.2 1.11.3 cm, short-pubescent with hairs 0.10.2 mm long,
glabrescent; pericarp 0.71.3 mm thick; stalk 0.30.7 cm long.
Vernacular name. Sarawakperupak (Kelabit).
Distribution. Endemic to Borneo (Sabah, Sarawak, Brunei, and Kalimantan).
Ecology. In primary well-drained forest on low hills and ridges, also in kerangas forest,
on tuff, sandy, and sandy loam soils, up to 1300 m altitude. Flowering mainly in June
November; fruiting in JulyDecember.
3. HORSFIELDIA Willd.
(T. Horsfield, 17731859, sometime a medical doctor and naturalist in Indonesia)
darah-darah (Malay), pisiah (Bidayuh, Padawan),
umpo (Bidayuh, Sadong), upang (Bidayuh, Bau)
Sp. Pl. 4 (1806) 872; Warburg, Monogr. Myrist. (1897) 262; Gamble, J. As. Soc. Beng. 75, 2 (1912)
206; Ridley, FMP 3 (1924) 54; J. Sinclair, Gard. Bull. Sing. 16 (1958) 368, ibid. 27 (1974) 133, ibid. 28
(1975) 1; Backer & Bakhuizen f., FJ 1 (1964) 138; Whitmore, TFM 1 (1972) 322; Anderson, CLTS
(1980) 263; W.J. de Wilde, Gard. Bull. Sing. 37 (1984) 115, ibid. 38 (1985) 55, 185, ibid. 39 (1986) 1;
Blumea 32 (1987) 459, ibid. 41 (1996) 377; Whitmore, Tantra & Sutisna, CLK 2, 1 (1990) 263; Coode
et al. (eds.), CLBD (1996) 221; Argent et al. (eds.), MNDT-CK 2 (1997) 447.
Dioecious shrubs or trees; occasionally buttressesed. Twigs usually glabrescent, subterete

or sometimes angular or ridged; bark usually striate, lenticellate. Leaves distichous or
in 3 or more rows along the twigs; blades up to 45 cm long, membranous to coriaceous,
often brittle when dry, pubescent or glabrescent, lower surface not glaucous, in some
species non-traumatic cork warts present; intercostal venation reticulate, usually lax above,
never forming a dense, close network as in Knema. Inflorescences rarely on the older
branches (H. sabulosa), paniculate, peduncled, usually branched several times but not
from the base, pubescent or glabrescent; at base of peduncle with few minute cataphylls;
females smaller; bract caducous. Flowers usually pedicelled, glabrous or pubescent, solitary
or in clusters, those in one inflorescence generally of about the same stage of development;
pedicel at base articulated or not; perianth thin-coriaceous to succulent, 24-lobed, inside
glabrous, greenish to yellowish, never red, the lobes cleft to various depths, at anthesis
never spreading; bracteole absent. Male flowers: buds usually small, globose, transversely
ellipsoid, reniform, pear-shaped, or clavate, laterally compressed or not; synandrium cupshaped, or almost globose to broadly ellipsoid, or trigonous, laterally compressed or not,
sessile or with short androphore; anthers 225, straight or curved or the apical parts
incurved into the cavity of the central column, entirely or largely connate, and adnate
by their backsides to the narrow or broad, variously excavated central column. Female
flowers: buds usually larger than the males; ovary globose or ovoid, glabrous or pubescent,
352
style absent, stigma minutely 2-lobed. Infructescences paniculate. Fruits globose or usually
ellipsoid; pericarp usually fleshy, glabrous or pubescent, or subglabrescent; perianth
sometimes persistent. Seeds ellipsoid, rarely globose, not pointed at one end; testa not
variegated; aril completely covering the seed, entire or at apex shallowly lobed or
convoluted; endosperm with fatty oil but no starch.
Distribution. More than 100 species ranging from Sri Lanka through NE India and S
China to Malesia and the Caroline Islands and eastwards to the Solomon Islands and
N Australia; absent from the Lesser Sunda Islands. Apart from a few widely distributed
species, e.g. H. irya, most species are of limited distribution. In Sabah and Sarawak
represented by 39 species, of which 9 species and 1 subspecies are endemic.
Ecology. In primary rain forest, often persisting in secondary growth, also in marshy
forest. Stilt roots present in some species. Some species occur in montane altitudes, and
the wide altitudinal range of the genus more or less contrasts with those of the other
genera.
Key to Horsfieldia species

(for male flowering specimens)
1.
Perianth predominantly 2-lobed, sometimes that of odd flower in one inflorescence

3- or 4-lobed.............2
Perianth predominantly 3- or 4-lobed.............5
2.
Leaves membranous, usually drying with whitish blotches of irregular shape and size.
Androecium: anthers for the greater part connate, forming a saucer-shaped column;
androphore distinct, tapering...........13. H. irya
Leaves coriaceous or membranous, without whitish blotches. Androecium
different........3
3.
Leaves coriaceous, lower surface with thin tomentum and with dark dots and
streaks........6. H. crassifolia
Leaves more or less membranous, lower surface glabrous or early glabrescent, without
dark dots or streaks.........4
4.
Androecium depressed globose, largely consisting of anthers; apical cavity of column

flat and shallow; anthers 7........35. H. sucosa
Androecium broadly obovoid, consisting of a large sterile basal part and at apex
with 3 or 4 small anthers............33. H. sterilis
5.
Leaves in 3 or more rows along the twigs. Leaf bud short and broad (less than 4
times longer than broad)..........6
Leaves distichous (in rare cases a few on the same plant in 3 rows). Leaf bud more
slender.........8
6.
Leaves more or less clustered towards the end of the twigs. Leaf bud and inflorescence
pubescent with hairs 0.51.5 mm long. Bark of older twigs often blackish, conspicuously
353
flaking...............................30. H. sabulosa
Leaves clustered or not. Leaf bud and inflorescence pubescent with hairs up to 0.3
mm long. Bark of older twigs not or slightly flaking..........7
7.
Bark of twig brown, not contrasting with the colour of drying petiole....
........10. H. fragillima (in part)
Bark of twig pale, greyish or straw, contrasting with the blackish brown colour of
the drying petiole.....22. H. pallidicaula (in part)
8.
Leaves scabrous above........12. H. grandis

Leaves not scabrous above........9
9.
Petiole almost absent. Twigs 57(10) mm diameter. Leaves 2570 cm long, with
persistent tomentum beneath..............31. H. sessilifolia
Petiole present. Twigs generally more slender. Leaves various........10
10. Flower bud ellipsoid, 2.83.7 mm long; perianth at anthesis clefts to 1/51/3. Leaves
more or less parchment-like, bullate (matted above with finely wrinkled surface)
........37. H. tristis
Flower bud either broadly ellipsoid, subglobose, broadly obovoid, or short pear-shaped,
c. 3.5 mm long or less, or (depressed)globose; perianth at anthesis clefts to (1/4)
1/3 or more. Texture of leaves various, not bullate....................11
11. Leaves with (sub)persistent tomentum beneath. (In H. gracilis and H. wallichii
sometimes hairs remaining only on and near midrib and lateral veins).......12
Leaves glabrous or early glabrescent beneath (in some species with hairs remaining
on the lower midrib)...........18
12. Leaves with dots and/or streaks beneath.......13
Leaves without dots or streaks beneath........14
13. Flower bud broadly obovoid, 22.5 mm long; pedicel tapering, 0.31 mm long. Twig
36(9) mm diameter. Leaves thinly pubescent beneath, often somewhat
glabrescent.........38. H. wallichii (in part)
Flower bud subglobose, 11.5 mm diameter; pedicel slender, terete, 0.51.5 mm long.
Twig 23 mm diameter. Leaves with conspicuous tomentum beneath...................
.........23. H. paucinervis
14. Flower bud c. 1 mm diameter, with persistent tomentum or late-glabrescent...
.....17. H. motleyi
Flower bud 12.5 mm diameter, glabrous or glabrescent........15
15. Pedicel not or indistinctly articulated at the base (character not quite clear in H.
rufo-lanata and H. gracilis)...............16
Pedicel articulated at the base............27. H. reticulata
16. Flower bud c. 1 mm diameter. Twig at apex 1.52.5 mm diameter. Leaves: intercostal
venation indistinct above; lower surface with hairs remaining on lateral veins and
354
midrib only...................11. H. gracilis

Flower bud 1.52.3 mm diameter. Twig at apex 3.57(13) mm diameter. Leaves:
intercostal venation distinct above; persistent tomentum distinct beneath.........17
17. Leaves 1845 cm long; lateral veins 1825 pairs, sunken above. Flower bud 1.5
2 mm diameter; anther 810......32. H. splendida
Leaves 1023 cm long; lateral veins 1116 pairs, raised above. Flower bud 22.3
mm diameter; anthers c. 15....................29. H. rufo-lanata
18. Twig pale, greyish or straw, contrasting with the dark brown colour of the drying
petiole....................19
Twig (dark) grey-brown, not contrasting with the drying petiole.................22
19. Androecium triangular in cross section. Anthers free for the upper half or more.
........21. H. oligocarpa
Androecium (sub)circular in cross section. Anthers largely connate......20
20. Leaves chartaceous to coriaceous; upper surface drying bright brown. Mostly in
kerangas forest on sandy soils........................5. H. carnosa
Leaves membranous or thinly chartaceous, upper surface drying (blackish) brown.
Usually growing in forest on richer soils, including sandy soils..................21
21. Fruit 4 cm long or less....22. H. pallidicaula (in part)
Fruit 5 cm long or more. (Male flowers not known).........7. H. discolor
22. Twig (also the older ones) ridged or nearly winged.........4. H. brachiata
Twig not ridged, sometimes faintly ridged, lined, or angular in the apical part only..23
23. Flower bud short pear-shaped, 22.5 mm long, subsessile with the pedicel thickish,
much shorter than the perianth, 0.31 mm long. Leaves with fine dots and streaks
beneath. Twig hollow......................38. H. wallichii (in part)
Flower bud various, the pedicel proportionally longer and more slender. Leaves with
or without dots (never streaks) beneath. Twig solid or faintly hollow.......................24
24. Androecium 3- or 4-angular in cross section; anthers more or less erect, free for
about halfway or more. Leaf without blackish dots beneath. Pedicel indistinctly
articulated....25
Androecium circular, subcircular, or subtriangular with rounded angles in cross section;
anthers more or less curved, almost entirely connate. Leaf with or without dots beneath.
Pedicel articulated or not.............................32
25. Leaves with midrib and lateral veins more or less level or sunken above.....26
Leaves with midrib and lateral veins raised above.....27
26. Twig and inflorescence glabrescent. Leaf apex acute or acute-acuminate; lateral veins
faint above. Anthers 46..............28. H. ridleyana
Twig late-glabrescent, inflorescence with persistent tomentum. Leaf apex blunt; lateral
veins flat or slightly raised above, clearly visible. Anthers 9 or 10...20. H. obtusa
355
27. Leaves and midrib glabrous beneath; apex long acute-acuminate. Twig rather smooth,
lower down longitudinally cracked.............8. H. disticha
Leaves early glabrescent beneath, but midrib sometimes late-glabrescent; apex acuteacuminate. Twig striate, lower down coarsely striate or finely cracked..................28
28. Twig 13(4) mm diameter. Leaves thinly membranous to subchartaceous; petiole
0.10.15(0.2) cm diameter. Inflorescence delicate, up to 10 cm long..........29
Twig 25(13) mm diameter. Leaves chartaceous to thickly coriaceous; petiole 0.15
0.4(0.8) cm diameter. Inflorescence up to 15(20) cm long.........30
29. Leaf bud pubescent with hairs (0.1)0.2(3) mm long. Twig at apex glabrescent.
Inflorescence pubescent with sparse stellate hairs 0.10.2 mm long, glabrescent. Leaves
glabrous, upper surface drying olivaceous to brown. Flower bud subglobose, tapering
into the pedicel................................36. H. tenuifolia
Leaf bud, apical part of twig, petiole, midrib beneath and inflorescence pubescent
with woolly stellate-dendroid hairs (0.2)0.5 mm long. Upper surface of leaves drying
dull olivaceous. Flower bud globose or depressed globose..15. H. macilenta
30. Twig and leaves stout; midrib broad above, at the transition to the petiole at least
3 mm wide. Inflorescence 1020 cm long.....14. H. laticostata
Twig and leaves less robust; midrib above towards the insertion of the petiole narrow,
less than 3 mm wide. Inflorescence up to 15(20) cm long..................31
31. Leaves 1628 cm long; base more or less rounded or short-attenuate; lateral veins
1619 pairs, very prominent above.......18. H. nervosa
Leaves 728 cm long; base short- to long-attenuate; lateral veins 615 pairs, raised
to various degree above. (On drying, colour of leaves above and beneath usually
much contrasting, generally more so than in the related species)..25. H. polyspherula
32. Leaf
long
Leaf
long
bud, apical part of twig, and inflorescence tomentose with hairs c. 0.2 mm
or more....33
bud, apical part of twig, and inflorescence tomentose with hairs 0.1(0.2) mm
or less.........................38
33. Flower bud ellipsoid, 2.53.5 mm long; androecium longer than broad. Leaves thick
and brittle, lower surface usually with conspicuous pale golden hair scars
(lens!).......9. H. endertii
Flower bud depressed globose or (sub)globose, (1)1.42.2(2.5) mm diameter;
androecium not longer than broad. Leaves various, without golden hair scars...34
34. Leaves with dots beneath (lens!). Pedicel articulated.........3. H. borneensis
Leaves without dots beneath. Pedicel not articulated...............35
35. Male flower bud 22.5 mm diameter; androecium sessile, broadly saucer-shaped.
Leaves 2045 cm long. Lowland forest...........10. H. fragillima (in part)
Male flower bud (1)1.42.2 mm diameter; androecium (depressed)globose, sessile
or with short androphore, the apical cavity small, concealed by the apices of the
anthers. Leaves 435 cm long. Hill to montane forests at 8002000 m altitude.36
356
36. Androecium with slender androphore 0.40.8 mm long, not hidden by the anthers.
..........2. H. androphora
Androecium (sub)sessile, androphore absent or up to 0.5 mm long, largely hidden
by the anthers.................37
37. Leaves chartaceous or membranous, up to 35 cm long; upper surface drying olivaceous;
apex acute-acuminate. Inflorescence up to 20 cm long...........................
.....1. H. amplomontana
Leaves coriaceous, 414 cm long; upper surface drying blackish; apex obtuse to
subacute. Inflorescence 416 cm long.........16. H. montana
38. Leaves with dots beneath..........41
Leaves without dots beneath (dots should not be confused with smaller, blackish
points)........39
39. Perianth coriaceous, lobes at base (0.3)0.41 mm thick. Androecium broadly obovoid,
longer than broad; androphore rather broad, tapering, (0.1)0.20.3 mm long. Leaves
coriaceous....................39. H. xanthina
Perianth thinner, lobes at base 0.20.3 mm thick. Androecium subglobose, depressed
globose-ovoid, ellipsoid, or broadly obovoid, not or little longer than broad; androphore
narrow, only c. 0.2 mm long. Leaves membranous or chartaceous...........40
40. Perianth of flower bud clefts for c. 1/2 at anthesis.......34. H. subalpina
Perianth of flower bud clefts for 2/34/5 at anthesis..19. H. obscura
41. Twig 2.54(10) mm diameter. Leaves 921 cm long or more. Flower bud depressed
globose, 1.62.2 mm diameter..........26. H. punctatifolia
Twig 1.52(4) mm diameter. Leaves 614(17) cm long. Flower bud subglobose,
c. (1.5)2 mm diameter........24. H. penangiana
Key to Horsfieldia species

(for female flowering and fruiting specimens)
1.
Perianth 2-lobed...................2
Perianth predominantly 34-lobed................5
2.
Leaves membranous, upper surface usually with irregularly shaped whitish blotches
here and there. Fruits globose. Seeds globose......................13. H. irya
Leaf texture various, upper surface usually without whitish blotches. Fruits subglobose,
ovoid, obovoid, oblong, or ellipsoid. Seeds ellipsoid or almost globose.3
3.
Leaves coriaceous; lower surface usually densely short-pubescent and with scattered
brownish or blackish dots and streaks. Twig grey-brown, not much contrasting with
the colour of the drying petiole..................6. H. crassifolia
Leaves membranous; lower surface glabrous or early glabrescent, without brownish
or blackish dots or streaks. Twig greyish or straw, rather contrasting with the blackish
colour of the drying petiole...........................4
357
4.
Female inflorescences more or less spike-like, 510 cm long. Perianth persistent under
the fruit....................33. H. sterilis
Female inflorescences branched, 12 cm long. Perianth not persisting under the
fruit...........35. H. sucosa
5.
Leaves in 3 or more rows along the twig................6

Leaves distichous.........................8
6.
Leaves generally bunched towards the apex of the twig. Petiole 2.55 cm long.
.......30. H. sabulosa
Leaves not bunched at the apex of twig. Petiole (0.2)0.42.5 cm long..................7
7.
Leaves 1030 cm long. Twig pale greyish or straw, contrasting with the blackish
colour of the drying petiole. Female flower buds 2.53 mm long. Fruits 1.54 cm
long......22. H. pallidicaula (in part)
Leaves 2045 cm long. Twig brown, in colour not contrasting with the drying petiole.
Female flower buds 45 mm long. Fruits 68 cm long........10. H. fragillima (in part)
8.
Apical portion of twig angular, i.e. distinctly lined or ridged from petiole to
petiole...........4. H. brachiata
Apical portion of twig terete, subterete or faintly angular, neither distinctly lined
nor ridged..........9
9.
Leaves on lower surface with persistent or subpersistent tomentum.................10

Leaves on lower surface glabrous or early glabrescent (midrib sometimes lateglabrescent)......................18
10. Leaves c. 50 cm long; petiole 0.20.3 cm long............31. H. sessilifolia

Leaves usually less than 40 cm long; petiole much longer...............11
11. Older leaves on both surfaces with scabrous hair scars. Fruits 11.4 cm long.........
........12. H. grandis
Leaves not scabrous. Fruits generally larger...................12
12. Leaves on lower surface with scattered dark brown or blackish dots and/or
streaks...................38. H. wallichii (in part)
Lower leaf surface with or without dots, but always without streaks.............13
13. Perianth with persistent tomentum, not persistent under mature fruit......17. H. motleyi
Perianth glabrous or early glabrescent, persistent under mature fruit or not.......14
14. Ovary pubescent. Fruit sometimes pubescent only towards the base..........15
Ovary and fruit glabrous...............16
15. Twig moderately stout, towards apex 3.55(7) mm diameter. Leaves 1023 cm long;
lateral veins 1016 pairs. Fruits largely glabrescent; perianth not persistent......
.........29. H. rufo-lanata
Twig stouter, toward apex 47(13) mm diameter. Leaves 1845 cm long; lateral
veins 1825 pairs. Fruits pubescent, with persistent perianth......32. H. splendida
358
16. Twig rather stout, towards apex 2.56(9) mm diameter. Leaves 1835 cm long; surface
sometimes bullate; lateral veins (9)1720 pairs. Fruits 2.32.7 cm long.......
......................................................27. H. reticulata
Twig slender, towards apex 1.53 mm diameter. Leaves generally smaller; surface
not bullate; lateral veins 517 pairs. Fruits 11.5 cm long.................17
17. Leaves membranous; lateral veins 1217 pairs; brownish or blackish dots on lower
leaf surface absent...............11. H. gracilis
Leaves thinly chartaceous; lateral veins 59(11) pairs; brownish or blackish dots
on lower leaf surface present..........23. H. paucinervis
18. Lower leaf surface with more or less regularly spaced, brownish or blackish dots
or with dots and streaks (dots originating from hair scars, not to be confused with
usually smaller and irregularly spaced dots or punctation of different origin)....19
Lower leaf surface without blackish dots (sometimes with enlarged hair scars)....22
19. Tomentum of leaf buds, twig apices and young inflorescences composed of hairs
c. 0.2 mm long or more...............20
Tomentum of leaf buds, twig apices and young inflorescences consisting of hairs
up to 0.1 mm long...................21
20. Twigs usually conspicuously hollow. Leaves on lower surface often with persistent
tomentum. Pedicel at base not articulated. Perianth generally persistent under the
fruit...........38. H. wallichii (in part)
Twigs not conspicuously hollow. Lower leaf surface early glabrescent. Pedicel at base
articulated. Perianth not persistent under the fruit...........3. H. borneensis
21. Twig slender, towards apex 1.52(4) mm diameter. Leaves 614(17) cm long; lateral
veins 811 pairs. Fruits c. 1.5 cm long; pericarp c. 1.5 mm thick.....................
.............24. H. penangiana
Twig stout, towards apex 2.54(10) mm diameter. Leaves 921 cm long; lateral
veins 1116 pairs. Fruits 4.58 cm long; pericarp 1020 mm thick...26. H. punctatifolia
22. Bark of twig drying to a pale colour, grey-brown or grey-yellowish, contrasting with
the dark brown or blackish colour of the drying petiole...................23
Bark of twig drying dark brown, not or slightly contrasting with the colour of drying
petiole...................27
23. Leaves membranous, upper surface drying brown or blackish brown, the lower surface
somewhat paler. Perianth persistent under the fruit........22. H. pallidicaula (in part)
Leaves usually chartaceous, upper surface drying bright brown or olivaceous, lower
surface usually brown or chocolate. Perianth not persistent under the fruit..24
24. Twig slender, towards apex 23 mm diameter. Leaves 716 cm long, lower surface
bright brown or chocolate, contrasting well with the grey-olivaceous upper
surface.......21. H. oligocarpa
Twig stout, towards apex 310 mm diameter. Leaves 1335 cm long, lower and upper
surfaces not conspicuously contrasting in colour...........................25
359
25. Fruits 5 cm long or more. Upper leaf surface not finely wrinkled on drying....
......7. H. discolor
Fruits 2 cm long or less. Upper leaf surface finely wrinkled on drying.........26
26. Leaves oblong to elliptic-oblong. Fruits 1.62 cm long..............5. H. carnosa
Leaves lanceolate to elliptic-oblong. Fruits up to 1.5 cm long................37. H. tristis
27. Tomentum of leaf buds and young inflorescences very short, consisting of hairs up
to 0.1 mm long. Leaf lateral veins on upper leaf surface flat or sunken, or slightly
raised................28
Tomentum of leaf buds and young inflorescences composed of hairs 0.1 mm long
or more. Leaf lateral veins raised or not above; if hairs only c. 0.1 mm long, then
the lateral veins, at least in the lower half, distinctly raised above...........30
28. Leaves membranous............19. H. obscura
Leaves membranous, chartaceous or coriaceous....................29
29. Leaves on lower surface without distinct large hair scars. Fruits 35 cm long.
...........34. H. subalpina
Leaves on lower surface usually with distinct enlarged hair scars. Fruits c. 5 cm
long..........39. H. xanthina
30. Pedicel at base not articulated (this character best seen in male flowers). Fruits with
persistent perianth or not...............................31
Pedicel at base articulated. Perianth not persistent in fruits.............35
31. Leaves 1545 cm long. Fruits c. 6 cm long or more; perianth persistent for a long
time....................32
Leaves 520 cm long. Fruits 24.2 cm long; perianth not persistent..................33
32. Lateral veins 1112 pairs............1. H. amplomontana
Lateral veins 2030 pairs.....................10. H. fragillima (in part)
33. Leaves membranous; apex acute-acuminate. Fruits 2.43 cm long...........2. H. androphora
Leaves chartaceous to thickly coriaceous; apex rounded to subacute. Fruits 24.2
cm long...........34
34. Leaves chartaceous to coriaceous; lower surface without large hair scars. Female flower
buds c. 2 mm long. Fruits 22.7 cm long...........................16. H. montana
Leaves usually thickly coriaceous, usually with large and distinct hair scars on lower
surface. Female flower buds 2.53 mm long. Fruits 34.2 cm long..........9. H. endertii
35. Midrib on upper leaf surface towards the transition to the petiole c. 3 mm broad
or more............14. H. laticostata
Midrib at the base narrow................36
36. Leaves 1628 cm long; base short-attenuate to rounded; lateral veins 1619
pairs.........18. H. nervosa
360
Leaves 528 cm long; base either rounded or short- or long-attenuate; lateral veins
518 pairs.....................37
37. Lateral veins on upper leaf surface sunken or flattish, or slightly raised........38
Lateral veins distinctly raised above...............39
38. Leaf apex rounded..................20. H. obtusa
Leaf apex acute-acuminate.....................28. H. ridleyana
39. Tomentum of leaf buds and young inflorescences comprising hairs 0.10.2(3) mm
long. Leaves drying dull greyish-brown, the colours of the upper and lower surfaces
not much contrasting. Fruits 1.72 cm long...............36. H. tenuifolia
Tomentum of leaf buds and young inflorescences consisting of hairs c. 0.2 mm long
or more; or if hairs c. 0.1 mm long, then the olivaceous to dark brown colour of the
dry leaf above much contrasting with the cinnamon colour beneath. Fruits 1.96 cm....40
40. Twig slender, towards apex 13 mm diameter. Leaves generally membranous, 10
18(27) cm long......................15. H. macilenta
Twig generally stouter, towards apex 25 mm diameter. Leaves chartaceous, of various
sizes.......................41
41. Twig early glabrescent; older bark more or less longitudinally cracked. Leaf apex
long acute-acuminate. Fruits 2.83.2 cm long; pericarp hard and woody, 810 mm
thick.................8. H. disticha
Twig usually rather late-glabrescent; older bark striate, not cracked. Leaf apex acuteacuminate, the acumen not conspicuously long. Fruits 1.96 cm long; pericarp not
so, 215 mm thick.................25. H. polyspherula
1. Horsfieldia amplomontana W.J. de Wilde

(Latin, amplum = large; resembling H. montana, but much larger)
Gard. Bull. Sing. 39 (1986) 34; Whitmore, Tantra & Sutisna l.c. 263. Type: Clemens 30536, Borneo,
Sabah (holotype L; isotypes BM, K).
Tree up to 20 m tall. Bark grey, fissured, soft, c. 5 mm thick; inner bark white, soft,
c. 5 mm thick; sap sticky. Sapwood white. Twigs solid, terete or faintly angular, not
ridged, towards apex 3.56(10) mm diameter, drying brown, not contrasting with the
drying petiole, glabrescent, tomentum comprising hairs 0.31(1.5) mm long; lenticels
inconspicuous; older bark not flaky. Leaf buds 1522 34 mm, tomentose with hairs
0.51(1.5) mm long. Leaves distichous, membranous to chartaceous, upper surface not
scabrous, glabrescent, drying olivaceous, lower surface glabrous, without blackish dots,
without pale hair scars; oblong, 1535 511 cm, base attenuate to rounded, apex acuteacuminate; midrib flattish to moderately raised above; lateral veins 1112 pairs, flattish
to raised above; petiole 0.81.5 cm long. Inflorescences pubescent with shaggy hairs
c. 0.5 mm long; males 3 or 4-times branched, many-flowered, 1021 1016 cm, peduncle
1540 mm long; females not seen, in fruit 78 cm long; bract 27 mm long, caducous.
Male flowers: pedicel not articulated, 0.82 mm long; buds slightly depressed globose,
361
1.52 22.2 mm, not or slightly collapsing on drying; perianth 34-lobed, lobes c.
0.2 mm thick, at anthesis clefts to 1/22/3; androecium (sub)sessile, depressed globose,
rounded or sagged at base, 0.61 1.11.8 mm, in transverse section circular; apical
excavation small, concealed by the apices of the anthers, anthers 1013, sessile, free
apices up to 0.1 mm long, curved and more or less entirely connate, concealing the
more or less 3-radiate, 0.20.5 mm deep cavity of the column; androphore narrow, 0.1
0.4 mm long, hidden by the anthers. Fruits 13 per infructescence, ellipsoid, 78 4.5
5 cm, glabrous, tubercled; pericarp c. 15 mm thick; stalk c. 3 mm long, at base not
articulated; perianth persistent. Seeds ellipsoid; aril red.
Distribution. Endemic to Borneo; confined to Mt. Kinabalu, Sabah.
Ecology. In primary and degraded hill to lower montane forests on sandstone, at altitude
of 10001500 m. Flowering in NovemberDecember and February; fruiting in November.
2. Horsfieldia androphora W.J. de Wilde

(Greek, andro = male, phorus = bearing; the well developed androphore of the androecium)
Gard. Bull. Sing. 39 (1986) 32; Whitmore, Tantra & Sutisna l.c. 263. Type: Nooteboom & Chai 1710,
Borneo, Sarawak, Kapit district, Bt. Goram (holotype L; isotypes K, SAR).
Tree up to 20 m tall. Bark chocolate to reddish brown, narrowly cracked or longitudinally

furrowed and cut into rectangular blocks; sap watery, more or less colourless (in flowering
tree) or blood red (in fruiting tree). Twigs solid, terete or only faintly angular, not ridged,
towards apex 24(5) mm diameter, rusty tomentose with hairs 0.30.6 mm long, lateglabrescent; bark drying brown, not contrasting with the drying petiole, not flaky; lenticels
not conspicuous. Leaf buds 812 23 mm, pubescent with hairs 0.30.6 mm long.
Leaves distichous, membranous, upper surface drying dark brown, not bullate, not
scabrous, glabrous, lower surface early glabrescent except the midrib, without blackish
dots and golden hair scars; elliptic to oblong, 918 3.56.5 cm, base attenuate, apex
acute-acuminate; midrib raised above, late-glabrescent beneath; lateral veins 913 pairs,
raised above; petiole 11.2 cm long. Inflorescences tomentose with hairs 0.20.6 mm
long; bract c. 3 mm long, caducous; males many-flowered, 3(4)-times branched, 614
3.59 cm, peduncle 1020 mm long; females 34 cm long. Male flowers: pedicel
0.52 mm long, slender, at base not articulated; buds (sub)globose, 1.42.2 mm diameter;
perianth glabrous, 3-lobed, at anthesis clefts to 1/3 to nearly 1/2, lobes c. 0.2 mm thick;
synandrium depressed globose, in transverse section circular, 0.61 0.81.3 mm;
androphore slender, 0.40.8 mm long; anthers 711, curved, sessile, and almost entirely
connate, concealing the 0.20.3 mm deep apical cavity of the column. Fruits 25 per
infructescence, ellipsoid, 2.43 1.42 cm, glabrous, finely tubercled; dry pericarp 1.5
2 mm thick; stalk c. 2 mm long; perianth not persistent. Seeds ellipsoid; outer seed
coat whitish grey; aril orange.
Vernacular name. Sarawakliang (Kelabit).
Ecology. In hill to lower montane mossy forests, wooded sandstone ridges, at altitudes
of 8001200 m. Flowering in March and October; fruiting in March and June.
362
3. Horsfieldia borneensis W.J. de Wilde

(of Borneo)
Gard. Bull. Sing. 39 (1986) 27, l.c. (1987) 468; Whitmore, Tantra & Sutisna l.c. 263. Type: Bojang S.
14610, Borneo, Sarawak (holotype L; isotypes K, SAR).
Tree up to 30 m tall. Bark rough, deeply fissured, flaking in squares, usually dark brown
or reddish or blackish, strips or flakes up to 5 cm wide, up to 1 cm thick; inner bark
510 mm thick, red-brown; sap red. Sapwood reddish white to pale red; heartwood redbrown. Twigs subterete, not conspicuously hollow, not ridged, towards apex (1.5)2
4(10) mm diameter, brown, not contrasting with the drying petiole, rusty tomentose with
hairs 0.20.4 mm long, glabrescent; lower down finely striate, not distinctly lenticellate,
when older finely cracked or flaky or not. Leaf buds 1017 24 mm, densely pubescent
with hairs c. 0.3 mm long. Leaves distichous, chartaceous to thinly coriaceous, upper
surface drying dull olivaceous to brown, not bullate, not scabrous, glabrous, lower surface
drying pale brown or chocolate, tomentose with shortish densely branched dendroid hairs
0.30.4 mm long (especially on the midrib), early glabrescent, without golden hair scars,
always with scattered blackish dots; elliptic-oblong to oblong-lanceolate, 718 26 cm,
base short-attenuate, apex acute to short-acuminate; midrib slightly raised above, early
or late-glabrescent; lateral veins 1016 pairs, flat or sunken (or close to the midrib slightly
raised) or in thinner-leaved specimens slightly raised above; intercostal venation hardly
visible on both surfaces; petiole 1.22.5 cm long, late-glabrescent. Inflorescences on older
twigs behind the leaves, woolly pubescent with rusty hairs up to 0.7 mm long; bract
1.55 mm, caducous; males fairly large, c. 4-times branched, (8)1320 (5)1018
cm, peduncle 1535 mm long; females stout, on older twigs c. 1.5 cm thick well below
the leaves, rather many-flowered, 810 45 cm, peduncle c. 15 mm long. Male flowers
in loose clusters of 26; pedicel 11.5 mm long, glabrous, articulated at base; buds
subglobose to broadly ellipsoid or obovoid, 1.31.8 1.21.7 mm, apex broadly rounded,
base narrowly rounded; perianth glabrous, 3-lobed, at anthesis clefts to c. 1/3 to nearly
1/2, not collapsing on drying; androecium subglobose, 0.71.2 0.61.3 mm, circular
to rounded-subtriangular in cross section, sessile, apex broadly rounded, slightly impressed
in the centre to 1/51/4 or less; anthers 710, almost completely sessile and entirely
connate, at apex incurved over the cavity of the column. Female flowers solitary or
24 together; pedicel rather thick, 11.5 mm long, at base articulated; buds subglobose
or broadly ellipsoid, c. 33.5 3 mm; ovary subglobose 2(2.5) 2 mm, glabrous.
Infructescences up to 13 cm long. Fruits 17 per infructescence, ovoid, somewhat laterally
flattened and slightly flanged, apex and base rounded, 46 34.5 cm, glabrous, drying
often with a glaucous tinge, rather smooth; dry pericarp 1015 mm thick; stalk stout,
46 mm long; perianth not persistent. Seeds ellipsoid; aril orange.
Distribution. Endemic to Borneo (Sabah, Sarawak, E and NE Kalimantan).
Ecology. In primary lowland mixed dipterocarp forest and swamp forest, on sandy clay
soils, up to 200 m altitude. Flowering in April and AugustSeptember; fruiting throughout
the year.
4. Horsfieldia brachiata (King) Warb.

(Latin, brachiatus = much-branched; the male inflorescence)
363
Monogr. Myrist. (1897) 325; Gamble l.c. 218; Ridley l.c. 59; Whitmore l.c. 325; J. Sinclair l.c. (1975) 8;
Anderson l.c. 263; W.J. de Wilde l.c. 39 (1986) 3; Turner l.c. 358; Coode et al. (eds.) l.c. 221; Argent et
al. (eds.) l.c. 447. Basionym: Myristica brachiata King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 311.
Lectotype: Griffith 4351, Malacca (hololectotype CAL; isolectotypes A, CGE, K, M, P, S, U). Synonyms:
H. subglobosa (Miq.) Warb. var. brachiata (King) J. Sinclair l.c. 16 (1958) 431; H. brachiata (King)
Warb. var. brachiata J. Sinclair l.c. (1975) 9.
Tree up to 35 m tall. Bark more or less smooth, pale brown to dark brown, shallowly
fissured, laminated or scaly, or cracked; inner bark 810 mm thick, pinkish to reddish
brown; sap reddish. Sapwood whitish to pale brown. Twigs more or less angular or
flattened, towards apex 27(18) mm diameter, mostly lined or ridged or nearly winged
between the petioles, at first rusty tomentose with hairs (0.1)0.20.4 mm long, generally
early glabrescent; bark lower down striate, (dark) brown, not contrasting with the drying
petiole, not flaky. Leaf buds 815 34 mm, pubescent with hairs 0.20.4 mm long.
Leaves distichous, membranous, upper surface glabrous, drying olivaceous to brown,
not bullate, not scabrous, lower surface drying light brown, early glabrescent but midrib
sometimes late-glabrescent, without dark dots; elliptic-oblong to oblong-lanceolate, 12
26(30) 49(11) cm, base cuneate, apex acute-acuminate; midrib raised above; lateral
veins 1220 pairs, raised above; intercostal venation not distinct; petiole 0.81.3(2) cm
long. Inflorescences sparsely to densely pubescent with hairs 0.20.5 mm long, sometimes
partly glabrescent; bract 35 mm long, pubescent, caducous; males 34-times branched,
718(22) 516(18) cm, peduncle 618 mm long, flowers in loose clusters of 3
6; females 38 26 cm. Flowers: perianth 3(4)-lobed, glabrous; pedicel pubescent
in various degree with hairs 0.10.2 mm long, or glabrescent, at base more or less
articulated. Male flowers: pedicel (1)1.52.5 mm long; buds subglobose or broadly
obovoid, in transverse section slightly angular or rounded, 11.5 1.21.8 mm, apex
broadly rounded, base rounded to short-cuneate; perianth at anthesis clefts to 1/22/3,
not or only slightly collapsing on drying; androecium depressed globose to obovoid,
(0.5) 0.71 0.81.2 mm, more or less 3-angular in transverse section; anthers 6
10, for c. halfway free, curved towards the centre, column deeply excavated; androphore
short. Female flowers: pedicel 11.5 mm long; buds broadly ellipsoid, 2.22.5 1.8
2 mm; ovary ovoid, 11.4 0.81.2 mm, glabrous. Fruits 412(20) per infructescence,
broadly ellipsoid, base broadly rounded, 22.8(4) 1.82.2(3) cm, apex narrowly
rounded, glabrous, drying brown, not warted nor lenticellate; dry pericarp 1.54(7) mm
thick; stalk 1.53 mm long; perianth not persistent. Seeds ellipsoid.
Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, and Borneo (Sabah,
Sarawak, Brunei, and Kalimantan).
Ecology. In primary and degraded lowland and hill rain forests on various types of welldrained and water-logged soils, up to 400 m altitude. Flowering and fruiting throughout
the year.
5. Horsfieldia carnosa Warb.
Fig. 3.
(Latin, carnosus = succulent; the leaves)

Monogr. Myrist. (1897) 348; Merrill l.c. (1921) 268; Masamune l.c. 300; J. Sinclair l.c. 28 (1975) 21;
Anderson l.c. 264; W.J. de Wilde l.c. (1985) 222; Whitmore, Tantra & Sutisna l.c. 263; Coode et al.
364
vi
365
(eds.) l.c. 221. Lectotype: Beccari 1242, Borneo, Sarawak (FI Acc. 7625, fruit). Synonym: Myristica
carnosa (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 87.
Tree up to 10 m tall. Bark often flaky or shallowly fissured; inner bark yellowish, thin;
sap watery, clear. Sapwood whitish. Twigs pale brown, terete, stout, neither distinctly
lined nor ridged, towards apex 310(16) mm diameter, initially tomentose with hairs
c. 0.1 mm long, glabrescent; older twigs striate, with a tendency to flake, drying pale,
contrasting with the dark brown drying petioles; lenticels not conspicuous. Leaf buds
913 23 mm, tomentose with hairs c. 0.1 mm long. Leaves distichous, chartaceous
to coriaceous, upper surface drying bright dark brown, bullate (with a finely wrinkledgranulate structure), lower surface brown, early glabrescent (glabrous), without blackish
dots; elliptic-oblong to oblong, 1335 511 cm, base attenuate; midrib flat above; lateral
veins 1318 pairs, flat above; petiole 11.6 cm long. Inflorescences tomentose with hairs
up to 0.1 mm long; bract 410 mm long, caducous; males many-flowered, 34-times
branched, 617 514 cm, peduncle 1030 mm long; females borne on older branches,
many-flowered, 12 cm long, flowers clustered. Flowers glabrous; perianth 3-lobed; pedicel
not articulated at the base. Male flowers in loose clusters of 39; pedicel 11.5 mm
long; buds subglobose, 1.92.1 1.82 mm; perianth at anthesis clefts to 1/31/2, lobes
c. 0.2 mm thick, not collapsing on drying, glabrous; androecium subglobose, 11.2
11.3 mm, circular in transverse section; anthers 911, largerly connate, completely sessile
(free apices more or less absent), curved, at apex concealing the 0.20.4 mm deep apical
cavity of the column; androphore narrow, 0.20.5 mm long, completely hidden by the
anthers. Female flowers: buds ellipsoid, c. 3.5 mm long. Fruits 211 per infructescence,
ellipsoid, 1.62 1.21.5 cm, glabrous, drying brown with a finely granular surface;
dry pericarp c. 1.5 mm thick; stalk 12 mm long; perianth not persistent. Seeds ellipsoid;
aril orange.
Distribution. Endemic to Borneo (Sabah, Sarawak, Brunei and W Kalimantan).
Ecology. In kerangas forest, peatswamp forest, and Agathis-Casuarina forest, on white
sandy soils, up to 100 m altitude. Flowering mainly in JulyNovember; fruiting throughout
the year.
6. Horsfieldia crassifolia (Hook. f. & Thomson) Warb.

(Latin, crassus = thick, folium = leaf; thick-leaved)
Monogr. Myrist. (1897) 323; Gamble l.c. 217; Merrill l.c. (1921) 268; Ridley l.c. 59; Masamune l.c.
301; J. Sinclair l.c. 16 (1958) 386, l.c. (1975) 23; Anderson l.c. 264; W.J. de Wilde l.c. (1985) 219;
Whitmore, Tantra & Sutisna l.c. 263; Turner l.c. 358; Coode et al. (eds.) l.c. 221; Argent et al. (eds.) l.c.
448. Basionym: Myristica crassifolia Hook. f. & Thomson, Fl. Ind. (1855) 160. Type: Griffith 4350,
Peninsular Malaysia (K). Synonyms: M. paludicola King l.c. 328; H. fulva (King) Warb. var. paludicola
(King) Warb. l.c. (1897) 299.
Tree up to 25 m tall; stilt roots or low buttresses occasionally occur. Bark greyish, fissured,
flaking into small rectangular scales. Twigs terete, grey-brown and not much contrasting
with the drying petioles, towards apex 26(8) mm diameter, yellow-brown to rusty woolly
tomentose with hairs 0.20.5 mm long, glabrescent; bark striate, not flaky, lenticels distinct
or not. Leaf buds 712 23 mm, pubescent with hairs 0.20.5 mm long. Leaves distichous,
366
coriaceous, upper surface drying dull (greenish) brown, lower surface thinly covered with
subpersistent tomentum consisting of interwoven or spaced stellate scales c. 0.1 mm high,
or glabrescent with distinct hair scars, with scattered blackish dots or streaks; elliptic
to oblong, 1020(28) 3.57(10) cm, base rounded to attenuate, apex rounded to subacute
or rarely emarginate; midrib flattish above; lateral veins 1116 pairs, flattish or sunken
above; intercostal venation indistinct or invisible on both surfaces; petiole 0.92(3) cm
long. Inflorescences late-glabrescent or with persistent, rather dense, woolly tomentum
consisting of dendroid hairs 0.20.5 mm long; bract 25(7) mm long, caducous; males
35-times branched, broadly paniculate, 620 415 cm, peduncle 420 mm long; females
314 cm long. Flowers in loose clusters of 27, glabrous; perianth 2-lobed; pedicel at
base not articulated. Male flowers: pedicel 0.31 mm long; buds globose or slightly
transversely ellipsoid, (0.8)11.3 11.5 mm; perianth at anthesis clefts to 1/31/
2 (2/3); androecium globose or more or less transversely ellipsoid, 0.40.5 0.5
0.8 mm; anthers (3)46, well-spaced, with broad connectives (giving the androecium
an angular appearance), free for almost the upper half or more; androphore 0.2(0.3)
mm long, slender. Female flowers: pedicel 1.52.5 mm long; buds 23 22.5 mm;
perianth at anthesis clefts for 1/51/3 only; ovary glabrous, 1.5 1.21.5 mm. Fruits
(1)210 per infructescence, ovoid to obovoid, 1.52.2 1.21.8 cm, base and apex rounded,
glabrous, sometimes with few lenticel-like tubercles; dry pericarp 1.52 mm thick; stalk
25(7) mm long; perianth persistent. Seeds ellipsoid.
Vernacular name. Sarawakkumpang paya (Iban).
Distribution. S Thailand, Sumatra, Peninsular Malaysia, Singapore, and Borneo (Sabah,
Ecology. Mostly in marshy habitats, such as freshwater and peatswamp forests, on sandy
soils, up to 200 m altitude. Flowering and fruiting throughout the year.
7. Horsfieldia discolor W.J. de Wilde

(Latin, discolor = of different colour; the two sides of the leaf blade)
Blumea 32 (1987) 469; Whitmore, Tantra & Sutisna l.c. 264. Type: Kostermans 7414, Borneo, E
Kalimantan (holotype L; isotype BO).
Tree up to 18 m tall. Bark red-brown, light brown or blackish, smooth or fissured; inner
bark red-brown or orange-brown, 57 mm thick; sap clear pink or red. Sapwood white
or yellow with reddish streaks. Twigs stout, terete, not ridged in between the bases of
the petioles, towards apex (2)34 mm diameter, tomentose with scale-like stellate hairs
c. 0.1 mm long, soon glabrescent; bark pale coloured, contrasting with the dark brown
drying petiole, lower down coarsely striate, not cracked or flaky; lenticels absent or few
or many, inconspicuous. Leaf buds slender, c. 10 2 mm, pubescent with scale-like hairs
up to 0.1 mm long. Leaves distichous, thinly chartaceous, extremely brittle when dry,
upper surface not scabrous, not finely wrinkled, drying olivaceous-brown, lower surface
early glabrescent (glabrous), drying bright brown, without blackish dots; oblong to oblonglanceolate, (9)1318 (3.5)47.5 cm, base attenuate; midrib flat, slender; lateral veins
1013 pairs, flat above, slender and inconspicuous beneath; petiole drying dark brown,
367
11.5 cm long. Male and female inflorescences and flowers not known, but judging
from the flower of related species, the perianth presumably subglobose, 34-lobed;
androecium subcircular in cross section. Infructescences coarse, short, little or not
branched, 1.53.5 cm long, early glabrescent, bearing 12(3) fruits. Fruits subglobose
to ellipsoid, 5.58(9) 46.5 cm, glabrous, drying blackish brown, wrinkled and without
or with a few wart-like lenticels; dry pericarp 1020 mm thick; stalk 25 mm long; perianth
not persistent. Seeds ellipsoid-oblong, 44.5 3 cm; aril bright orange to red, laciniate
at apex for 1/41/5.
Ecology. In hill mixed dipterocarp forest on loamy soils over sandstone, at 700900 m
altitude; also in alluvial (riverbank) forest and kerangas forest on sandy soils. Fruiting
in March, JuneJuly, and November.
8. Horsfieldia disticha W.J. de Wilde

(Latin, distichus = in two opposite rows; the leaves)
Gard. Bull. Sing. 39 (1986) 10, l.c. (1987) 467; Anderson l.c. 264; Whitmore, Tantra & Sutisna l.c. 264;
Coode et al. (eds.) l.c. 222. Type: Sinclair (& Kadim) 10453, Borneo, Brunei, Andulau FR (holotype L;
isotypes A, B, BM, E, K, NY, SAR, SING).
Tree up to 20 m tall. Bark with longitudinal, shallow furrows. Twigs solid, slender, terete
to faintly angular, not ridged, towards apex 23(4) mm diameter, on drying dark greybrown or dull reddish brown, not contrasting with the drying petiole, initially tomentose
with dull rusty hairs 0.20.3 mm long, early glabrescent; bark coarsely longitudinally
cracked, lenticels sparse but distinct. Leaf buds slender, c. 10 2 mm, pubescent with
hairs 0.20.3 mm long. Leaves distichous, chartaceous, upper surface drying olivaceous,
not bullate, not scabrous, glabrous, lower surface pale chocolate, glabrous, without blackish
dots; oblong-lanceolate, 813.5 2.53.5 cm, base more or less rounded to short-attenuate,
apex long acute-acuminate; midrib raised above, at base narrower than 3 mm; lateral
veins 913 pairs, raised above; petiole 11.5 cm long. Male and female flowers not
seen but perianth 3-lobed as judged from the perianth scars on the fruits. Infructscences
36 24 cm, borne on older branches behind the leaves, bearing 36 fruits. Fruits
ovoid-ellipsoid, 2.83.2 2.12.5 cm, glabrous, drying dark brown, not tubercled or
lenticellate; pericarp woody when dry, 810 mm thick; stalk 1015 mm long; perianth
not persistent. Seeds ellipsoid.
Distribution. Endemic to Borneo (Brunei; can be expected to occur in Sabah and/or
Sarawak).
Ecology. In lowland mixed dipterocarp forest. Fruiting in August.
9. Horsfieldia endertii W.J. de Wilde

(F.H. Endert, 18911953, sometime forest officer in Indonesia)
Gard. Bull. Sing. 39 (1986) 24; Whitmore, Tantra & Sutisna l.c. 264. Type: Endert 3996, Borneo, E
Kalimantan, W Kutai (holotype L; isotypes A, BO, K).
368
Tree up to 40 m tall. Bark cracked to finely fissured, brown-black to dark brown; inner
bark reddish. Sapwood whitish. Twigs solid, terete, not ridged, towards apex 2.5
4(8) mm diameter, drying dark brown, not contrasting with the drying petiole, at
first pubescent with rough, deep-rusty hairs 0.30.6 mm long, glabrescent; bark coarsely
striate, with conspicuous lenticels, older bark tending or not to flake. Leaf buds 10
20 34 mm, pubescent with hairs 0.30.6 mm long. Leaves distichous, thickly coriaceous,
brittle on drying, upper surface drying olivaceous or brown, not scabrous, lower surface
drying brown, not much contrasting with the upper surface, without blackish dots or streaks
but usually with conspicuous golden hair scars, glabrous; elliptic to lanceolate, 8
17(26) 36(10) cm, base rounded to short-attenuate, apex rounded to subacute;
midrib broad and raised above, towards the base late-glabrescent on both sides; lateral
veins 815 pairs, sunken or slightly raised above; intercostal venation generally invisible
on both surfaces; petiole 0.61.6 cm long. Inflorescences rusty pubescent with shaggy
hairs 0.51 mm long; bract 37 mm long, pubescent, caducous; males 2- or 3-times
branched, few-flowered, (1.5)310 (1)25 cm, peduncle 520 mm long, flowers in
clusters of (1)26; females few-flowered, 24 cm long. Flowers: perianth glabrous, 3
4-lobed; pedicel puberulous with minute hairs 0.10.3 mm long, especially in the lower
half, at base not articulated. Male flowers: pedicel 23 mm long; buds obovoid to ellipsoid,
2.53.5 22.5 mm, apex and base rounded; perianth at anthesis clefts to 1/31/2, slightly
collapsing on drying; androecium more or less sessile, truncate-ellipsoid, 22.8 1.4
1.6 mm, subtriangular in cross section; anthers (10)1214, sessile, 22.8 mm long, almost
entirely connate, more or less curved; column solid except for the narrow apical cavity
of 1/51/3 deep; androphore narrow, 0.10.3 mm long, hidden by the anthers. Female
flowers glabrous; buds 2.53 22.5 mm; perianth at anthesis clefts to 1/31/2; ovary
c. 2 1.5 mm, glabrous; pedicel 1.52 mm long, towards the base pubescent with hairs
c. 0.2 mm long. Fruits 14(8) per infructescence, ellipsoid, 34.2 1.62.4 cm, base
rounded, apex rounded to somewhat acute, glabrous; dry pericarp 24 mm thick; stalk
35 mm long; perianth not persistent. Seeds ellipsoid; aril bright orange.
Vernacular name. Sarawakbinarak (Murut).
Ecology. In lower to upper montane forests on ridges and wind-swept crests, on sandy,
black or brownish soils, at 12002100 m altitude. Flowering and fruiting throughout the
year.
10. Horsfieldia fragillima Airy Shaw
Fig. 4.
(Latin, fragilis = fragile; the dry leaves)

Kew Bull. 1939, no. 10 (1940) 542; Masamune l.c. 301; J. Sinclair l.c. (1975) 31; Anderson l.c. 264;
W.J. de Wilde l.c. (1986) 29; Whitmore, Tantra & Sutisna l.c. 264; Coode et al. (eds.) l.c. 222; Argent et
al. (eds.) l.c. 448. Type: Richards 2602, Borneo, Sarawak, Marudi district, Mt. Dulit (holotype K;
isotypes A, SING).
Tree up to 30 m tall. Bark chocolate, reddish brown, or blackish brown, slightly fissured,
mostly with few flakes, or scaly, or peeling off in thin flakes; inner bark c. 3 mm thick,
pale pink or reddish to yellowish. Sapwood soft, whitish, yellow-pink or pinkish white.
369
vi
Twigs solid, terete or subterete, not ridged, drying brown, not contrasting with the drying
petiole, towards apex 2.57(15) mm diameter, tomentose with hairs c. 0.3 mm long,
early glabrescent; bark with conspicuous lenticels, with a slight tendency to slightly flaking.
Leaf buds 1220 35 mm, pubescent with hairs c. 0.2 mm long. Leaves distichous
or sometimes partly in 3 rows, not bunched at the top of the twigs; membranous to
coriaceous, upper surface drying olivaceous or brown, not bullate, not scabrous, lower
surface early glabrescent, often with some minute tomentum remaining at the base of
the midrib, without blackish dots, without golden hair scars; oblong to oblong-lanceolate,
2045 6.512(15.5) cm, base rounded to attenuate; midrib raised above; lateral veins
2030 pairs, raised above; petiole 0.41.3(2) cm long. Inflorescences tomentose with
hairs 0.2 mm long or more; bract up to 12 mm, caducous; males large, to 5-times branched,
1530 1020 cm, peduncle 2560 mm long; females 613 cm long. Flowers glabrous;
perianth 34-lobed. Male flowers: pedicel 12 mm long, at base not articulated; buds
depressed globose, 1.42 22.5 mm; perianth at anthesis clefts to nearly 1/2; androecium
sessile, almost saucer-shaped, broadly depressed in the centre, 0.51 11.5 mm; anthers
79, sessile, more or less curved, almost entirely connate; column with broad apical cavity
to 1/2 deep; androphore minute. Female flowers: buds 45 mm long. Fruits up to 8
per infructescence, broadly ellipsoid, 68 46 cm, glabrous; dry pericarp 1020 mm
thick; perianth persistent for a long time; stalk 24 mm long. Seeds ellipsoid; aril reddish
orange.
Ecology. In primary lowland mixed dipterocarp forest, on hill slopes and flat seasonally
flooded lands, up to 400 m altitude, most commonly recorded from sandy, leached clayey
loam over sandstone, and sandy clay soils, often along or near streams. Flowering in
FebruaryMay; fruiting throughout the year.
Uses. Fruits recorded as edible, very acid and resinous.
11. Horsfieldia gracilis W.J. de Wilde

(Latin, gracilis = slender; the twigs)
Gard. Bull. Sing. 38 (1985) 211, Blumea 41 (1996) 377; Whitmore, Tantra & Sutisna l.c. 264; Coode
et al. (eds.) l.c. 222. Type: Ilias S. 16604, Borneo, Sarawak, Miri district (holotype L; isotypes K,
SAR).
Tree c. 5 m tall. Twigs terete, slender, towards apex 1.52.5 mm diameter, initially tomentose
with rough hairs 0.30.5 mm long, late-glabrescent; bark not cracked or flaky; lenticels
inconspicuous or absent. Leaf buds 67 1.5 mm, pubescent with persistent rough hairs
0.30.5 mm long. Leaves distichous, membranous, upper surface drying olivaceous,
glabrous, not bullate, not scabrous, lower surface tomentose with subpersistent scattered
rough hairs 0.3(0.5) mm long or subglabrous, without blackish dots or streaks; oblonglanceolate, 1221 46(6.5) cm, base subattenuate; midrib raised above, with persistent
tomentum beneath; lateral veins 1217 pairs, slightly raised to flat above; intercostal
venation indistinct above; petiole 0.81.4 cm long, late-glabrescent. Inflorescences: males
paniculate, 35 cm long; females 11.5 cm long. Male flowers: pedicel c. 0.5 mm long,
371
articulated at the base; buds depressed globose, c. 0.6 1 mm; perianth glabrous, 3
4-lobed, at anthesis clefts to slightly over halfway; androecium depressed globose, c.
0.5 0.91 mm; anthers 1618, central column with small cavity at the apex. Female
flowers: buds c. 2 1.5 mm. Fruits 13 per infructescence, ellipsoid, 1.41.5 1
1.1 cm, glabrous, drying dark brown, finely granulate; dry pericarp 0.51 mm thick; stalk
11.5 mm long; perianth persistent. Seeds ellipsoid.
Distribution. Endemic to Borneo (Sarawak, Miri district; also in Brunei).
Ecology. In primary lowland mixed dipterocarp forest.
12. Horsfieldia grandis (Hook. f.) Warb.

(Latin, grandis = large; the large leaves)
301; J. Sinclair l.c. (1958) 400, l.c. (1975) 48; Anderson l.c. 264; W.J. de Wilde l.c. (1985) 203; Whitmore,
Tantra & Sutisna l.c. 264; Turner l.c. 358; Coode et al. (eds.) l.c. 222; Argent et al. (eds.) l.c. 448.
Basionym: Myristica grandis Hook. f., Trans. Linn. Soc. 23 (1860) 157. Type: Low s.n., Borneo, Sabah
(holotype K). Synonym: M. rubiginosa King l.c. 302.
Tree up to 25 m tall. Bark smooth, fissured, scaly, flaky, or cracked, strips c. 1 mm

thick, c. 1 cm wide, hard and thin; inner bark 23 mm thick, slash rich red-brown; sap
reddish, watery. Sapwood soft, whitish to yellowish pink, heartwood pale brown. Twigs
rather stout, terete, towards apex (3)410 mm diameter, initially densely tomentose with
rough hairs 11.5 mm long, late-glabrescent, not ridged; bark sometimes cracked and
flaky; lenticels not conspicuous. Leaf buds 715 mm long, pubescent. Leaves distichous,
membranous or chartaceous, surface somewhat bullate, upper surface drying olivaceous
or brown, largely with rough persistent tomentum, sometimes glabrescent, always scabrous
by rough persistent hair bases, lower surface rough tomentose with persistent hairs 1
2 mm long, without dark dots; elliptic-oblong to oblong-oblanceolate, 1240 520 cm,
base attenuate or subcordate; midrib raised above; lateral veins (8)1016(19) pairs,
usually sunken above; petiole 0.61.5 cm long, pubescent. Inflorescences pubescent with
hairs 1.53 mm long; bract (1)312 mm long, caducous; males 625 2.510(15)
cm, peduncle up to 12 mm long; females 1.55 cm long. Flowers: perianth 34-lobed,
glabrous; pedicel glabrous, at base not articulated. Male flowers: pedicel (0.5)12 mm
long; buds depressed globose, 1.21.8(2) mm diameter; perianth at anthesis clefts to
1/31/2; androecium depressed globose, 0.51 0.81.5 mm; anthers 810, almost
completely sessile, incurved towards the apex; column with cavity c. 0.2 mm deep;
androphore narrow, 0.20.4 mm long. Female flowers: buds broadly ellipsoid, 23.2 mm
long. Fruits 210 per infructescence, more or less clustered, obovoid-ellipsoid, 11.4
0.81.1 cm, glabrous; dry pericarp c. 1.5 mm thick; stalk c. 1 mm long; perianth
persistent. Seeds almost globose, 0.81 cm diameter; aril orange.
Vernacular name. Sabahdarah-darah (Malay).
Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo (Sabah, Sarawak,
Brunei, and Kalimantan).
372
Ecology. Under- or middle-storey trees, rarely reaching timber size, of primary and
secondary lowland to hill forests on sandy and clay soils, up to 600 m altitude. Flowering
13. Horsfieldia irya (Gaertn.) Warb.

(irya, a local plant name from Sri Lanka)
301; J. Sinclair l.c. (1958) 382, l.c. (1975) 61; Backer & Bakhuizen f. l.c. 138; Whitmore, l.c. 326;
Anderson l.c. 264; W.J. de Wilde l.c. (1985) 55; Corner l.c. 564; Whitmore, Tantra & Sutisna l.c. 264;
Turner l.c. 358; Coode et al. (eds.) l.c. 222; Argent et al. (eds.) l.c. 448. Basionym: Myristica irya
Gaertn., Fruct. 1 (1788) 195. Type: Gaertners drawing, plant from Ceylon. Synonyms: M. javanica
Blume, Bijdr. (1825) 576; M. spherocarpa Wall., Pl. As. Rar. (1830) 79; M. irya (Gaertn.) Warb. var.
wallichii King l.c. 309; H. lemanniana (A. DC.) Warb. l.c. (1897) 326; H. subglobosa (Miq.) Warb. l.c.
(1897) 328; M. vrieseana Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 49; M. irya (Gaertn.) Warb. var.
longifolia King l.c. 309; H. labillardieri Warb. l.c. (1897) 283; H. acuminata Merr., Phil. J. Sc. 17, 1920
(1921) 253; H. nunu Kanehira, Bot. Mag. Tokyo 46 (1932) 451; H. amklaal Kanehira, Bot. Mag. Tokyo
47 (1933) 670; H. congestiflora A.C. Smith, J. Arn. Arb. 22 (1941) 64.
Tree up to 40 m tall; bole often fluted, with prop-roots or buttresses up to 3 m high,

2 m out, 10 cm thick, or without buttresses. Bark smooth, fissured or cracked, flaky
or peeling off in small pieces; inner bark cream or whitish, up to 7 mm thick. Sapwood
cream or whitish, ochreish, straw-coloured, or pinkish; heartwood absent or only slightly
darker, pale brown. Twigs terete, usually thinly ridged, towards apex (2)310 mm diameter,
initially tomentose with minute, grey or rusty hairs 0.10.5 mm long, glabrescent; bark
often blackish, not flaky; lenticels usually conspicuous. Leaf buds 10(15) mm long,
pubescent with hairs 0.10.5(1) mm long. Leaves distichous, often curved towards the
tip, membranous, upper surface drying greenish brown to blackish, usually with irregular
whitish blotches of unknown origin, lower surface early glabrescent, without dark dots;
elliptic-oblong to lanceolate, 1030(35) 37(9) cm, base rounded or attenuate; midrib
flattish above; lateral veins 1020 pairs, flattish above; petiole 0.71.6 cm long. Inflorescences
pubescent with hairs 0.10.5(1) mm long, persistent or glabrescent; bract 1.54 mm
long, caducous; males 418 37(10) cm, peduncle 545 mm long; females 26(8)
cm long. Flowers largely glabrous; perianth 2-lobed; pedicel at base not articulated.
Male flowers: pedicel 0.11.5 mm long; buds subglobose, laterally somewhat compressed
or not, 11.3(1.5) (1)1.21.5 mm; perianth at anthesis clefts to 1/22/3; androecium
transversely broadened, 0.81.2 11.5 mm; anthers 69, for the greater part connate,
forming a saucer-shaped column, at apex free for 0.20.3 mm; androphore cup-shaped,
tapered. Female flowers: buds ellipsoid, 1.52.3 mm long. Fruits 28 per infructescence,
globose, 1.52.2 cm diameter, glabrous, drying dark brown or blackish; dry pericarp 1
2 mm thick; stalk 510 mm long; perianth not persistent. Seeds globose; aril orange,
orange-red or red.
Vernacular names. Sabahlempoyang paya (preferred name). Sarawakpenarahan
pianggu (Malay).
373
Distribution. Widespread from Sri Lanka through Malesia to the Solomon Islands.
Throughout Borneo.
Ecology. In primary and old secondary forests. Most frequent in coastal or riverine habitats
on alluvial, sandy, loamy, or clayey soils, up to 450 m altitude.
14. Horsfieldia laticostata (J. Sinclair) W.J. de Wilde

(Latin, latus = broad, costa = midrib; the broad midrib)
Gard. Bull. Sing. 39 (1986) 15; Whitmore, Tantra & Sutisna l.c. 264. Basionym: H. brachiata (King)
Warb. var. laticostata J. Sinclair l.c. (1975) 12; Anderson l.c. 263. Type: Sinclair 10265, Borneo, Sarawak,
Serian district, Sg. Sabal Tapang (holotype K; isotypes A, E, L, SAR, SING).
Tree up to 35 m tall. Bark hard, fissured or flaky, brown to reddish brown; inner bark
reddish, laminated. Sapwood whitish. Twigs stout, solid, not ridged, terete, towards apex
38(15) mm diameter, grey-brown to blackish on drying, not contrasting with the drying
petiole, tomentose with hairs 0.10.4 mm long, early glabrescent; lower down coarsely
striate, almost flaky, lenticels usually distinct. Leaf buds 1523 35 mm, tomentose
with hairs 0.10.4 mm long. Leaves distichous, thinly to thickly coriaceous, upper surface
drying dull olivaceous, glabrous, not bullate, not scabrous, lower surface chocolate,
contrasting with the upper surface, without blackish dots, early glabrescent; obovateoblong to oblong, (15)2033 612 cm, base attenuate, apex shortly acute-acuminate;
midrib raised above, towards the base flattish and conspicuously broadened to 3
4(5) mm wide at the transition to the petiole, glabrous or with remnants of tomentum
towards the base; lateral veins 1120(24) pairs, raised above but towards the blade
margin sunken; intercostal venation distinct or not; petiole stout, 0.61.5 cm long, 0.15
0.4(0.8) cm diameter. Inflorescences sparsely pubescent with hairs 0.20.4 mm long;
bract 28 mm, caducous; males stout, 45-times branched, many-flowered, 1025 8
22 cm, axis towards the base 34 mm diameter, peduncle 335 mm long; females 3
7 cm long, few-flowered. Flowers in clusters of up to 10; perianth 3-lobed, glabrous;
pedicel sometimes with few hairs at the base, 0.10.2 mm long, distinctly articulated.
Male flowers: pedicel (1)1.5(2) mm long; buds broadly obovoid, 1.31.5 mm diameter;
base rounded or sometimes more or less attenuate; perianth at anthesis clefts to c. halfway,
not collapsing on drying; androecium broadly obovoid, c. 0.7 mm diameter, triangular
in transverse section; anthers 6, suberect, acutish, mutually largely free, c. 0.3(0.4) mm
long; androphore more or less tapering, c. 0.3(0.4) mm long. Female flowers glabrous
or with few minute hairs c. 0.1 mm long; buds broadly ellipsoid, c. 2.5 mm diameter;
perianth at anthesis clefts to c. 1/3, lobes 0.30.4 mm thick; ovary c. 1.5 mm long, glabrous;
pedicel 0.51 mm long. Fruits 14 per infructescence, ellipsoid, 2.84 2.22.5 cm,
apex and base rounded, glabrous; dry pericarp 45 mm thick; stalk 24 mm long; perianth
not persistent. Seeds ellipsoid, shining white, spotted.
Vernacular name. Sabahpisau-pisau (Kedayan).
Ecology. In primary peatswamp and kerangas forests, on sandy acid, nutrient-poor,
waterlogged soils, up to 400 m altitude. Flowering and fruiting throughout the year.
374
15. Horsfieldia macilenta W.J. de Wilde

(Latin, macilentus = meagre; the delicate twigs)
Gard. Bull. Sing. 39 (1986) 13; Whitmore, Tantra & Sutisna l.c. 264; Turner l.c. 358. Type: Aban SAN
37103, Borneo, Sabah (holotype L; isotype SAN).
Tree up to 17 m tall. Bark whitish or greenish yellow; inner bark red; sap red. Twigs
slender, terete, not ridged, drying brown, not contrasting with the drying petiole, towards
apex 13 mm diameter, tomentose with woolly stellate-dendroid hairs (0.2)0.5 mm long,
late-glabrescent; older bark not cracked; lenticels inconspicuous. Leaf buds 68 1.5
mm, tomentose with woolly stellate-dendroid hairs (0.2)0.5 mm long. Leaves distichous,
membranous, upper surface drying dull olivaceous, not bullate, not scabrous, glabrous,
lower surface early glabrescent except the midrib and the very base, without dark dots
and streaks; oblong to oblong-lanceolate, 1018(27) 36.5 cm, base rounded to shortattenuate, apex acute-acuminate; midrib raised above, at base narrow, on lower surface
late-glabrescent; lateral veins 1015(18) pairs, distinctly raised above; petiole 11.5
cm long, 0.10.15(0.2) cm diameter, tomentose with woolly stellate-dendroid hairs
(0.2) 0.5 mm long, late-glabrescent. Inflorescences sparsely woolly pubescent with
stellate-dendroid hairs c. (0.2)0.5 mm long; bract 25 mm long, caducous; males manyflowered, 2- or 3(4)-times branched, 49(10) 35(7) cm, peduncle 312 mm long;
females similar, or shorter. Flowers glabrous or sometimes with a few minute hairs less
than 0.1 mm long towards the base of the pedicel; perianth 3-lobed; pedicel glabrous
or sparsely pubescent in the lower portion with hairs up to 0.1 mm long, articulated
at the base. Male flowers in clusters of 58; pedicel (0.8)1(1.3) mm long; buds depressed
globose, 0.71.2 11.2 mm; perianth at anthesis clefts to nearly 1/2, lobes 0.20.3
mm thick; androecium subglobose, 0.40.5 0.50.7 mm, in transverse section more
or less triangular; anthers 57, suberect, 0.30.4 mm long, the upper half mutually free;
column largely hollow; androphore tapering, 0.20.3 mm long. Female flowers: buds
subglobose, c. 1.5 mm diameter. Fruits 2 or 3 together in a short infructescence 23
cm long, glabrous, ellipsoid, 2.22.4 1.51.6 cm, base contracted into a narrowed portion;
dry pericarp c. 2 mm thick, drying brown, not or sparingly tubercled; stalk c. 5 mm
long; perianth not persistent. Seeds ellipsoid.
Distribution. Sumatra, Peninsular Malaysia and Borneo (Sabah and Sarawak).
Ecology. In primary lowland mixed dipterocarp forest and swamp forest, up to 200 m
altitude. Flowering in JuneNovember.
16. Horsfieldia montana Airy Shaw

(Latin, montanus = growing in the mountain)
Kew Bull. 1939, no. 10 (1940) 542; Masamune l.c. 301; Anderson l.c. 265; W.J. de Wilde l.c. (1986) 35;
Whitmore, Tantra & Sutisna l.c. 265; Coode et al. (eds.) l.c. 222. Type: Richards (Native Collector)
2509, Borneo, Sarawak, Belaga district, Dulit Range (holotype K; isotype SING).
Tree up to 24 m tall. Bark finely and shallowly fissured, or sometimes flaky, chocolate,
red-brown, dark brown or dark grey; inner bark pale yellowish, reddish, or brownish;
sap red, watery. Sapwood pale orange or whitish. Twigs solid, terete or faintly angular,
375
towards apex 1.54(7) mm diameter, drying grey-brown to dark brown, not contrasting
with the drying petiole, tomentose with shaggy hairs 0.21 mm long, glabrescent; lenticels
inconspicuous or absent; bark flaky or not. Leaf buds 510 mm long, tomentose with
hairs 0.21 mm long. Leaves distichous, chartaceous to coriaceous, upper surface glabrous
except the midrib which is late-glabrescent towards the base, drying blackish, not bullate,
not scabrous, lower surface glabrous, drying chocolate-brown, not much contrasting with
the upper surface, without blackish dots or conspicuous golden hair scars; elliptic to
elliptic-oblong, 414 26 cm, base short-attenuate, apex rounded to subacute; midrib
raised above; lateral veins 611 pairs, flattish to raised above; petiole 0.51.6 cm long,
late-glabrescents. Inflorescences on older branches behind the leaves, either sparsely
pubescent with hairs 0.20.4 mm long, or densely pubescent with shaggy hairs 0.51
mm long; bract 2.56 mm long, caducous; males 412(16) 310 cm, peduncle 5
20 mm long; females 26 cm long. Flowers glabrous; perianth 34-lobed; pedicel in
males not articulated at the base, in females more or less distinctly articulated. Male
flowers: pedicel 11.5 mm long; buds subglobose, (1.2)1.42 mm diameter; perianth
at anthesis clefts to 1/3 to nearly 1/2, not or slightly collapsing on drying, lobes
(0.2) 0.3 mm thick; androecium (sub)sessile, depressed globose, (0.5)0.61.1 0.8
1.1 mm; anthers (8)911, more or less curved, almost entirely connate, free apices up
to 0.1 mm long; column with shallow apical cavity; androphore narrow, 0.30.5 mm
long, largerly hidden by the anthers. Female flowers: buds ellipsoid-obovoid, c. 2 mm
long; ovary glabrous. Fruits 29 per infructescence, ellipsoid, 22.7 1.31.7 cm, base
more or less contracted toward the stalk, glabrous, drying brown; pericarp 1.52 mm thick;
stalk 24 mm long, at base more or less articulated; perianth not persistent. Seeds ellipsoid.
Ecology. In crest forest, kerangas forest on ridges, montane mossy forest, and Agathis
forest, on black soils or sandy soils, at (800)13002000 m altitude. Flowering and fruiting
17. Horsfieldia motleyi Warb.

(James D. Motley, ?1859, engineer and plant collector, mainly in Borneo)
Monogr. Myrist. (1897) 304; Merrill l.c. (1921) 268; Masamune l.c. 301; J. Sinclair l.c. (1975) 81;
Anderson l.c. 265; W.J. de Wilde l.c. (1985) 209; Whitmore, Tantra & Sutisna l.c. 265; Argent et al.
(eds.) l.c. 452. Type: Motley 355, Borneo, Kalimantan, Banjarmasin (holotype K; isotype CGE).
Synonyms: Myristica motleyi (Warb.) Boerl. l.c. 85; H. macrobotrys Merr. l.c. (1929) 76.
Tree up to 25 m tall; bole sometimes with steep buttresses up to 5 m high, c. 30 cm

out, c. 10 cm thick. Bark usually distinctly fissured, often scaly, strips 34 cm wide,
up to 10 cm thick, dark brown, chocolate, or red-brown; inner bark 1012 mm thick,
undulate in cross section, dark brown or brown-red, laminated; sap red, watery or sometimes
sticky, appearing fast. Sapwood whitish, reddish white; heartwood reddish or pinkish
or brown. Twigs terete, towards apex (2.5)35(10) mm diameter; at first rusty tomentose
with hairs 0.5(1) mm long, late-glabrescent; lenticels not conspicuous; older twigs not
flaky. Leaf buds 1520 mm long, pubescent with hairs c. 0.5 mm long. Leaves distichous,
membranous, surface not scabrous, upper surface drying olivaceous to brown, glabrous,
376
not bullate, lower surface rather densely tomentose with persistent hairs 0.31 mm long,
without blackish dots or streaks; elliptic to oblong, 927 412 cm, base rounded to
attenuate, apex somewhat obtuse to acute-acuminate; midrib flattish above, sometimes
late-glabrescent; lateral veins 921 pairs, above late-glabrescent, usually sunken; petiole
1.32.2 cm long, late-glabrescent. Inflorescences yellow-brown to rusty pubescent with
hairs 0.51 mm long; bract 25 mm long, caducous; males 1220 1014 cm, peduncle
2040 mm long; females 36(10) cm long. Flowers: perianth 3-lobed; pedicel at base
not articulated; persistent pubescence comprising hairs 0.10.3 mm long. Male flowers:
pedicel 11.5 mm long; buds subglobose or broadly obovoid, 0.81 0.71 mm; perianth
at anthesis clefts to about halfway; androecium (including androphore) depressed obovoid,
0.50.7 (0.3)0.50.6 mm; anthers 5, almost sessile, 0.20.4 mm long, column with
apical cavity to c. 1/10 deep; androphore conspicuous, about as long as the anthers. Female
flowers pubescent; buds c. 2.4 1.82 mm; ovary glabrous. Fruits 515 per infructescence,
broadly ellipsoid, 1.92.3 1.61.8 cm, glabrous; dry pericarp c. 3 mm thick; stalk 2
4 mm long; perianth not persistent. Seeds ellipsoid.
Ecology. In various types of primary and disturbed mixed dipterocarp forests on welldrained sandy or sandy clay soils, often on ridge tops, up to 750 m altitude. Flowering
18. Horsfieldia nervosa W.J. de Wilde

(Latin, nervosus = nerved; the conspicuously many-nerved leaves)
Gard. Bull. Sing. 39 (1986) 16; Whitmore, Tantra & Sutisna l.c. 265. Type: Ilias & Ashton S. 16652,
Borneo, Sarawak (holotype L; isotypes A, K, SAN, SAR).
Tree up to 16 m tall; buttresses thin, up to 35 cm tall. Bark pale ochre and brown
mottled, smooth. Twigs solid, terete or towards apex subterete, not ridged, drying greybrown, not contrasting with the drying petiole, 3.56(8) mm diameter, initially tomentose
with hairs 0.30.6 mm long, glabrescent; lenticels inconspicuous; bark flaky or not. Leaf
buds 1620 mm long, tomentose with hairs c. 0.5 mm long. Leaves distichous, thinly
coriaceous, upper surface not bullate, not scabrous, drying olivaceous, midrib and lateral
veins late-glabrescent, lower surface chocolate, contrasting in colour with the upper surface,
without blackish dots, early glabrescent with the hairs c. 0.5 mm long often remaining
on veins and midrib; oblong, 1628 5.59 cm, base short-attenuate to rounded; midrib
slender, at transition with petiole c. 1.5 mm wide, distinctly raised above, late-glabrescent;
lateral veins 1619 pairs, conspicuously raised above, glabrescent; intercostal venation
laxly reticulate, indistinct or hardly visible; petiole 1.52.2 cm long, 0.30.5 cm diameter,
late-glabrescent. Inflorescences on the older branches behind the leaves, pubescent with
shaggy hairs up to 0.6 mm long; bract caducous; males c. 10 10 cm, peduncle 10
20 mm long; females c. 4 cm long. Flowers: perianth 3-lobed, glabrous; pedicel towards
the base pubescent with hairs 0.10.2 mm long, articulated. Male flowers: pedicel c.
1.5 mm long; buds globose, c. 1.2 mm diameter; perianth at anthesis clefts to nearly
1/2, not collapsing on drying; androecium (including androphore) more or less broadly
377
obovoid, 0.60.7 0.6 mm, triquetrous in transverse section; anthers 5 or 6, at least

the upper half free, more or less erect; column with deep cavity at the apex; androphore
more or less tapering to the base, c. 0.3 mm long. Female flowers: buds subglobose,
c. 2.5 mm diameter; perianth subglabrous with some scattered hairs c. 0.1 mm long toward
the base, at anthesis clefts to slightly over 1/3; ovary broadly ovoid, c. 1.5 mm diameter,
glabrous, stigma c. 0.3 0.6 mm, 2-lobed. Fruits not seen.
Distribution. Endemic to Borneo (Sarawak, known by two collections only).
Ecology. In primary mixed dipterocarp forest on yellow podsolic soils, up to 100 m altitude.
Flowering in November.
19. Horsfieldia obscura W.J. de Wilde

(Latin, obscurus = indistinct; a species difficult to recognize)
Gard. Bull. Sing. 39 (1986) 42, Blumea 32 (1987) 468; Whitmore, Tantra & Sutisna l.c. 265. Type:
Kostermans 13773, Borneo, E Kalimantan (holotype L; isotypes A, BO, K, NY, P, PNH, SING).
Tree up to 20 m tall. Bark red-brown, rough, c. 5 mm thick, peeling off irregularly

in strips; inner bark c. 10 mm thick, red to brown-red; sap red. Sapwood reddish to
brown-red. Twigs solid, terete, not ridged, drying dark brown, not contrasting with the
drying petiole, towards apex 23(4) mm diameter, initially tomentose with hairs up to
0.1 mm long, early glabrescent; bark not flaky; lenticels moderately large. Leaf buds slender,
710 mm long, pubescent with hairs up to 0.1 mm long. Leaves distichous, membranous,
not bullate, not scabrous, upper surface drying dark brown, lower surface pale brown,
glabrous, without blackish dots; elliptic-oblong to oblong, 1015 47 cm, base shortattenuate, apex shortly acute-acuminate; midrib flat to slightly raised above; lateral veins
1013 pairs, very thin and flat above; intercostal venation laxly reticulate, very faint
on both surfaces; petiole 11.5 cm long, glabrous. Inflorescences sparsely pubescent with
hairs up to 0.1 mm long; bract caducous; males 710 58 cm, 34-times branched,
many-flowered, peduncle (2)612 mm long; females c. 3.5 cm long, stout. Flowers
glabrous; perianth 34-lobed; pedicel subglabrous, sometimes faintly articulated. Male
flowers in clusters of 35; pedicel 12 mm long; buds subglobose, 22.2 22.3 mm;
perianth at anthesis clefts to 2/33/4(4/5), lobes at base 0.20.3 mm thick; androecium
depressed globose-ovoid, apex narrowly, base broadly rounded, c. 1 11.2 mm, subcircular
in transverse section; anthers 810, almost sessile, more or less curved, almost entirely
connate, the free apices up to 0.1(0.2) mm long; apical cavity of column 0.10.2 mm
deep; androphore c. 0.2 mm long. Fruits 2 or 3 on a stout infructescence c. 3.5 cm
long, broadly ellipsoid, 55.5 3.54 cm, glabrous, drying dark brown, with few small
tubercles; dry pericarp 710 mm thick; perianth not persistent. Seeds ellipsoid; aril red.
Distribution. Borneo (Sarawak and E Kalimantan) and the Philippines (Palawan).
Ecology. In ridge forest on limestone and coral limestone, and lowland mixed dipterocarp
forest on yellow sandy soils, at 150730 m altitude. Flowering in August and November;
fruiting in May.
378
20. Horsfieldia obtusa W.J. de Wilde

(Latin, obtusus = blunt; the blunt or rounded leaf tip)
Gard. Bull. Sing. 39 (1986) 9; Whitmore, Tantra & Sutisna l.c. 265. Type: Native Collector BS 821,
Borneo, Sarawak (L).
Tree. Twigs solid, terete or faintly angular, not ridged, towards apex 2.54(6) mm
diameter, drying dark brown, not contrasting with the drying petiole, initially dark rusty
tomentose with hairs 0.20.4 mm long, late-glabrescent; bark finely cracked; lenticels
inconspicuous. Leaf buds c. 12 mm long, densely dark rusty pubescent with hairs 0.2
0.4 mm long. Leaves distichous, thinly coriaceous, not bullate, not scabrous, upper surface
glabrous (except the midrib), drying olivaceous, lower surface early glabrescent, drying
purplish brown, with yellowish enlarged hair scars, without blackish dots; oblong, 8
10 33.5 cm, base attenuate, apex rounded; midrib at base narrow, raised above, lateglabrescent; lateral veins c. 10 pairs, flat or slightly raised above; intercostal venation
laxly reticulate, not or hardly visible; petiole c. 1 cm long, late-glabrescent. Inflorescences
on older branches behind the leaves, densely rusty pubescent with hairs 0.20.4(0.5)
mm long; bract caducous; males 59 35 cm, 3-times branched, many-flowered; peduncle
310 mm long. Male flowers: pedicel glabrous, 11.5 mm long, at base articulated;
buds subglobose, 1.31.5 1.51.6 mm; perianth glabrous, 34-lobed, at anthesis clefts
to nearly 1/2, more or less woody, brittle, not collapsing on drying; androecium (including
androphore) broadly obovoid, c. 0.8 1 mm, in transverse section 34-angled; anthers
9 or 10, more or less erect, free in the upper half; androphore more or less tapering,
c. 0.3 mm long; apical cavity of column distinct. Female flowers and fruits not known.
Distribution. Endemic to Borneo. Only known by the type specimens from Sarawak,
precise locality not indicated.
21. Horsfieldia oligocarpa Warb.

(Greek, oligos = few, karpos = fruit; with few fruits)
Monogr. Myrist. (1897) 354; Merrill l.c. (1921) 268; Masamune l.c. 301; W.J. de Wilde l.c. (1986) 22;
Whitmore, Tantra & Sutisna l.c. 265; Coode et al. (eds.) l.c. 222. Lectotype (de Wilde, 1986): Beccari
2066, Borneo, Sarawak, Kuching district, Mt. Matang (hololectotype FI acc. no. 7620; isolectotypes K,
P). Synonyms: Myristica oligocarpa (Warb.) Boerl. l.c. 87; H. polyspherula (Hook. f.) J. Sinclair var.
oligocarpa (Warb.) J. Sinclair l.c. (1975) 104.
Tree up to 20 m tall. Bark grey, dark brown to reddish brown, broadly fissured; inner
bark pink-brown, soft, c. 10 mm thick; sap red. Sapwood pink-yellow. Twigs terete,
towards apex 23(4) mm diameter, glabrescent, drying pale yellowish or whitish-brown,
contrasting with the dark colour of the drying petiole; lenticels small, distinct; older bark
not flaky. Leaf buds 710 mm long, pubescent with hairs c. 0.2 mm long, the tomentum
early shed in the form of small crust-like pieces. Leaves distichous, chartaceous, not
bullate, not scabrous, upper surface glabrous, drying grey-greenish, dull, lower surface
generally chocolate-brown, contrasting with the upper surface, glabrous, without blackish
dots; elliptic-oblong to oblong, 716 2.56 cm, base attenuate, apex acute-acuminate;
midrib raised above, glabrous; lateral veins 811(15) pairs, raised above; intercostal
venation indistinct on both surfaces; petiole 0.61.2 cm long, drying blackish brown.
379
Inflorescences sparsely pubescent with hairs 0.20.3 mm long, sometimes glabrescent;

bract caducous; males 36 1.54 cm, 23-times branched, peduncle 510 mm long,
few-flowered; females not seen, according to the infructescences c. 1.54 cm long. Male
flowers: pedicel glabrous, 11.5 mm long, at base articulated; buds globose to broadly
obovoid, 11.7 mm diameter; perianth glabrous, 3-lobed, at anthesis clefts to nearly 1/
2, more or less coriaceous, not collapsing on drying; androecium (including androphore)
more or less obovoid, c. 1 mm across, triangular in transverse section; anthers 6 or
7, largely mutually free; androphore tapering, c. 0.5 mm long. Female flowers not seen.
Fruits 14 per infructescence, ellipsoid, 1.82.7 1.41.9 cm, glabrous, drying dark
brown, hardly lenticellate; pericarp 23 mm thick; stalk 25 mm long; perianth not
persistent. Seeds ellipsoid.
Distribution. Endemic to Borneo (Sarawak and Brunei).
Ecology. In peatswamp and heath forests on nutrient-poor soils (white sand, white podsolic
sand, yellow sand, sandstone terraces, and peat), up to 100 m altitude. Flowering in
June and August; fruiting in AugustOctober.
22. Horsfieldia pallidicaula W.J. de Wilde

(Latin, pallidus = pale; caulis = stem; with pale stem)
Gard. Bull. Sing. 38 (1985) 191; Whitmore, Tantra & Sutisna l.c. 265. Type: Jacobs 5413, Borneo,
Sarawak, Baram district (holotype L; isotypes CANB, G, K, SAR, SING).
Tree up to 20 m tall. Bark smooth, slightly flaky or fissured, dark grey. Twigs terete,
not ridged, towards apex 210 mm diameter, drying pale whitish or greyish brown,
contrasting with the blackish colour of the drying petiole, initially tomentose with hairs
0.10.3 mm long, early glabrescent; bark slightly flaky or not; lenticels inconspicuous.
Leaf buds 710 mm long, pubescent with hairs (0.1)0.2(0.3) mm long. Leaves either
in 2, 3 or 5 rows, not bunched at the top of twigs, membranous, not bullate, not scabrous,
upper surface drying dark brown, lower surface paler, without blackish dots, early
glabrescent; oblong to oblong-lanceolate, 1030 49.5 cm, base long-attenuate, apex
acute-acuminate; midrib flat or slightly raised above, glabrous; lateral veins 1018 pairs,
flat or slightly raised above; intercostal venation laxly reticulate, indistinct above; petiole
12.5 cm long, often greyish and flaky in the lower half. Inflorescences glabrescent,
tomentum weak, comprising hairs 0.10.2 mm long; bract 13 mm long, caducous; males
39 2.57 cm, (2)3-times branched, many-flowered, peduncle up to 7 mm long (hence
inflorescences often also branched from near the base); females few-flowered, 13 cm
long. Flowers: perianth 34-lobed, in females sometimes 2-lobed, glabrous; pedicel
glabrous, at base not articulated. Male flowers: pedicel 12 mm long, slender; buds
globose or broadly obovoid, 1.52(2.2) mm diameter; perianth at anthesis clefts to 1/
3 to 1/2, not or slightly collapsing on drying; androecium short-ellipsoid or depressed
globose, 0.81.2 0.81.2 mm, circular in cross-section; anthers 810, sessile; column
with narrow apical cavity 0.10.2 mm deep; androphore up to 0.2 mm long. Female
flowers: buds broadly ellipsoid or subglobose, 2.53 2.5 mm. Fruits 15 per infructescence,
broadly ellipsoid, 1.54 13 cm, glabrous, drying blackish, finely granulate, sometimes
tuberculate; pericarp various; stalk 13 mm long; perianth persistent (always?). Seeds
ellipsoid.
380
Distribution. Endemic to Borneo (Sabah, Sarawak and Kalimantan). Three varieties are
known.
Key to varieties
1. Fruits 3.74 2.73 cm; pericarp 68(10) mm thick. Leaves (23)2530 cm long.........
var. macrocarya W.J. de Wilde
(Greek, makro = large, karyon = nut; with large nut)
l.c. (1985) 193. Type: Ding Hou 474, Borneo, Sarawak (L).
Endemic to Borneo (Sarawak).

Fruits 1.52.2 11.8 cm; pericarp 13 mm thick. Leaves up to 25 cm long.............2
2.
Fruits 1.51.7 11.2 cm; pericarp 11.5 mm thick. Leaves up to 23 cm long............

var. microcarya (J. Sinclair) W.J. de Wilde
(Greek, mikro = small, karyon = nut; with small nut)
l.c. (1985) 193. Type: Wood & Kapis SAN 16971, Borneo, Sabah (holotype: K; isotypes
KEP, L, SAN, SING). Synonym: H. bracteosa Henderson var. microcarya J. Sinclair l.c.
(1975) 20.
Endemic to Borneo (W Sabah and E Kalimantan). In lowland mixed dipterocarp

forest up to 100 m altitude. Fruiting in July.
Fruits 22.2 1.8 cm; pericarp 23 mm thick. Leaves up to 25 cm long..............
var. pallidicaula
W.J. de Wilde l.c. (1985) 192.
Endemic to Borneo (Sabah, Sarawak; possibly also in W Kalimantan). In primary
lowland and hill mixed dipterocarp forests on sandstone, up to 700 m altitude.
Flowering throughout the year; fruiting in June.
23. Horsfieldia paucinervis Warb.

(Latin, pauci = few, nervium = nerve; the few-nerved leaves)
Anderson l.c. 265; W.J. de Wilde l.c. (1985) 212, l.c. (1996) 379; Whitmore, Tantra & Sutisna l.c. 265.
Type: Beccari 3279 (female flowers), Borneo, Sarawak (FI). Synonym: Myristica paucinervis (Warb.)
Boerl. l.c. 87.
Tree up to 8 m tall. Twigs terete, slender, towards apex 23 mm diameter, initially reddish
to yellow-brown tomentose with shaggy hairs 1.52 mm long, late-glabrescent; bark dark
grey, not cracked; lenticels not conspicuous. Leaf buds 810 mm long, pubescent. Leaves
distichous, thinly chartaceous, surface not scabrous, not bullate, upper surface drying
olivaceous, glabrous, lower surface thinly tomentose with persistent hairs 0.51 mm long,
with blackish dots (always?) but without streaks; elliptic to elliptic-oblong, 715 3
6.5 cm, base rounded to attenuate, apex acute to acute-acuminate; midrib flat and indistinct
above, late-glabrescent; lateral veins 59(11) pairs, flat or sunken above; petiole 0.6
1.2 cm long, pubescent. Inflorescences pubescent with hairs (0.7)12 mm long; bract
caducous; males 59 35 cm, peduncle up to 15 mm long; females 46 cm long. Flowers:
perianth 3-lobed, glabrous; pedicel glabrous, at base not articulated. Male flowers: pedicel
slender, 0.51.5 mm long; buds subglobose, 0.81 11.5 mm; perianth at anthesis clefts
381
to 1/3 to nearly 1/2; androecium small, more or less depressed globose, 0.30.5 0.4
0.8 mm, more or less circular in transverse section; anthers 4 or 5, sessile; apical cavity
of column c. 1/5 deep; androphore 0.10.2 mm long. Female flowers: buds ovoid-globose,
c. 2 mm diameter, ovary glabrous. Fruits (immature) with persistent perianth, glabrous,
oblong, c. 1 0.7 cm; stalk c. 2 mm long.
Distribution. Endemic to Borneo (Sarawak).
Ecology. In coastal kerangas and secondary forests on eroded white sands, up to 50
m altitude. Flowering throughout the year.
24. Horsfieldia penangiana J. Sinclair

(of Penang, Peninsular Malaysia)
Gard. Bull. Sing. 16 (1958) 408, l.c. (1975) 94; W.J. de Wilde l.c. (1986) 52, l.c. (1996) 379;
Turner l.c. 359. Type: Curtis 2406, Peninsular Malaysia, Penang (holotype SING; isotypes BM,
K).
subsp. obtusifolia W.J. de Wilde

(Latin, obtusus = blunt, folium = leaf; with blunt leaf apex)
l.c. (1996) 379. Type: Dayang Awa & Othman S. 47061, Borneo, Sarawak (holotype L; isotypes K,
SAR).
Tree up to 25 m tall. Twigs solid, terete, slender, towards apex 1.52(4) mm diameter,
drying grey-brown to brown, not contrasting with the drying petiole, tomentose with hairs
up to 0.1 mm long, glabrescent; bark not flaky; lenticels small. Leaf buds slender, 6
9 mm long, tomentose with very short hairs up to 0.1 mm long. Leaves distichous,
membranous to thinly coriaceous, not bullate, not scabrous, upper surface drying olivaceous
to blackish brown, lower surface early glabrescent, with scattered dark (rarely pale) brown
dots; elliptic-oblong to oblong, 614 (17) 24 cm, base attenuate, apex blunt or rounded;
midrib flattish to raised above; lateral veins 811 pairs, thin, flat and inconspicuous above;
petiole 0.81.3 cm long, glabrescent. Inflorescences with sparse tomentum comprising
very short hairs c. 0.1 mm long; bract 1.52.5 mm long, caducous; males 27 1.5
4.5 cm, peduncle 220 mm long; females 25 cm long. Male flowers: pedicel glabrous,
0.81.5(2) mm long, slender, at base more or less articulated or not; buds subglobose,
more or less circular to faintly triangular in transverse section, c. (1.5)2 mm diameter;
perianth 3-lobed, glabrous, at anthesis clefts to c. 1/3 to nearly halfway; androecium
depressed globose, subcircular in transverse section; anthers 12(15), sessile, more or
less curved, almost entirely connate, free apices up to 0.3 mm long, slightly incurved;
column with shallow cavity at the apex; androphore up to 0.1 mm long. Fruits 26 per
infructescence, ovoid-ellipsoid, c. 1.5 1 cm, glabrous, without lenticel-like tubercles;
dry pericarp c. 1.5 mm thick; stalk 34 mm long, at base articulated; perianth not persistent.
Distribution. Endemic to Borneo (Sarawak and E Kalimantan).
Ecology. In mixed dipterocarp forest on ridge tops and hills, at (50)300800 m altitude.
382
25. Horsfieldia polyspherula (Hook. f. emend. King) J. Sinclair

(Latin, poly = many; sphaera = globe or ball; flowers like many small balls)
Gard. Bull. Sing. 16 (1958) 422; l.c. (1975) 101 p.p. (for the type var. only); W.J. de Wilde l.c. (1986)
17, l.c. (1996) 381; Whitmore, Tantra & Sutisna l.c. 266; Coode et al. (eds.) l.c. 222. Basionym: Myristica
polyspherula Hook. f., Fl. Brit. Ind. 5 (1886) 108, p.p.; King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 312,
emend. Lectotype (J. Sinclair, 1975): Griffith 4354, Peninsular Malaysia (hololectotype K; isolectotypes
BM, CAL, CGE, FI, G, P, U).
Tree up to 40 m tall, with or without buttresses. Bark shallowly boat-shaped fissured

or flaky; inner bark yellow or pinkish, fibrous. Wood moderately soft, whitish yellow.
Twigs solid, not ridged, terete or subterete, towards apex 25(13) mm diameter, drying
grey-brown to blackish, not contrasting with the drying petiole, tomentose with rusty,
mealy, dendroid hairs 0.10.6 mm long, late-glabrescent; lenticels distinct or not; bark
striate, not cracked. Leaf buds 617 mm long, tomentose with hairs 0.10.6 mm long.
Leaves distichous, thickly membranous or chartaceous, not bullate, not scabrous, on drying
usually brittle, upper surface drying olivaceous or greenish brown, the midrib glabrescent,
lower surface drying light brown or chocolate, usually much contrasting with the upper
surface, without blackish dots, early glabrescent but with the midrib often late-glabrescent;
elliptic-oblong, ovate-oblong to oblong-lanceolate, 728 2.59 cm, base subrounded
or attenuate, apex acute-acuminate; midrib raised above, at base narrow; lateral veins
615 pairs, distinctly raised above except close to the leaf margin, glabrous; petiole 0.6
1.5 cm long, 0.150.3 cm diameter, late-glabrescent. Inflorescences sparsely to densely
woolly pubescent with hairs c. 0.6 mm long; bract 1.57 mm long, caducous; males 4
15 312 cm, peduncle 315 mm long; females smaller, up to 8 cm long. Flowers:
perianth 3-lobed, glabrous or glabrescent; pedicel sometimes minutely pubescent towards
the base, articulated. Male flowers: pedicel slender, 0.81.5(2) mm long; buds subglobose,
11.8 mm diameter, base rarely more or less tapering into the pedicel, glabrous; perianth
at anthesis clefts to 1/22/3, not collapsing on drying; androecium (including androphore)
more or less broadly obovoid, 0.50.8 0.61 mm, usually sharply triangular in transverse
section; anthers (3)47, free at least in the upper half, more or less curved or suberect,
0.30.5 mm long, apex somewhat acute; column bowl-shaped, passing into the tapering
androphore of 0.20.4 mm long. Female flowers glabrous or sparingly pubescent with
hairs c. 0.1 mm long; buds 23 mm across; ovary glabrous. Fruits 16 per infructescence,
subglobose or ellipsoid, 1.96 1.45 cm, base rounded or slightly attenuate, glabrous;
dry pericarp 215 mm thick; stalk 14 mm long; perianth not persistent.
Distribution. Sumatra, Peninsular Malaysia, Borneo, and southern parts of the Philippines.
A variable species, with three varieties, all occurring in Sabah and Sarawak.
Key to varieties
1.
Fruits (3.5)46 35 cm; pericarp (5)815 mm thick. Leaves 920 3.57 cm; lateral
veins (6)915 pairs........................................
var. maxima W.J. de Wilde
(Latin, maximus = largest; referring to the fruit)
l.c. (1986) 22; Whitmore, Tantra & Sutisna l.c. 266; Coode et al. (eds.) l.c. 223. Type: Chai
S. 36228, Borneo, Sarawak (holotype L; isotype SAR).
383
Endemic to Borneo (Sabah, Sarawak, Brunei and E Kalimantan). In mixed

dipterocarp forest and Agathis forest on sandy waterlogged soils, sandy loam or
yellow clay-loam soils; also known from kerangas forest; up to 500 m altitude.
Fruiting throughout the year.
Fruits up to 3.5 cm long; pericarp 25 mm thick. Leaves 728 2.59 cm; lateral veins
620 pairs.....................................................................................2
2. Fruits (1.7)1.92.5(2.8) 1.42 cm; pericarp 24 mm thick. Male flower buds
11.5 mm diameter; anthers (3 or 4)7. Female flower buds 22.5 mm long. Leaves
719 2.56 cm; lateral veins 615 pairs. Terminal leaf buds covered with hairs
(0.2)0.30.6 mm long................................
var. polyspherula
W.J. de Wilde l.c. (1986) 19; Whitmore, Tantra & Sutisna l.c. 266; Coode et al. (eds.) l.c. 223.
Sumatra, Peninsular Malaysia, Borneo, and the Philippines. In Borneo, recorded

from Sabah, Sarawak, Brunei and E Kalimantan. In lowland forest, most often
on sandy soils; also in Casuarina forest (E Borneo), freshwater swamp forest,
ridge-top forest, kerangas forest (Sarawak, rare); up to 900 m altitude. Flowering
throughout the year, but most collections in JuneSeptember; fruiting throughout
the year.
Fruits (2.5)2.83.5 2.22.7 cm; pericarp 35 mm thick. Male flower buds 1.2
1.8 mm diameter; anthers 6 or 7. Female flower buds c. 3 mm long. Leaves 13
28 4.59 cm; lateral veins 1115(20?) pairs. Terminal leaf buds covered with
hairs 0.10.4 mm long.....................
var. sumatrana (Miq.) W.J. de Wilde
(of Sumatra)
l.c. (1986) 20; Whitmore, Tantra & Sutisna l.c. 266; Turner l.c. 359; Coode et al. l.c. 223.
Basionym: Myristica glabra Blume var. sumatrana Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865)
49. Type: Korthals s.n., W Sumatra (holotype L; isotypes A, K, U). Synonyms: H. brachiata
(King) Warb. var. sumatrana (Miq.) J. Sinclair l.c. (1975) 13, p.p.; M. collettiana King, l.c.
312.
Sumatra, Peninsular Malaysia, Singapore, and Borneo. In lowland mixed dipterocarp

forest, ridge forest, montane forest; also in heath and peatswamp forests, on sandy
and sandy-loamy soils, red soils, yellow clayey soils; up to 1100 m altitude.
Flowering and fruiting throughout the year.
26. Horsfieldia punctatifolia J. Sinclair

(Latin, punctatus = dotted, folium = leaf; the punctate leaves)
Gard. Bull. Sing. 16 (1958) 413, l.c. (1975) 104; Whitmore l.c. 327; Anderson l.c. 265; W.J. de Wilde
l.c. (1986) 54; Whitmore, Tantra & Sutisna l.c. 266; Turner l.c. 359; Coode et al. (eds.) l.c. 223;
Argent et al. (eds.) l.c. 452. Type: Sinclair SFN 40211 (= Sinclair 7989), Singapore (holotype SING;
isotypes B, BO, DD, E, K, L, P).
Tree up to 30 m tall. Bark smooth but shallowly fissured or cracked, or brittle and scaly;
inner bark pinkish to red; sap reddish, watery. Sapwood white, yellowish, or pinkish;
heartwood dark brown. Twigs solid, terete, not ridged, towards apex 2.54(10) mm
diameter, drying dark grey-brown, not contrasting with the drying petiole, early grey-
384
brown tomentose with hairs up to 0.1 mm long, glabrescent; bark not flaky; lenticels
present. Leaf buds 812 mm long, pubescent with hairs up to 0.1 mm long. Leaves
distichous, membranous, not bullate, not scabrous, upper surface drying olivaceous or
brown, lower surface glabrous, with scattered blackish dots; elliptic-oblong to oblong,
921 39 cm, base attenuate, apex acute-acuminate; midrib flat or slightly raised above;
lateral veins1116 pairs, thin and flat above; petiole 11.7 cm long, glabrous. Inflorescences
sparsely pubescent with hairs up to 0.1 mm long or glabrescent; bract 24 mm long,
caducous; males 410 28 cm, peduncle 520 mm long; females 36 cm, fewer-flowered.
Flowers glabrous; perianth 34-lobed; pedicel glabrous, not articulated. Male flowers:
pedicel 12(2.5) mm long, slender; buds depressed globose, 1.42 1.62.2 mm; perianth
at anthesis clefts to 3/44/5; androecium depressed, broadly ovoid, 0.60.7 11.5 mm,
subtriangular with rounded angles in cross section, subsessile; anthers 79, more or less
curved, almost entirely connate, free apices up to 0.1 mm long, concealing apical cavity
of column up to 0.2 mm deep. Female flowers: buds ellipsoid, 2.83.5 mm long; ovary
glabrous. Fruits 13 per infructescence, broadly ellipsoid, (4.5)58 (3)3.54.5 cm,
glabrous; dry pericarp 1020 mm thick; stalks 46 mm long; perianth not persistent.
Seeds ellipsoid; aril red.
Brunei, C and E Kalimantan).
Ecology. In primary lowland and hill forests, swamp forest and kerangas forests, on a
variety of soils including grey, brown, sandy clay, and sandstone- and dacite-derived soils,
up to 1100 m altitude. Flowering and fruiting throughout the year.
27. Horsfieldia reticulata Warb.

(Latin, reticulatus = netted; the intercostal venation of the leaf)
Anderson l.c. 265; W.J. de Wilde l.c. (1985) 218, l.c. (1996) 381; Whitmore, Tantra & Sutisna l.c. 266.
Type: Beccari 3475, Borneo, Sarawak (holotype FI; isotype K). Synonyms: Myristica reticutata (Warb.)
Boerl. l.c. 85; H. affinis W.J. de Wilde l.c. (1985) 217.
Tree up to 20 m tall. Bark rough, dark brown, lenticellate; inner bark red. Twigs terete,
not ridged, towards apex 2.56(9) mm diameter, tomentose with hairs 0.51 mm long,
late-glabrescent; bark when old dark grey, cracked but not flaky; lenticels inconspicuous.
Leaf buds 1015 mm long, pubescent with hairs 0.51 mm long. Leaves distichous,
membranous or chartaceous, both surfaces not scabrous, sometimes bullate, upper surface
drying brown, glabrous, lower surface sparsely tomentose with subpersistent, mixed sessile
and dendroid hairs 0.51 mm long, without blackish dots or streaks; elliptic-oblong to
lanceolate, 1835 411 cm, base rounded to attenuate, apex acute-acuminate; midrib
flattish or slightly raised above, often late-glabrescent; lateral veins (9)1720 pairs,
slender, sunken or raised above; petiole (0.7)1.22 cm long, pubescent. Inflorescences
pubescent with shaggy hairs 0.52 mm long; bract 39 mm long, caducous; males 12
25 814 cm, peduncle 1560 mm long. Male flowers: pedicel slender, 0.52 mm long,
glabrous, at base articulated; buds depressed globose, 1.22 1.52(2.5) mm; perianth
glabrous, 35-lobed, at anthesis clefts to 1/41/3; androecium depressed globose, more
or less circular in transverse section, 0.60.8 11.5 mm; anthers 1012(15), sessile;
385
apical cavity of column up to halfway deep; androphore 0.20.4 mm long, hidden or

not by the anthers. Female flowers not seen. Fruits 47 per infructescence, ellipsoid,
2.32.7 1.72.1 cm, glabrous, drying brown, pale wart-like tubercles present; dry pericarp
1.52 mm thick; stalk 34 mm long; perianth persistent. Seeds ellipsoid.
Distribution. Endemic to Borneo (Sarawak, Brunei and Kalimantan).
Ecology. In lowland and hill mixed dipterocarp forests on alluvial, sandy clay, and
sandstone-derived soils, up to 600 m altitude. Flowering in MayNovember; fruiting in
July.
28. Horsfieldia ridleyana (King) Warb.

(H.N. Ridley, 18551956, botanist and former Director of the Singapore Botanic Gardens)
Monogr. Myrist. (1897) 331; Gamble l.c. 221; Ridley l.c. 60; Burkill, EPMP 1, (1935) 1199; J. Sinclair
l.c. (1958) 432, l.c. (1975) 108; W.J. de Wilde l.c. (1986) 7; Whitmore, Tantra & Sutisna l.c. 266; Turner
l.c. 359; Coode et al. (eds.) l.c. 223. Basionym: Myristica ridleyana King, Ann. Roy. Bot. Gard. Calc. 3
(1891) 311. Syntypes: Scortechini s.n. & 862, Peninsular Malaysia, Perak (holotype CAL; isotypes
BM, K, L).
Tree up to 25 m tall. Bark dark brown to red-brown, shallowly and rectangularly fissured,
shallowly cracked, flaky or shallowly dippled and fissured; inner bark reddish, fibrous,
laminated. Sapwood pale, whitish, or creamy pink. Twigs solid, not ridged, terete or
somewhat angular, towards apex 1.53.5(5) mm diameter, drying dark brown and not
contrasting with the drying petiole, rusty tomentose with hairs 0.20.4 mm long, early
glabrescent; lenticels either small and distinct, or absent; older bark not flaky. Leaf buds
614 mm long, pubescent with hairs 0.20.5 mm long. Leaves distichous, thinly chartaceous
to coriaceous, not bullate, not scabrous, upper surface glabrous, drying olivaceous or
brown, lower surface glabrous, drying light reddish-brown, sometimes contrasting with
the upper surface, without blackish dots; elliptic-oblong to lanceolate, 515(16) 2
4.5 cm, base attenuate, apex acute to acute-acuminate; midrib at base narrow, flattish
above; lateral veins 715 pairs, sunken to flattish above, inconspicuous; petiole 0.7
1.5 cm long, early glabrescent. Inflorescences sparsely pubescent with hairs 0.10.3 mm
long, or glabrescent; bract caducous; males 26 24 cm, peduncle (2)515 mm long;
females fewer-flowered, 1.54 cm long. Flowers: perianth 34(rarely 2)-lobed, glabrous;
pedicel slender, glabrous, at base articulated. Male flowers: pedicel 12 mm long; buds
subglobose or short-ellipsoid, 11.2 0.81.2(1.3) mm, base rounded or short-attenuate;
perianth at anthesis clefts to 1/3 to nearly 1/2, not or slightly collapsing on drying;
androecium (including androphore) broadly obovoid, 0.60.8 0.40.7 mm, in transverse
section 34-angular (rarely more or less ellipsoid in flowers with 2-lobed perianth); anthers
46, more or less erect, mutually almost free, 0.30.4 mm long; androphore tapering.
Female flowers: buds ellipsoid, 1.51.8 1.5 mm; ovary glabrous. Fruits 16 per
infructescence, ellipsoid, 1.52 11.4 cm, glabrous; dry pericarp c. 1.5 mm thick; stalk
25 mm long; perianth not persistent. Seeds ellipsoid.
Distribution. Peninsular Malaysia and Borneo (Sabah, Sarawak and Brunei).
Ecology. Usually in forest on poor soils, e.g., heath forest (with Dacrydium beccarii),
386
forest on quartzite-ridge, and forest on sandstone ridge with very shallow soils (together
with Gymnostoma, Tristaniopsis and Cotylelobium), up to 1100 m altitude. Flowering
in JuneOctober; fruiting throughout the year.
29. Horsfieldia rufo-lanata Airy Shaw

(Latin, rufus = reddish; lanatus = woolly; the indumentum)
Kew Bull. 1939, no. 10 (1940) 440; Masamune l.c. 301; J. Sinclair l.c. (1975) 111; Anderson l.c. 265;
W.J. de Wilde l.c. (1985) 216; Whitmore, Tantra & Sutisna l.c. 266. Type: Richards 1667, Borneo,
Sarawak (holotype K; isotype SING).
Tree up to 17 m tall. Twigs terete, not ridged, towards apex 3.55(7) mm diameter,
initially tomentose with reddish brown hairs 11.5 mm long, late-glabrescent; bark dark
brown, longitudinally cracked, not flaky; lenticels indistinct. Leaf buds 1520 mm long,
pubescent with hairs 11.5 mm long. Leaves distichous, chartaceous, surface slightly
or not bullate, not scabrous, upper surface drying olivaceous, glabrous except the lateral
veins, lower surface densely or sparsely tomentose with hairs 0.31.5 mm long, without
blackish dots; elliptic to oblong, 1023 510(12) cm, base nearly rounded to subattenuate,
apex acute to acute-acuminate; midrib more or less raised above, late-glabrescent; lateral
veins 1116 pairs, raised above, pubescent or late-glabrescent; intercostal venation
indistinct above; petiole 11.6 cm long, pubescent. Inflorescences woolly pubescent with
shaggy hairs 23 mm long; bract 38 mm long; males 812 57 cm, peduncle 15
30 mm long; females few-flowered, c. 3 cm long. Male flowers: pedicel glabrous, 0.5
1 mm long, at base indistinctly articulated; buds subglobose, hardly or not depressed,
22.3 mm diameter, rather firm; perianth glabrous, 34-lobed, at anthesis clefts to nearly
1/2; androecium globose, c. 1.2 mm diameter, circular in transverse section; anthers c.
15, sessile; column broad, solid except for small apical cavity to 1/5 deep; androphore
c. 0.2 mm long. Female flowers not known. Fruits 24 per infructescence, broadly ovoidellipsoid, 22.2 1.51.6 cm, glabrescent, remnants of minute hairs c. 0.1 mm long
present at the base; pericarp drying brown, without lenticels or tubercles, c. 2 mm thick;
stalk 13 mm long; perianth not persistent.
Ecology. In hill mixed dipterocarp forest on sandstone and lower montane forest, at 900
1400 m altitude. Flowering in June and September; fruiting in September.
30. Horsfieldia sabulosa J. Sinclair
Fig. 5.
(Latin, sabulosus = growing in sandy places)

Gard. Bull. Sing. 27 (1974) 133; Anderson l.c. 265; W.J. de Wilde l.c. (1985) 185; Whitmore, Tantra &
Sutisna l.c. 266; Coode et al. (eds.) l.c. 223. Type: Sinclair & Kadim 10491, Borneo, Brunei, Bt. Labi
(holotype K; isotypes A, B, E, K, L, NY, SAR, SING).
Tree up to 37 m tall. Bark dark grey, longitudinally fissured; inner bark reddish brown,
laminated, fibrous; sap red, copious. Sapwood soft, pinkish. Twigs stout, terete, with
wart-like thickened markings of leaf scars, (3)510(20) mm diameter, at first grey-
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brown to rusty tomentose with hairs 0.51(1.5) mm long, late-glabrescent; bark blackish,
cracked and often conspicuously flaking; lenticels inconspicuous. Leaf buds stout, 10
15 (3)45 mm, densely rusty pubescent with hairs 0.51.5 mm long. Leaves in 3
5 rows, generally bunched towards the apex of the twigs, coriaceous, upper surface glabrous,
more or less shining or not, drying olivaceous to brown, lower surface pale brown,
glabrescent, with scattered blackish, roundish or sometimes elliptic or line-shaped dots;
elliptic-oblong to sublanceolate, 921 26 cm, base attenuate, apex acute to shortacuminate; midrib flat above; lateral veins 1220 pairs, flat above and below; petiole
2.55 cm long, pubescent with hairs 0.51.5 mm long, late-glabrescent. Inflorescences
on older twigs of 1520 mm thick behind the leaves, emerging from the axils of woody
wart-like petiole scars on the older bark, densely pubescent with hairs 11.5 mm long;
bract more or less boat-shaped, 415 mm long, outside pubescent with hairs 0.51.5 mm
long, inside glabrous, late-caducous; males 1013 56 cm, peduncle 1525 mm long;
females few-flowered, 35 cm long. Flowers: perianth glabrous, 34-lobed; pedicel
glabrous, at base articulated. Male flowers: pedicel 1.5(2) mm long; buds
(depressed)globose, 2 22.3 mm; perianth at anthesis clefts to (1/2)1/3, not collapsing
on drying; androecium subsessile, depressed globose, 11.2 1.71.8 mm; anthers 12
(15), sessile; apical cavity of column shallow. Female flowers: ovary glabrous. Fruits
16 per infructescence, ovoid, 35 2.54 cm, glabrous, drying dark brown or blackish;
dry pericarp hard, 58(10) mm thick; stalk 28 mm long; perianth not persistent.
Ecology. In mixed dipterocarp forest on sandy or peaty soils, deep yellow sands, sandy
loam, or heavy yellow clay soils; also in Agathis forest and ridge forest, up to 100 m
altitude. Flowering in MayJuly; fruiting in MayDecember.
31. Horsfieldia sessilifolia W.J. de Wilde

(Latin, sessilis = without stalk; folium = leaf; with sessile leaves)
Gard. Bull. Sing. 38 (1985) 201; Whitmore, Tantra & Sutisna l.c. 267. Type: Turkey S. 27808, Borneo,
Sarawak, Sibu district, Loba Kabang FR, Bt. Ingan (holotype L; isotypes K, SAR, SING).
Tree up to 30 m tall, 35 cm diameter, with stilt roots. Twigs stout, terete, not ridged,
towards apex 57(10) mm diameter; bark lower down glabrous, striate and densely set
with conspicuous lenticels. Leaves distichous, chartaceous, not scabrous, upper surface
glabrous, drying dark brown, lower surface densely pubescent with interwoven hairs
0.1(0.3) mm long, without dark brown dots; oblong-lanceolate, more or less parallelsided, c. (25)50(70) 14 cm, base broadly rounded to subcordate, apex acute-acuminate;
midrib flattish above; lateral veins 30(35) pairs, flat or slightly sunken above; petiole
0.20.3 cm long, or leaves sessile. Male inflorescences not seen. Female inflorescences
45 3.5 cm, many-flowered; branches rusty pubescent with woolly hairs 0.30.5 mm
long; peduncle c. 1 0.8 cm; bract c. 1 cm long, inside subglabrous, outside pubescent
as the inflorescence, caducous. Female flowers: pedicel broad, c. 1 mm long, minutely
pubescent; buds obovoid-ellipsoid, stout, 4.55 44.5 mm; perianth 3-lobed, coriaceous,
pubescent with hairs c. 0.1 mm long in the lower half, at anthesis clefts to 1/81/10
only, lobes c. 0.5 mm thick; ovary ovoid, subglabrous. Fruits not seen.
389
Vernacular name. Sarawakkumpang tembaga (Malay, Iban).

Distribution. Endemic to Borneo (Sarawak; known only by the type specimens).
Ecology. In lowland mixed swamp forest, below 100 m altitude. Flowering in June.
32. Horsfieldia splendida W.J. de Wilde
Fig. 6.
(Latin, splendens = beautiful)

Gard. Bull. Sing. 38 (1985) 213; Whitmore, Tantra & Sutisna l.c. 267. Type: Othman S. 33723, Borneo,
Sarawak (holotype L; isotypes SAN, SAR).
Tree up to 30 m tall. Bark blackish, brown, or red-brown, fissured or flaky; inner bark
soft, light brown or reddish, laminated; sap reddish. Sapwood yellowish or whitish. Twigs
stout, terete, not ridged, towards apex 47(13) mm diameter, initially tomentose with
woolly hairs 11.5 mm long, late-glabrescent; bark dark brown, flaky; lenticels indistinct.
Leaf buds 15(20) mm long, pubescent with hairs 11.5 mm long. Leaves distichous,
membranous or chartaceous, surface not bullate, not scabrous, upper surface drying
olivaceous or brown, early glabrescent, lower surface conspicuously tomentose with
dendroid, more or less evenly spaced, about equal-sized hairs 0.51.5 mm long, without
blackish dots; elliptic-oblong to oblanceolate, 1845 6.517 cm, base almost rounded
to attenuate, apex acute-acuminate; midrib slightly raised above, late-glabrescent; lateral
veins 1825 pairs, sunken above, glabrescent, distinct; intercostal venation distinct above;
petiole 1.21.7 cm long, pubescent. Inflorescences woolly pubescent with hairs 11.5
mm long; bract 0.41 cm long, caducous; males 4-times branched, many-flowered, 6
16 312(16) cm, peduncle 520 mm long; females 35 cm long. Male flowers glabrous
or thinly pubescent; pedicel slender, 11.5 mm long, pubescent with hairs 0.30.5 mm
long, subglabrescent, at base not articulated; buds depressed subglobose, 1.42 1.5
2 mm; perianth glabrous, 34-lobed, at anthesis clefts to 1/3 to nearly halfway; androecium
depressed subglobose, 0.50.6 11.3 mm; anthers 810, sessile, column with apical
cavity to 1/3(1/2) deep; androphore 0.3(0.5) mm long. Fruits 27 per infructescence,
broadly ellipsoid, 22.2 1.51.8 cm, pubescent with hairs c. 0.5 mm long; dry pericarp
c. 3 mm thick; stalk c. 1 mm long; perianth persistent.
Ecology. In lowland mixed dipterocarp forest, edges of swamp forest, kerangas forest
(with Dryobalanops fusca dominant), and lower montane forest, on brown soils, yellowish
soils, and tuff-plateau, at 6001500 m altitude. Flowering and fruiting throughout the
year.
33. Horsfieldia sterilis W.J. de Wilde

(Latin, sterilis = sterile; the few reduced anthers)
Gard. Bull. Sing. 38 (1985) 224; Whitmore, Tantra & Sutisna l.c. 267. Type: Aban SAN 30597, Borneo,
Sabah, Tawau district (holotype L; isotypes K, SAN, SING).
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Tree or shrub, up to 12 m tall. Bark grey-brown, non-fissured; inner bark with orangered sap. Sapwood pale yellow. Twigs terete, towards apex 2.54(8) mm diameter, drying
greyish brown, sometimes contrasting with the blackish colour of the drying petiole,
tomentose with hairs 0.10.2 mm long, glabrescent; bark not tending to crack or to flake;
lenticels conspicuous or not. Leaf buds slender, 712 mm long, tomentose with hairs
0.10.2 mm long. Leaves distichous, membranous, upper surface glabrous, drying blackish
brown, lower surface glabrous, without dark dots; elliptic to oblong-lanceolate, 1337
4.59 cm, base attenuate, apex acute-acuminate; midrib flat above, glabrous; lateral
veins 1118 pairs, flattish or slightly raised above; petiole 0.72 cm long. Inflorescences
tomentose with weak hairs 0.10.2 mm long, subglabrescent; bract 38 mm long, caducous;
males lax, 1020 58 cm, with few side branches, peduncle 313 cm long, with several
bract scars, or branched from the base; females elongate, slightly branched, almost spikelike, 510 cm long. Male flowers: pedicel 11.5 mm long, glabrous, at base not articulated;
buds broadly obovoid-subglobose, 1.51.7 mm diameter, base subattenuate, slender;
perianth glabrous, 2lobed, at anthesis clefts to c. 1/4; androecium broadly obovoid, c.
0.8 0.60.7 mm, subcircular in transverse section; anthers 3 or 4 (or 6 or 8), sessile,
occupying only the apical part of the androecium, with narrow thecae, column with apical
cavity absent or very inconspicuous; androphore conspicuous, tapering, c. 0.4 mm long.
Fruits 410 per spike-like infructescence, ellipsoid, 2.21.8 cm long, glabrous, drying
blackish, not tuberculate; pericarp c. 2 mm thick; stalk 34 mm long; perianth persistent.
Distribution. Endemic to Borneo (SE Sabah and C Kalimantan).
Ecology. In mixed dipterocarp forest on hillsides and riverbanks, at 100500 m altitude.
Flowering from AprilJuly.
34. Horsfieldia subalpina J. Sinclair

(Latin, sub- = somewhat, alpinus = growing in alpine zone of mountains; growing at rather
high altitude)
Gard. Bull. Sing. 16 (1958) 410, l.c. (1975) 131; W.J. de Wilde l.c. (1986) 40; Turner l.c. 359. Type:
Wray 467, Peninsular Malaysia, Perak, G. Batu Putih (holotype L; isotypes BM, CAL, K).
subsp. kinabaluensis W.J. de Wilde

Gard. Bull. Sing. 39 (1986) 41; Whitmore, Tantra & Sutisna l.c. 267. Type: Clemens 33136, Borneo,
Sabah (holotype L; isotypes A, BM, K, SING).
Tree up to 30 m tall. Bark slightly fissured, reddish brown; inner bark fibrous, whitish
turning to brown, and then yellowish. Sapwood white to yellowish, medium-hard, heartwood
not differentiated. Twigs solid, not ridged, terete, drying dark grey-brown, not contrasting
with the drying petiole, towards apex 2.55(12) mm diameter, tomentose with hairs up
to 0.1 mm long, early glabrescent; bark not flaky; lenticels usually conspicuous. Leaf
buds slender, 1220 mm long, pubescent with hairs up to 0.1 mm long. Leaves distichous,
membranous to chartaceous, not bullate, not scabrous, upper surface drying olivaceousbrown, lower surface glabrous, without blackish dots or distinct hair scars; elliptic-oblong
to oblong, 1527 510 cm, base attenuate, apex sometimes somewhat blunt; midrib
392
flat or slightly raised above; lateral veins 918(20) pairs, flattish above; petiole 0.5
1.5 cm long, glabrous. Inflorescences on older twigs behind the leaves, sparsely pubescent
with hairs c. 0.1 mm long; bracts 25 mm long, caducous; males 514 310 cm, peduncle
(6)1030 mm long; females 27 cm long. Flowers: perianth 34-lobed, glabrous; pedicel
glabrous, at base not articulated. Male flowers: pedicel 1.52 mm long; buds broadly
ellipsoid, 1.72 1.51.8 mm; perianth at anthesis clefts to halfway, lobes at base 0.2
0.3 mm thick; androecium broadly ellipsoid, 1.11.2 0.81 mm; anthers 8 or 9, sessile,
more or less curved, almost entirely connate, concealing apical cavity of the column
up to 0.30.5 mm deep; androphore hidden by the anthers. Female flowers: pedicel c.
1.5 mm long; buds ellipsoid, 22.5 mm long; ovary glabrous. Fruits 26 per infructescence,
broadly ellipsoid, 35 cm long, top narrowly rounded, glabrous; dry pericarp 45 mm
thick; stalk 37 mm long; perianth not persistent.
Distribution. Endemic to Borneo (Sabah: Mt. Kinabalu and vicinity; one doubtful
collection, Chai S. 35461, from Sarawak).
Ecology. In lower to upper montane forests on clayish soils, at 14002000 m altitude.
35. Horsfieldia sucosa (King) Warb.

(Latin, succosus = juicy; the sappy twigs)
Monogr. Myrist. (1897) 322; Gamble l.c. 217; Ridley l.c. 58; J. Sinclair l.c. (1958) 416 (p.p. incl. lectotype,
excl. fig. 45 = H. sparsa), l.c. (1975) 139 p.p.; Whitmore l.c. 328; Anderson l.c. 265; W.J. de Wilde l.c.
(1985) 188; Turner l.c. 359; Argent et al. (eds.) l.c. 452. Basionym: Myristica sucosa King, Ann. Bot.
Gard. Calc. 3 (1891) 301. Lectotype (W.J. de Wilde, 1985): Kings Collector 4647, Peninsular Malaysia,
Perak, Gopeng (holotype K; isotypes CAL, G, L). Synonyms: H. bracteosa Henderson, Gard. Bull. Str.
Settl. 7, 2 (1933) 120, J. Sinclair l.c. 16 (1958) 419; H. bracteosa Henderson var. bracteosa J. Sinclair
l.c. (1975) 18.
subsp. bifissa W.J. de Wilde

(Latin, bifissus = 2-cleft; the perianth)
l.c. (1985) 190; Whitmore, Tantra & Sutisna l.c. 267. Type: Soegeng 58, Borneo, Kalimantan, Kutai
(holotype L; isotype BO).
Tree up to 20 m tall. Bark smooth, undulately fissured; inner bark c. 10 mm thick, laminated,
reddish brown or yellow-brown. Sapwood whitish or pale brown, or reddish. Twigs terete,
towards apex 210 mm diameter, drying whitish or straw-coloured, contrasting with the
blackish petiole, tomentose with hairs 0.10.3 mm long, very early glabrescent; bark with
a tendency to flake; lenticels conspicuous on the young parts. Leaf buds 1015 mm long,
pubescent with hairs 0.10.3 mm long. Leaves in 2 or 3 rows, membranous to thinly
chartaceous, upper surface glabrous, drying blackish brown, often with a greenish tinge,
lower surface early glabrescent, without blackish dots, but rather smaller blackish points
often present; elliptic-oblong to lanceolate, 1428 4.58.5 cm, base attenuate or longattenuate, apex acute-acuminate; midrib flat or only slightly raised above, glabrous; lateral
veins 1317 pairs, flat or slightly raised above; petiole 12 cm long. Inflorescences on
older twigs behind the leaves, pubescent with hairs 0.10.2 mm long, late-glabrescent;
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bract 1.54 mm long, late-caducous; males lax or condensed, 719 516 cm, peduncle
1020 mm long; females branched, few-flowered, 12 cm long. Flowers: perianth mostly
2-lobed, glabrous; pedicel usually glabrous, at base articulated. Male flowers: pedicel
12 mm long, slender; buds depressed globose, 1.21.4 1.51.7 mm; perianth at anthesis
clefts to 1/3 to nearly 1/2; androecium depressed globose, 0.30.4 0.80.9 mm, broadly
ellipsoid in transverse section; anthers 7, sessile; column broad, more or less saucershaped; androphore narrow, 0.10.3 mm long. Female flowers: buds broadly ellipsoid,
2.83.5 mm long; ovary glabrous. Fruits 14 per infructescence, broadly ovoid-ellipsoid,
23.5 2 cm, glabrous, drying blackish, sometimes more or less tuberculate; pericarp
c. 4 mm thick; stalk 12 mm long; perianth not persistent.
Ecology. In lowland mixed dipterocarp forest on sandy loam soils, Shorea laevifolia forest
on sandy ridge, and once recorded in forest over limestone, up to 800 m altitude. Flowering
36. Horsfieldia tenuifolia (J. Sinclair) W.J. de Wilde
Fig. 7.
(Latin, tenuis = thin, folium = leaf; thin-leaved)

Gard. Bull. Sing. 39 (1986) 11; Whitmore, Tantra & Sutisna l.c. 267; Coode et al. l.c. 223. Basionym:
H. polyspherula (Hook. f. emend. King) J. Sinclair var. tenuifolia J. Sinclair l.c. (1975) 105. Type:
Bujang S. 13686, Borneo, Sarawak (holotype SING; isotypes K, L, S, SAR).
Tree up to 15 m tall. Bark dark brown to greenish brown, narrowly fissured, not flaky;
inner bark dark red; sap red. Sapwood whitish. Twigs solid, not ridged, terete, drying
not or little contrasting with the drying petiole, towards apex 13(4) mm diameter, initially
tomentose with hairs (0.1)0.2 mm long, glabrescent; bark striate or finely cracked; lenticels
inconspicuous or absent. Leaf buds 58 mm long, pubescent with hairs 0.10.2(0.3)
mm long. Leaves distichous, membranous to thinly chartaceous, not bullate, not scabrous,
upper surface drying olivaceous to brown, not much contrasting with the lower surface,
glabrous, lower surface glabrous, without blackish dots; elliptic to oblong-lanceolate, 6.5
16.5 36.5 cm, base attenuate, apex acute-acuminate; midrib raised above, glabrous
or sometimes late-glabrescent, at base narrow; lateral veins 511 pairs, distinctly raised
above; petiole 0.81.6 cm long, 0.10.2 cm diameter. Inflorescences sparsely tomentose
with stellate hairs 0.10.2 mm long, glabrescent; bract 23 mm, caducous; males 3
5 24 cm, peduncle 210 mm long; females slender, few-flowered, 23.5 cm long.
Flowers: perianth 34-lobed, glabrous; pedicel with few hairs in the lower half, at base
indistinctly or not articulated. Male flowers: pedicel 12 mm long; buds subglobose,
tapering into the pedicel, 0.81.3 11.5 mm; perianth at anthesis clefts to 1/3 to nearly
1/2; androecium almost obovoid, 0.50.7 0.50.8 mm, in transverse section triquetrous;
anthers 46, more or less erect, free for about halfway, central column largely hollow;
androphore tapering, 0.20.3 mm long. Female flowers: buds ellipsoid, c. 2 mm long;
ovary glabrous. Fruits 13 per infructescence, ellipsoid, 1.72 1.41.5 cm, glabrous,
drying dark brown, not tuberculate; pericarp 11.5 mm thick; stalk 35 mm long; perianth
not persistent.
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Ecology. In lowland mixed dipterocarp forest on ridges and slopes, up to 300 m altitude,
on yellow clay, yellow sandy clay, yellow loam, and brownish soils. Flowering and fruiting
from MaySeptember.
37. Horsfieldia tristis W.J. de Wilde

(Latin, tristis = dull-coloured; the drying leaves)
Gard. Bull. Sing. 38 (1985) 197; Whitmore, Tantra & Sutisna l.c. 267. Type: Lai S. 37470, Borneo,
Sarawak (holotype L; isotypes K, KEP, MO, SAN, SAR, n.v.).
Tree up to 15 m tall. Twigs terete, not ridged, towards apex 35(7) mm diameter, pale
brown to yellowish, contrasting with the drying petiole, tomentose with hairs up to 0.1
mm long, early glabrescent; bark cracked or flaky; lenticels smallish. Leaf buds slender,
1015 mm long, tomentose with hairs up to 0.1 mm long. Leaves distichous, parchmentlike, membranous to thinly chartaceous, upper surface not scabrous, drying dull olivaceousbrown and finely wrinkled, lower surface glabrous, not much contrasting in colour with
the upper surface, without blackish dots; elliptic-oblong to lanceolate, 1432 48.5
cm, base attenuate, apex acute-acuminate; midrib flat or slightly raised above, glabrous;
lateral veins 1117 pairs, flat or slightly raised or sunken above; petiole 0.61.2 cm long.
Inflorescences sparsely tomentose with minute hairs c. 0.1 mm long; males 512 3.5
7 cm, peduncle 725 mm long; females (in fruit) 23 cm long, bracts caducous. Male
flowers: pedicel slender, 1.55 mm long, glabrous, at base not articulated; buds broadly
ellipsoid, 2.83.7 23 mm; perianth 34-lobed, glabrous, at anthesis clefts to 1/5
1/4 (to nearly 1/3); androecium obovoid-ellipsoid, in transverse section suborbicular or
bluntly 34-angular, 22.7 1.52 mm; anthers 1220, almost sessile; apical cavity of
the column 0.50.8 mm deep; androphore 0.20.3 mm long, largely hidden by the anther
bases. Female flowers not seen. Fruits 48 per infructescence, ellipsoid, c. 1.5 1.2
cm, apex subacute, glabrous, drying blackish, without lenticels or tubercles; pericarp c.
1.5 mm thick; stalk 12 mm long; perianth not persistent.
Distribution. Sumatra and Borneo (Sarawak and S Kalimantan).
Ecology. In forest on flat lands, up to 100 m altitude. Flowering in August and November,
fruiting in September.
38. Horsfieldia wallichii (Hook. f. & Thomson) Warb.

(N. Wallich, 17861854, Danish botanist at Calcutta, India)
Monogr. Myrist. (1897) 305; J. Sinclair l.c. (1958) 405, l.c. (1975) 156; Anderson l.c. 265; W.J. de
Wilde l.c. (1985) 204; Whitmore, Tantra & Sutisna l.c. 267; Turner l.c. 359; Coode et al. (eds.) l.c.
223. Basionym: Myristica wallichii Hook. f. & Thomson, Fl. Ind. (1855) 161. Lectotype: Griffith
s.n., Malacca (K).
Tree up to 30 m tall; bole straight; crown dense, monopodial. Bark shallowly or deeply
longitudinally fissured, dark grey, not flaky; inner bark bright red, pink, reddish brown,
or deep red. Sapwood whitish, pale or dirty yellowish, light brown, brown or red-brown.
Twigs terete or drying somewhat flattened, not ridged, usually conspicuously hollow,
396
towards apex 36(9) mm diameter, tomentose with hairs 0.30.6 mm long, late-glabrescent;
without or with inconspicuous lenticels; older twigs sometimes flaky. Leaf buds 2030
mm long, pubescent with hairs 0.30.6 mm long. Leaves distichous, membranous to
coriaceous, surface not scabrous, not bullate, upper surface drying olivaceous or brown,
glabrous (with minute tomentum remaining on the midrib or not), lower surface early
glabrescent or tomentose with locally persistent hairs (0.3)0.50.8 mm long, always with
scattered brown-black dots and streaks of irregular size; ovate-oblong to oblong-lanceolate,
(14)1940 (4)4.512 cm, base rounded or short-attenuate, apex subobtuse to acuteacuminate; midrib slightly raised above, late-glabrescent; lateral veins (12)1528 pairs,
flat or sunken above; petiole 1.53.5 cm long. Inflorescences usually on older twigs
behind the leaves, tomentose with hairs 0.51 mm long; bract 310 mm long, caducous;
males 45-times branched, 1033 622 cm, peduncle 3070 mm long; females rather
stout, 37 cm long. Flowers glabrous or glabrescent; perianth 34-lobed; pedicel at base
not articulated. Male flowers: pedicel 0.30.6(1) mm long; buds broadly obovoid, 2
2.5(3) 2.42.5(3) mm, base more or less attenuate, glabrous; perianth at anthesis
clefts to 1/32/3; androecium subglobose, subsessile, apex more or less depressed with
a triradiate crack, faintly angular, 1.32 1.52 mm; anthers (12?)1523, sessile, closely
appressed; apical cavity of column up to c. halfway deep. Female flowers: buds ovoidellipsoid, 2.54 mm; ovary glabrous. Fruits 29 per infructescence, ovoid-ellipsoid, 4
6(7) 34.5 cm, glabrous; dry pericarp 1015 mm thick; stalk stout, 46 mm long;
perianth generally persistent. Seeds ellipsoid, with orange aril.
Ecology. In lowland and ridge-top forests, on red soils, granitic sandy soils, loam soils
with coral limestone, up to 500 m altitude. Flowering and fruiting throughout the year.
39. Horsfieldia xanthina Airy Shaw

(Greek, xanthos = yellow; the yellow flowers)
Kew Bull. 1939, no. 10 (1940) 541; Masamune l.c. 302; Anderson l.c. 266; W.J. de Wilde l.c. (1986)
45, l.c. (1996) 381. Type: Richards 1927, Borneo, Sarawak, Marudi district, Mt. Dulit (holotype K;
isotypes A, SING).
Tree up to 30 m tall. Twigs solid, not ridged, subterete, towards apex 2.56(16) mm
diameter, drying dark brown, not contrasting with the drying petiole, tomentose with
hairs c. 0.1 mm long, glabrescent; bark with a tendency to flake or not; lenticels conspicuous
or not. Leaf buds 813 mm long, pubescent with hairs 0.1(0.2) mm long. Leaves distichous,
coriaceous, not bullate, not scabrous, upper surface drying olivaceous to brown, usually
with distinct hair scars, lower surface drying usually with a reddish brown tinge, usually
with distinct hair scars, without blackish dots, early glabrescent; elliptic to oblonglanceolate, 835 3.513 cm, base attenuate to short-rounded, apex subacute to acuteacuminate; midrib slightly raised above; lateral veins 820 pairs, flat or slightly raised
above; petiole 0.71.2 cm long. Inflorescences on older twigs behind the leaves, pubescent
with hairs c. 0.1 mm long; bract caducous; males robust, (2)420 cm long, few-flowered,
peduncle 58 mm or 1520 mm (subsp. macrophylla) long, flowers in loose clusters
397
of 35(8), reflexed or not; females 11.5 or 57 (subsp. macrophylla) cm long, fewflowered. Flowers glabrous; perianth 34-lobed; pedicel glabrescent, towards the base
pubescent with hairs c. 0.1 mm long, at base not articulated. Male flowers: pedicel
rather thick, tapering or not, straight or more or less curved (flowers reflexed), 12.5
mm long; buds more or less obovoid or ellipsoid, 2.52.8 mm long, base somewhat tapering
into the pedicel; perianth at anthesis clefts to 1/2(2/3), not collapsing on drying, lobes
coriaceous 0.40.8(1) mm thick; androecium more or less laterally flattened, almost
broadly obovoid in outline, 11.3 mm long; anthers (3)48, more or less curved, almost
entirely connate, c. 1 mm long, sessile, free apices 0.10.3 mm long; column broad,
solid, at apex with shallow cavity; androphore broad, more or less tapering, (0.1)0.2
0.3 mm long. Female inflorescences and flowers known only in subsp. macrophylla;
ovary glabrous. Fruits 26 per infructescence, more or less ellipsoid, c. 5 cm long, glabrous;
stalk stout, c. 3 mm long; perianth not persistent.
Ecology. In hill mixed dipterocarp to lower montane forests.
Key to subspecies
Twig moderately slender, towards apex 2.54.5(8) mm diameter. Leaves 818 3.57
cm. Male inflorescences 25 cm long; flowers often more or less reflexed; anthers 6....
subsp. xanthina
W.J. de Wilde l.c. (1986) 46; Whitmore, Tantra & Sutisna l.c. 268.
Tree up to 17 m tall. Male flower buds subglobose or broadly ellipsoid, c. 2.5 2.22.5
mm; at anthesis perianth clefts to 1/22/3; pedicel 22.5 mm long; androecium slightly
laterally compressed. Young fruits ellipsoid-obovoid.
Endemic to Sarawak. In kerangas, lower montane and limestone forests, at 8001150
m altitude.
Twig stout, towards apex 3.56(16) mm diameter. Leaves 2235 713 cm. Male
inflorescences 1020 cm long; flowers erect; anthers 78................................
subsp. macrophylla W.J. de Wilde
(Greek, makro = large, phullon = leaf; large-leaved)
l.c. (1986) 47, l.c. (1996) 381; Whitmore, Tantra & Sutisna l.c. 268. Type: Clemens 50050, Borneo,
Sabah, Mt. Kinabalu (holotype K; isotypes BM, L).
Tree up to 30 m tall. Bark of twig not flaky. Bark brown and grey, fissures boat-shaped;
sap light red, watery. Male flowers: pedicel 11.5 mm long; buds broadly ellipsoid, 2.6
2.8 2.52.7 mm; perianth 3-lobed, at anthesis clefts to nearly halfway; androecium c.
1.30.8 mm long, subtriquetrous in transverse section. Female inflorerscences (0.5)18
cm long, once (or twice) branched. Female flowers: pedicel 1.52 mm long, glabrous or
with minute hairs towards the base; buds ovoid-ellipsoid, 44.5 33.5 mm; perianth 3lobed, at anthesis clefts to nearly halfway; ovary c. 2.2 2 mm, glabrous. Fruits glabrous,
c. 5 33.5 cm; pericarp 5(10) mm thick, woody towards inside.
Endemic to Borneo (Sarawak and Mt. Kinabalu in Sabah). In lower montane forest,
ridge forest, on igneous derived (andesitic) soils, at 11001300 m altitude. Flowering
in July and November; fruiting in November.
398
4. KNEMA Lour.
(Greek, knema = the spoke of a wheel; the stamens in the androecium)
Fl. Coch. (1790) 604; Warburg, Monogr. Myrist. (1897) 132, 543; Gamble, J. As. Soc. Beng. 75, 2
(1912) 236; Ridley, FMP 3 (1924) 66; J. Sinclair, Gard. Bull. Sing. 16 (1958) 254, ibid. 18 (1961) 102;
Backer & Bakhuizen f., FJ 1 (1964) 139; Whitmore, TFM 1 (1972) 330; W.J. de Wilde, Blumea 25
(1979) 321, ibid. 27 (1981) 223, ibid. 32 (1987) 115, ibid. 41 (1996) 382, ibid. 43 (1998) 241; Anderson,
CLTS (1980) 266; Corner, WSTM 3rd. edition 2 (1988) 564; Whitmore, Tantra & Sutisna, CLK 2, 1
(1990) 268; Coode et al. (eds.), CLBD (1996) 224; Argent et al. (eds.), MNDT-CK 2 (1997) 452;
PROSEA 5, 3 (1998) 317.
Dioecious trees; stilt roots sometimes present. Twigs towards apex variously pubescent,
glabrescent, smooth or striate, lower down sometimes tending to crack or flake. Leaves
up to 65 cm long, membranous to coriaceous, not brittle when dry; when young covered
with various tomentum, upper surface early glabrescent, lower surface glaucous or greyish,
glabrescent or with various persistent tomentum, dark dots present or absent; lateral veins
interarching at the leaf margin, usually distinct above, always distinct beneath; intercostal
venation reticulate, distinct and prominent above; petiole, in some species with cordate
leaf base, somewhat pulvinate in the apical portion. Inflorescences axillary to leaves
or fallen leaves, consisting of sessile or short-peduncled, simple or 2(3)-fid scar-covered
brachyblasts; bracts caducous; flowers in terminal subumbellate clusters, mostly numerous
in males, few in females. Male flowers pedicelled; bracteole minute, persistent or caducous,
located at the base or at about the middle part of the pedicel; buds ellipsoid, obovoid,
depressed globose or mitriform, at base attenuate, broadly rounded, or more or less saccate,
in section circular or trigonous; perianth 3(5)-lobed, at anthesis clefts to c. halfway to
nearly to the base, lobes spreading; androecium comprising a cylindrical or tapering
staminal column (androphore) of various lengths, and apical, circular, peltate, flat to convex
or mamillate staminal disc (synandrium) to which the anthers are radially attached by
their bases; anthers 325, sessile, or shortly stiped, horizontal or suberect, spaced or
touching each other. Female flowers subsessile or pedicelled, in bud obovoid, or ellipsoidoblong; perianth 3(4)-lobed, at anthesis clefts to c. halfway; ovary ovoid to globose,
variously pubescent; style short or absent, stigma 2many-lobed, each lobe 2severallobulate. Fruits sessile or stalked; pericarp thick, hard and fleshy or more or less woody,
with various tomentum, sometimes early glabrescent. Seeds not pointed at one end; testa
not variegated; aril shallowly laciniate or convoluted at the apex only (to c. 1/3), rarely
completely closed; endosperm with a fixed oil and starch.
Distribution. A genus comprising about 93 species distributed in continental SE Asia and
Malesia, from the Deccan Peninsula (India) to S China (Yunnan), eastwards to the Philippines,
Maluku and W New Guinea. Most species are found in Borneo, of which 44 species are
recorded for Sabah and Sarawak.
Ecology. Mostly under- and middle-storey trees of primary and degraded, usually ever
wet forests, up to 1800(2000) m altitude. In continental SE Asia also occur in seasonal
forest.
399
Key to Knema species

1. Lower leaf surface densely tomentose with persistent, appressed, tightly interwoven
hairs, silky to the touch.........................2
Lower leaf surface either glabrous (glabrescent), or with various tomentum, silky or
not silky to the touch..............4
2. Bark of twig longitudinally cracked and flaky.................4. K. elmeri
Bark of twig not cracked or flaky..............3
3. Tomentum of flowers consists of hairs c. 0.1 mm long. Fruits subglobose to broadly
ovoid, 3.55 cm long. Staminal disc conspicuously convex. Reticulation of intercostal
venation on the upper leaf surface coarse...........................37. K. sericea
Tomentum of flowers very inconspicuous, composed of stellate scale-like hairs c. 0.1
mm long or less. Fruits obovoid-oblong to oblong, 58 cm long. Staminal disc flat.
Reticulation of intercostal venation on the upper leaf surface finer.....................................
.............................................................................................1. K. ashtonii (var. cinnamomea)
4. Bark of twig longitudinally cracked, when older flaky..................5
Bark of twig not cracked or flaky.................13
5. Flowers wholly or largely glabrescent...............6
Flowers with persistent tomentum..............8
6. Fruits tomentose with persistent hairs c. 1 mm long.................32. K. psilantha
Fruits glabrescent..............7
7. Flower buds more or less mitriform (sharply angled), 715 mm long. Twig towards
apex 410 mm diameter, brown......................6. K. galeata (in part)
Flower buds not sharply angled, 3.55 mm long. Twig towards apex 1.53 mm diameter,
often yellowish..............23. K. membranifolia (in part)
8. Tomentum of twigs and flowers comprising hairs (2)34 mm long.....................
......20. K. longepilosa
Tomentum of twigs and flowers comprising hairs 0.12(2.5) mm long........9
9. Twig rather slender, towards apex (1)24(8) mm diameter. Female flower with pedicel
up to 2 mm long..........................10
Twig stout, towards apex 412 mm diameter. Female flower with pedicel 220 mm
long.........11
10. Male flower bud depressed ovoid to depressed globose, 34.5 mm diameter; pedicel
(1.5)23(6) mm long; bracteole apical; staminal disc flat or faintly convex.
.....16. K. latericia
Male flower bud depressed globose, 46 mm diameter; pedicel (4)510 mm long;
bracteole median; staminal disc mammillate........13. K. korthalsii (subsp. rimosa)
11. Male flower bud 610 mm long. Fruits 3.57 cm long; stalk 1030 mm long.....
.....27. K. pallens
400
Male flower bud less than 6 mm long. Fruits 24.5 cm long; stalk 03(5) mm
long......12
12. Leaves chartaceous or thinly coriaceous, upper surface drying brown; leaf base obtuse
or subcordate (or rarely narrowed)...............21. K. lunduensis
Leaves usually coriaceous, upper surface drying olivaceous-brown; leaf base attenuate
to rounded.................30. K. percoriacea
13. Lower leaf surface with minute reddish brown to blackish dots.................14
Lower leaf surface without such dots......................18
14. Twig slender, towards apex 11.5 mm diameter. Leaves glabrescent beneath. Fruits
obovoid-oblong, 1.71.8 cm long.............................25. K. muscosa
Twig stout, towards apex 1.56 mm diameter. Lower leaf surface with persistent or
subpersistent tomentum. Fruits of various shapes, 1.54 cm long...............15
15. Tomentum of lower leaf surface consisting of scattered, minute, stellate hairs c. 0.1
mm long, sometimes with a few larger ones scattered in-between.....15. K. kunstleri
of lower leaf surface consisting of stellate and stellate-dendroid hairs c. 0.21.2 mm
long...........................16
16. Leaves 415 cm long, upper surface drying dark brown. Perianth inside hairy.......
........33. K. pubiflora
Leaves 1036 cm long, upper surface drying olivaceous-brown. Perianth inside
glabrous............17
17. Pedicel of male flower 715 mm long, and that of female flower 58(10) mm long.
Perianth 34-lobed, at base inside without a disc. Fruits usually ellipsoid or obovoid,
(1.5)24 cm long; stalk 510 mm long...............2. K. conferta
Pedicel of male flower 2030 mm long, that of female flower 1012 mm long. Perianth
45-lobed, at the base inside with a disc. Fruits subglobose, usually somewhat broader
than long, 23 2.52.8 cm; stalk 1830 mm long................29. K. pedicellata
18. Fruits glabrescent. Flower buds pubescent or glabrescent. Bark of twigs usually
yellowish or purplish.............................19
Fruits pubescent or glabrescent. Flower buds with persistent tomentum. Bark of twigs
yellow-brown, grey-brown or blackish............................23
19. Twig towards apex 410 mm diameter. Male flower buds 610 mm long, more or
less mitriform ...........6. K. galeata (in part)
Twig towards apex 14.5 mm diameter. Male flower buds 2.57 mm long, depressed
globose, subglobose or ovoid...........................20
20. Twig towards apex at first tomentose with hairs c. 1 mm long, glabrescent...
.....23. K. membranifolia (in part)
Twig towards apex at first tomentose with hairs 0.10.5 mm long..............21
21. Leaves 1730 cm long. Pedicel stout. Fruits 56 cm long; pericarp c. 10 mm
thick.........43. K. viridis
401
Leaves up to 20(25) cm long. Pedicel slender. Fruits 25 cm long; pericarp 12.5

mm thick.........22
22. Tomentum of twig towards apex consisting of hairs 0.20.5 mm long. Fruit stalk
(2)58 mm long.........................19. K. linguiformis
Tomentum of twig towards apex consisting of hairs c. 0.1 mm long or less. Fruit
stalk 816 mm long........3. K. curtisii
23. Tomentum of twig towards apex comprising hairs more than 0.2 mm long................24
Tomentum of twig towards apex comprising hairs 0.10.2 mm long..................32
24. Lower leaf surface with persistent tomentum or late-glabrescent...................25
Lower leaf surface early glabrescent (sometimes with some tomentum persisting on
midrib and veins in K. glomerata)............................27
25. Tomentum of lower leaf surface consisting of more or less uniformly stalked dendroid
hairs. Fruits tomentose with hairs 13 mm long; stalk 01(5) mm long........
......18. K. laurina
Tomentum of lower leaf surface consisting of sessile-stellate and stellate-dendroid
hairs of various sizes. Fruits tomentose with hairs 0.21.5 mm long; stalk 210 mm
long...............26
26. Female flower buds 67.5 mm long. Fruits 23.8 cm long, tomentose with hairs 1
1.5 mm long; stalk 310 mm long. Male flower buds 2.55 mm diameter...
.....26. K. oblongata
Female flower buds 22.5 mm long. Fruits 1.42 cm long, tomentose with hairs 0.2
0.5 mm long; stalk 23 mm long. Male flower buds 1.52.5 mm diameter.......
.........41. K. tridactyla (in part)
27. Leaves above drying brown or blackish. Fruits tomentose with hairs c. 3 mm
long........36. K. rufa
Leaves above drying olivaceous-brown. Tomentum of fruits consists of hairs 0.1
0.5(1) mm long...................28
28. Leaves generally coriaceous; midrib and lateral veins prominently raised above;
reticulation of intercostal venation raised and distinct above.............29
Leaves generally chartaceous; midrib and lateral veins flat or slightly raised above;
reticulation of intercostal venation slightly raised and less distinct above............30
29. Bracteole attached on the pedicel at about halfway. Staminal disc longmammillate.....11. K. intermedia
Bracteole subapically attached on the pedicel. Staminal disc convex or lowmammillate.............42. K. uliginosa
30. Twig rather stout, towards apex 35 mm diameter. Fruits 1.54 cm long, tomentose
with hairs 0.51 mm long; stalk 26 mm long......13. K. korthalsii (subsp. korthalsii)
Twig slender, towards apex 1.52.5(3) mm diameter. Fruits 1.52.5(3) cm long,
tomentose with hairs 0.10.2(0.5) mm long; stalk 46(10) mm long...........31
402
31. Fruit stalk 46(10) mm long; fruits pubescent with scurfy hairs 0.10.2 mm long.
Leaves drying greenish brown above; reticulation of intercostal venation indistinct.
...........9. K. glomerata
Fruit stalk slender, 710 mm long; fruits with hairs up to 0.5 mm long. Upper surface
of leaves drying conspicuously green-olivaceous; reticulation of intercostal venation
distinct....5. K. emmae
32. Fruits largely glabrescent, somewhat flat, often ridged, at base somewhat saccate.
Leaves elliptic-oblong or obovate-oblong, upper surface drying dark brownish to
blackish, lower surface often tinged reddish brown or purplish. Anthers 3 or
4.............................................................................17. K. latifolia
Fruits either with persistent tomentum, or glabrescent and then not saccate at the
base. Leaves of various shapes; upper surface drying olivaceous or brown, lower
surface greyish. Anthers 525 ................33
33. Female flower buds 22.5 mm long. Fruits 1.42 cm long; stalk 23 mm long. Anthers
5 or 6 ......................41. K. tridactyla (in part)
Female flower buds 3.57 mm long. Fruits of various size; stalk usually more than
3 mm long. Anthers 6 or more..................................34
34. Twig stout, towards apex 46 mm diameter. Fruits 68 cm long, with blunt or rounded
apex. Reticulation of intercostal venation on the upper leaf surface very fine, areoles
c. 0.5 mm diameter or less.........................1. K. ashtonii (var. ashtonii)
Twig various, towards apex 16(8) mm diameter. Fruits smaller, if over 6 cm long,
then apex acute, not rounded. Reticulation of intercostal venation on the upper leaf
surface various.....................35
35. Perianth of male flower 12 mm thick at the sutures...............36
Perianth of male flower up to 1 mm thick at the sutures..............37
36. Twig towards apex usually blackish brown and more or less angled by ridges from
the bases of the petioles. Leaves thinly coriaceous; lateral veins 2230 pairs..44. K. woodii
Twig towards apex pale brown or greyish brown, not angled. Leaves coriaceous or
rigidly coriaceous; lateral veins 2550 pairs...................28. K. pectinata
37. Leaf base broadly rounded to cordate. Twig rather stout, towards apex 28 mm diameter.
Leaves up to 50 cm long.............................38
Leaf base attenuate, cuneate to rounded. Twig usually more slender, towards apex
13.5 mm diameter. Leaves up to 30 cm long........................39
38. Twig towards apex 36(8) mm diameter. Leaves (20)3050 (6)1017 cm; lateral
veins flat or slightly raised above. Fruits subglobose to ellipsoid-oblong, (3)46
cm long, apex rounded. Male flower buds pear-shaped.............34. K. pulchra
Twig towards apex 24 mm diameter. Leaves 1230 5.512 cm; lateral veins sunken
to flattish above. Fruits oblong, oblong-lanceolate to almost fusiform, (5)78 cm
long, apex long-acute. Male flower buds depressed globose....14. K. kostermansiana
39. Leaves coriaceous to rigidly coriaceous; intercostal venation on the upper leaf surface
forming a raised and very distinct network...............12. K. kinabaluensis
403
membranous, chartaceous to thinly coriaceous; intercostal venation on the upper leaf

surface distinct or not..............................40
40. Twig towards apex usually somewhat flat, drying yellow-brown. Intercostal venation
on the upper leaf surface forming a distinct very fine network, the areoles less than
0.5 mm diameter..............22. K. luteola
Twig towards apex terete or somewhat angular, drying brown to blackish. Intercostal
venation on the upper leaf surface generally coarser, distinct or not distinct...............41
41. Fruits glabrescent. Bracteole usually persistent. Lower leaf surface subglabrous, early
glabrescent.........7. K. glauca
Fruits with persistent tomentum. Bracteole persistent or caducous. Tomentum of lower
leaf surface persistent or glabrescent.......................42
42. Male flower buds pear-shaped, much tapered in the lower half......31. K. piriformis
Male flower buds obovoid or globose or subglobose...................43
43. Lower leaf surface early glabrescent. Style distinct, 0.51 mm long. Male flower
buds obovoid......39. K. stylosa
Lower leaf surface with persistent tomentum or late-glabrescent. Style up to 0.5 mm
long. Male flower buds obovoid or globose or subglobose.............................44
44. Tomentum of lower leaf surface consisting of dense, touching or interwoven hairs.......45
Tomentum of lower leaf surface comprising scattered hairs, not touching each other........46
45. Hairs of plant parts brown. Lateral veins usually raised above. Fruits 1.82.2 cm
long.......8. K. glaucescens
Hairs of plant parts yellowish or greyish white. Lateral veins flat or partially raised
above. Fruits c. (3)4 cm long............................35. K. riangensis
46. Twig slender, towards apex c. 1 mm diameter. Leaves up to 18 cm long, lower surface
with very inconspicuous tomentum of scattered scale-like stellate hairs of less than
0.1 mm long. Fruit 11.5 cm long, stalk 1020 mm long.....
.......38. K. stenophylla (subsp. longipedicellata)
Twig towards apex generally stouter, 13.5 mm diameter. Leaves up to 25 cm long,
lower surface tomentose with hairs 0.10.3 mm long. Fruit 1.53.5 cm long, stalk
up to 10(15) mm long..................47
47. Twig slender, towards apex 11.5(2) mm diameter. Reticulation of intercostal venation
on the upper leaf surface rather faint. Anthers 68. Fruit 1.52 cm
long.......24. K. mogeana
Twig towards apex generally stouter, 1.53 mm diameter. Reticulation of intercostal
venation on the upper leaf surface distinct. Anthers (7)913(15). Fruit 2 cm long
or more...............48
48. Twig towards apex 1.52(2.5) mm diameter. Leaves 920 cm long. Staminal column
(almost) glabrous. Fruit 22.7 cm long..........40. K. subhirtella
Twig towards apex 23.5 mm diameter. Leaves (7)1023 cm long. Staminal column
pubescent towards the base. Fruit 2.53.5 cm long......10. K. hirtella
404
1. Knema ashtonii J. Sinclair

(P.S. Ashton, 1934, sometime forest botanist at the Forestry Departments of Brunei
Darussalam and Sarawak; Professor of Dendrology, Harvard University, U.S.A.)
Gard. Bull. Sing. 18 (1961) 162 (excl. male flowers); W.J. de Wilde, Blumea 25 (1979) 384, ibid. 41
(1996) 382; Anderson l.c. 266; Whitmore, Tantra & Sutisna l.c. 268; Argent et al. (eds.) l.c. 454. Type:
Smythies, Wood & Ashton SAN 17386, Brunei, Temburong district (holotype K; isotypes A, BO, BRI,
BRUN, KEP, L, SAN, SING).
Tree up to 25 m tall. Twigs stout, towards apex ridged, 46 mm diameter, tomentose

with hairs c. 0.1 mm long, glabrescent; bark drying brown or blackish, not cracked or
flaked. Leaves thinly coriaceous, upper surface drying olivaceous, lower surface not
papillose, without brownish or blackish dots, glabrous or sparsely or densely greyish
or cinnamon tomentose with silky hairs 0.10.2 mm long; obovate-oblong to oblonglanceolate, 2555 715 cm, base subcordate to attenuate; midrib raised above; lateral
veins 2040 pairs, flat or depressed above; intercostal venation more or less trabeculate,
very fine on the upper surface; petiole 13 cm long. Inflorescences up to 1 0.5 cm;
males 410-flowered; flowers pubescent with minute stellate scale-like hairs less than
0.1 mm long. Male flowers: buds obovoid, 5.58 mm long; perianth at anthesis clefts
to c. 4/5; staminal disc flat; anthers 6 or more; bracteole inserted on the apical part
of the pedicel, caducous. Female flowers: buds 56 4 mm; pedicel 56 mm long.
Fruits 13 per infructescence, obovoid-oblong, oblong to long-ellipsoid, 58 34 cm,
with blunt or rounded apex, tomentose with persistent hairs c. 0.2 mm long; dry pericarp
c. 4 mm thick; stalk 1030 mm long, the bracteole scars situated about halfway to near
the apex; collar-like perianth scar present at the base of the fruit.
Distribution. Endemic to Borneo (Sabah, Sarawak, Brunei and Kalimantan).
Ecology. In primary mixed dipterocarp forest, often by streams, up to 600 m altitude.
Flowering in March to April; fruiting throughout the year.
Key to varieties
Lower leaf surface (nearly)glabrous or with scattered grey scaly hairs not silky to the
touch. Fruit stalk 1015(20) mm long; bracteole scars attached to the pedicel usually
above the middle. Pedicel of male flower c. 9 mm long.................
var. ashtonii
W.J. de Wilde l.c. (1979) 386, l.c. (1996) 382; Coode et al. (eds.) l.c. 224.
Bark smooth, ochre. Stilt roots occasionally present. Male flower buds c. 5.5 4.55
mm; pedicel c. 9 mm long; staminal column c. 2 mm long, staminal disc faintly convex,
including anthers c. 3 mm diameter. Fruits (5)67.5(8) cm.
Sabah, Sarawak and Brunei.
Lower leaf surface densely brownish grey to cinnamon-brown pubescent, silky to the
touch. Fruit stalk 2030 mm long; bracteole scar attached to the pedicel usually at about
the middle. Pedicel of male flower 2025 mm long.................
var. cinnamomea W.J. de Wilde
(Latin, cinnamomeus = cinnamon-coloured; the indumentum of lower leaf surface)
l.c. (1979) 386. Type: Suib S. 23449, Borneo, Sarawak (holotype L; isotypes K, SAR, SING).
Bark smooth, reddish to greyish brown, without furrows or flakes. Male flower buds
405
78 6 mm; pedicel 2025 mm long; staminal column tapering, 1.52 mm long,

staminal disc subtriangular, flat, including anthers c. 3.5 mm diameter. Fruits 58
cm long.
Sarawak and Kalimantan. Often found by streams on sandy, basalt-derived, and clayrich soils, at altitudes up to 600 m.
2. Knema conferta (King) Warb.

(Latin, confertus = pressed together; male flowers densely crowded)
Monogr. Myrist. (1897) 578; Gamble l.c. 243; Merrill l.c. (1921) 270, l.c. (1929) 74; Ridley l.c. 70;
Masamune l.c. 302; J. Sinclair l.c. (1958) 286, l.c. (1961) 194; W.J. de Wilde l.c. (1979) 472; Anderson
l.c. 266; Whitmore, Tantra & Sutisna l.c. 268; Turner, Gard. Bull. Sing. 47 (1995) 360; Argent et al.
(eds.) l.c. 455. Basionym: Myristica conferta King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 315. Lectotype
(W.J. de Wilde, 1979): Kings Collector 6211, Peninsular Malaysia, Perak, Larut (hololectotype CAL;
isolectotypes FI, G, K, L).
Tree up to 25 m tall; bole straight, occasionally with buttresses up to 3 m high, extending

over the ground c. 30 cm wide, c. 10 cm thick. Bark smooth or somewhat rough, peeling
off in narrow strips or in isodiametric scales 11.5 mm thick; inner bark 58 mm thick,
light red or red-brown. Sapwood whitish, or yellowish, or reddish brown; heartwood
reddish brown. Twigs stout, towards apex 25 mm diameter, densely tomentose with hairs
0.21 mm long, glabrescent; bark not tending to crack or flake. Leaves subcoriaceous,
upper surface drying olivaceous-brown, lower surface papillose, with scattered blackish
dots especially on the veinlets, persistently tomentose with stellate and stellate-dendroid
hairs (0.2)0.31 mm long, not silky to the touch; oblong to lanceolate, 1036 3
12 cm, base subcordate to cuneate, apex sometimes subobtuse; midrib raised above; lateral
veins 1228 pairs, more or less raised above; petiole 0.51.5 cm long. Inflorescences
simple or 2(3)-furcate, up to 1.5 cm diameter; (5)1040-flowered in males; flowers
tomentose with hairs (0.1)0.20.3 mm long. Male flowers: pedicel 715 mm long;
bracteole caducous, attached to the pedicel at or above the middle; buds subglobose,
more or less trigonous, 44.5 mm diameter, inside glabrous, greenish to yellowish; perianth
34-lobed, at anthesis clefts to the base, at base inside without a disc; staminal disc
more or less flat, including anthers 23 mm diameter; anthers (10)1217, shortly stiped,
horizontal; staminal column 0.81.5 mm long. Female flowers: pedicel 58(10) mm
long; buds obovoid, c. 5 mm long. Fruits 15 per infructescence, ellipsoid or obovoid,
1.54 13 cm, tomentose with hairs c. 0.5 mm long, late-glabrescent; dry pericarp
25 mm thick; stalk 510 mm long.
Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo (Sabah, Sarawak and
Kalimantan).
Ecology. In well-drained as well as swampy forests, up to 600 m altitude. Flowering
throughout the year; fruiting mainly in November to March.
3. Knema curtisii (King) Warb.

(Ch. Curtis, 18521928; plant collector; 18841903, superintendent of the Penang Botanic
Garden)
406
Monogr. Myrist. (1897) 567; Gamble l.c. 240; Ridley l.c. 69; J. Sinclair l.c. (1958) 281, l.c. (1961) 196
(excl. var. linguiformis); W.J. de Wilde l.c. (1979) 416; Anderson l.c. 266; Whitmore, Tantra & Sutisna
l.c. 268; Argent et al. (eds.) l.c. 455. Basionym: Myristica curtisii King l.c. 326. Lectotype: Scortechini
292, Peninsular Malaysia, Perak (hololectotype CAL; isolectotypes FI, G, K, L, SING).
Shrub or tree up to 35 m tall. Twigs slender, towards apex (0.5)12 mm diameter, yellowish,
tomentose with greyish hairs up to 0.1 mm long, glabrescent; bark not cracked or flaked.
Leaves membranous or chartaceous, upper surface drying greenish or brown, lower surface
finely papillose, without brownish dots, at first densely tomentose with hairs c. 0.1 mm
long, early glabrescent; elliptic or obovate or lanceolate, 3.515(20) 1.24(6.5) cm,
base mostly attenuate, apex sometimes blunt or rounded; midrib raised above; lateral veins
(6)814(20) pairs, slightly raised above; petiole 0.52 cm long. Inflorescences up to
0.5 cm diameter, (2)515-flowered in males; flowers tomentose with persistent hairs
0.10.3 mm long, or partially glabrescent. Male flowers: pedicel 511 mm long; bracteole
subpersistent, attached on the pedicel at (or below or above) the middle; buds subglobose,
often depressed, angular or not, 2.55(5.5) mm diameter, inside cream or pink; perianth
at anthesis clefts to 2/34/5; staminal disc flat or with minute mammilla, subtriangular,
including anthers 1.73 mm diameter; anthers 925(30), sessile, closely appressed;
staminal column 12 mm long. Female flowers: buds obovoid, 45 mm long; pedicel
510 mm long. Fruits 12 per infructescence, ellipsoid, 25 1.53 cm, apex acute,
or narrowed at both ends, tomentose with hairs 0.10.2 mm long, or glabrescent; dry
pericarp 12.5 mm thick; stalk 816 mm long.
Distribution. Sumatra, Peninsular Malaysia and Borneo.
Notes. A complex species comprising 4 varieties with considerable differences in general
habit, distribution and habitats. Three of the varieties occur in Sabah and Sarawak. Variety
amoena J. Sinclair is endemic to Brunei.
Key to varieties
1.
Leaves 515(20) cm long, upper surface drying greenish, apex acute or acuminate;
lateral veins distinct on both sides. Male flower buds tending to become glabrescent.
Anthers 915...............................................
var. curtisii
J. Sinclair, Gard. Bull. Sing. 18 (1961) 196; W.J. de Wilde l.c. (1979) 418, l.c. (1996) 383;
Turner l.c. 360; Coode et al. (eds.) l.c. 224.
Stem without stilt roots, sometimes with buttresses up to 50 cm high. Leaves

membranous, sometimes more or less parallel-sided; lateral veins (6)816(18)
pairs, intercostal venation distinct on both surfaces. Male flower buds in cross
section obtuse-angular, 2.54 mm diameter, initially tomentose with mixed sessile
stellate and dendroid hairs 0.10.3 mm long, glabrescent; perianth inside reddish.
Fruits scurfy, 24.5 cm long, usually ridged; stalk 816 mm long.
Sumatra, Peninsular Malaysia and Borneo (Sabah, Sarawak, Brunei and Kalimantan).
In primary and secondary forests, on hill slopes, on a variety of soils, up to
1000 m altitude. Flowering and fruiting throughout the year.
Leaves 3.510 cm long, upper surface drying greenish to dark brown; apex acuminate
or rounded; lateral veins indistinct. Tomentum of flowers persistent or partially
glabrescent. Anthers 1825(30)....................................2
407
2.
Male flower buds with persistent tomentum of mixed sessile-stellate and dendroid
hairs; angles rounded. Leaves drying greenish brown above; apex acute to acuminate......
var. arenosa J. Sinclair
(Latin, arenosum = sandy place; its natural habitat)
l.c. (1961) 198, f. 6D; W.J. de Wilde l.c. (1979) 420. Type: Kostermans 8653, Borneo,
Kalimantan, Nunukan Is. (holotype K; isotypes BM, BO, L, P, SING).
Tree up to 35 m tall, without stilt roots. Leaves membranous or chartaceous,

3.513 cm long; base cuneate to attenuate, apex rarely somewhat obtuse; lateral
veins 1218 pairs, indistinct on both surfaces. Male flower buds in cross section
rounded-angular, 35 mm diameter, initially tomentose with mixed stellate and
dendroid hairs 0.10.3 mm long, glabrescent; anthers 1823.
Sumatra and Borneo (Sarawak and Kalimantan). In ridge forest on sandy, sandy
loam or sandstone-derived soils, up to 600 m altitude. Flowering throughout the
year.
Male flower buds with tomentum of minute mainly sessile stellate hairs, tending to
be partly shed or easily rubbed off; angles sharp. Leaves drying dark brown above;
apex rounded or subacuminate with obtuse tip....................
var. paludosa J. Sinclair
(Latin, paludosus = swampy place; its natural habitat)
l.c. (1961) 201; W.J. de Wilde l.c. (1979) 420; Turner l.c. 360. Type: Corner SFN 26155,
Singapore, Jurong (holotype K; isotypes BO, CAL, SING).
Stilt roots usually present. Leaves usually chartaceous, 410 cm long; base cuneate,
apex either subacute with blunt tip, or obtuse or rounded; lateral veins 616 pairs,
faint on both sufaces. Male flower buds in cross section sharply angular, 35
mm diameter; initially tomentose with mixed sessile-stellate and dendroid hairs
0.10.2 mm long, partially tending to become glabrescent; anthers 2030. Fruits
ridged on the line of suture, 45 cm long; stalk c. 10 mm long.
Sumatra, Peninsular Malaysia, Singapore, and Borneo (Sarawak). Mainly in freshwater
swamp forest, up to 200 m altitude. Flowering in JuneOctober.
4. Knema elmeri Merr.

(A.D.E. Elmer, 18701942, plant collector in Sabah and the Philippines)
PEB (1929) 75; Masamune l.c. 302; J. Sinclair l.c. (1961) 202; W.J. de Wilde l.c. (1979) 383; Anderson
l.c. 266; Whitmore, Tantra & Sutisna l.c. 268; Coode et al. (eds.) l.c. 224. Type: Elmer 21527, Borneo,
Sabah, Tawau district (holotype UC; isotypes A, BM, BO, BR, C, G, K, L, M, NY, P, SING, U).
Tree up to 15 m tall, sometimes with stilt roots. Bark smooth or scaly or flaky, or cracked,
hard; inner bark white, or reddish, or red-brown. Sapwood brown. Twigs towards apex
more or less angled, 24(5) mm diameter, initially tomentose with hairs 0.10.2 mm
long, glabrescent; bark cracked and flaky. Leaves chartaceous, upper surface drying
greenish, lower surface densely tomentose with silvery to pale brown, persistent short
hairs 0.10.2 mm long, silky to the touch; elliptic to oblong, 1232 413 cm, base
rounded to acute; midrib sunken to prominent above; lateral veins 1220(25) pairs, raised
or flat above; petiole 0.81.8 cm long. Inflorescences sessile, up to 0.8 cm diameter,
525-flowered in males; flowers tomentose with dense hairs c. 0.1 mm long. Male flowers:
pedicel 714 mm long; bracteole subpersistent, attached to the pedicel at about the middle;
408
buds ovoid to depressed globose, often more or less sagged at the base, 34 44.5
mm, inside red or purple; perianth at sutures 0.50.8 mm thick, at anthesis clefts nearly
to the base; staminal disc, including anthers circular, convex in the centre, 22.5 mm
diameter; anthers 1014, sessile to stalked; staminal column 0.50.8 mm long, clasped
by the thickened perianth base. Female flowers: pedicel 12 mm long; bracteole attached
to the pedicel at about the middle; buds ellipsoid, c. 6 mm long. Fruits 16 per
infructescence, obovoid to ellipsoid-oblong, 22.5 1.11.6 cm, tomentose with hairs
0.10.2 mm long; dry pericarp at suture c. 2 mm thick; stalk 15 mm long.
Ecology. In primary and disturbed riverine and mixed dipterocarp forests, lower montane
forest, on clay-rich soils, sandy clay soils, also on limestone, up to 1500 m altitude.
5. Knema emmae W.J. de Wilde

(Emma van Nieuwkoop, sometime desk editor at the Rijkherbarium, Leiden, the Netherlands)
Blumea 41 (1996) 384. Type: Amdjah 144, Borneo, NE Kalimantan, G. Muru Tagal (holotype L; isotypes
BO, K, SING).
Treelet up to 5 m tall. Bark dark brownish; inner bark brownish; sap red. Sapwood
whitish. Twigs terete, towards apex 1.52 mm diameter, initially tomentose with hairs
0.5(1) mm long, glabrescent; bark lower down yellowish brown, not cracked or flaky;
lenticels minute. Terminal leaf buds c. 10 mm long, tomentose with hairs c. 0.5 mm
long. Leaves chartaceous, upper surface drying green-olivaceous, lower surface glabrous,
without blackish dots, not or only slightly papillose; elliptic-oblong or oblong-lanceolate,
1220 3.57 cm, base short-attenuate, apex somewhat blunt or acute-acuminate; midrib
raised above; lateral veins 1215 pairs, flat or slightly raised above; intercostal venation
reticulate, distinct; petiole c. 1 cm long. Inflorescences 0.10.2 cm diameter, 25-flowered;
peduncle to 1.5 mm long; flowers tomentose with persistent hairs (0.2)0.50.7 mm long.
Male flowers: pedicel 67 mm long; bracteole minute, subpersistent, inserted on the pedicel
slightly above the middle; buds depressed globose, c. 3 4 mm; perianth at sutures
c. 0.5 mm thick, at anthesis clefts to c. 3/4(4/5); staminal disc circular, flat, c. 2 mm
diameter; anthers 1415, small, sessile just below the rim of the disc; staminal column
0.50.8 mm long. Female flowers: pedicel c. 4 mm long; buds obovoid, c. 3.5 mm long.
Fruits 24 per infructescence, ovoid-oblong, 2.53 1.51.6 cm, apex with style remnant
c. 1 mm long; dry pericarp 0.5(1) mm thick; pubescent with hairs c. 0.5 mm long;
stalk 710 mm long, with scar of bracteole at about the middle.
Distribution. Endemic to Borneo (Sabah and NE Kalimantan).
Ecology. In hill mixed dipterocarp forest, at low altitude. Flowering and fruiting in June.
6. Knema galeata J. Sinclair

(Latin, galeatus = provided with a helmet; the helmet-shaped flower buds)
409
Gard. Bull. Sing. 18 (1961) 211; W.J. de Wilde l.c. (1979) 422; Anderson l.c. 267; Whitmore,
Tantra & Sutisna l.c. 268; Coode et al. (eds.) l.c. 224. Type: Anderson (Md. Hasan) S. 4855, Borneo,
Brunei, Berakas FR (holotype SAR; isotypes BRUN, KEP, SING).
Tree up to 30 m tall. Bark rather smooth, finely flaky or fissured; inner bark brownish.
Sapwood pink-yellow or pale brown, or whitish. Twigs towards apex 410 mm diameter,
yellowish, purplish or brown, at first rusty tomentose with hairs 12 mm long, glabrescent;
older bark brown, frequently tending to crack or flake. Leaves coriaceous, upper surface
drying olivaceous, glossy, lower surface early glabrescent, without blackish dots, greyish;
oblong to lanceolate, 1550 2.514 cm, base usually rounded; midrib raised above;
lateral veins 2035 pairs, raised above; petiole 1.23(4) cm long. Inflorescences borne
on older twigs 12 cm diameter, simple or with up to 10 protuberances, up to 2 1
cm, (5)1050-flowered in males; flowers at first tomentose with hairs 0.41 mm long,
glabrescent. Male flowers: pedicel (15)2030 mm long; bracteole subpersistent, inserted
on the pedicel at about the middle; buds mitriform, triquetrous in cross-section, 610
915 mm, inside white, pink or red; perianth lobes broad, reniform, at anthesis clefts
nearly to the base; staminal disc flat or more or less convex, including anthers subtriangular,
35 mm diameter; anthers 1423; staminal column 0.51 mm long. Female flowers:
pedicel 1225 mm long; buds subovoid, mitriform, 79 mm long. Fruits (1)25 per
infructescence, pear-shaped and often contracted towards the base, 25 1.53 cm, at
first tomentose with hairs c. 0.5 mm long, glabrescent; dry pericarp 23 mm thick; stalk
1530 mm long.
Ecology. In primary lowland mixed dipterocarp forest, open kerangas forest and swamp
forest, often on sandy, yellow podsolic, yellow clay, and sandy loam soils. Flowering
7. Knema glauca (Blume) Petermann

(Latin, glaucus = sea-green; the lower leaf surface)
Pflanzenreich (183845) 294; Warburg, Monogr. Myrist. (1897) 594 (for the var. typica only); Merrill
l.c. (1921) 270; Masamune l.c. 302; W.J. de Wilde l.c. (1979) 452, l.c. (1987) 126; Whitmore, Tantra &
Sutisna l.c. 268; Turner l.c. 360; Argent et al. (eds.) l.c. 455. Basionym: Myristica glauca Blume, Cat.
(1823) 111; King l.c. 320, p.p. Type: Blume s.n., W Java (holotype L; isotype BO). Synonyms: K.
glaucescens Jack var. glaucescens auct. non Jack: J. Sinclair l.c. (1958) 304, p.p.; K. cinerea (Poir.)
Warb. var. sumatrana (Blume) J. Sinclair l.c. (1961) 185, Backer & Bakhuizen f. l.c. 140, Whitmore,
TFM 1 (1972) 334.
Tree up to 30 m tall; stilt roots sometimes present, up to 1 m high. Bark smooth, or

flaky or scaly, or sometimes cracked or peeling off in strips c. 10 mm wide; inner
bark 510 mm thick, pinkish or red-brown; sap copious, pinkish. Sapwood whitish
or cream; heartwood reddish. Twigs towards apex terete or somewhat angular, drying
brown or blackish, 13.5 mm diameter, tomentose with minute hairs up to 0.1 mm
long, early glabrescent; bark not tending to crack or flake. Leaves membranous to
chartaceous, upper surface drying greenish to dark brown, lower surface glaucous or
greyish, without dark dots, at first with scattered weak greyish hairs up to 0.1 mm
410
long, early glabrescent; elliptic, lanceolate, to oblong-lanceolate, 620(25) 25.5(

11) cm, base subattenuate, rounded or subcordate; midrib more or less raised above;
lateral veins 1222 pairs, more or less raised above; intercostal venation coarse above;
petiole 0.72 cm long. Inflorescences sessile or with peduncle up to 3 mm long, simple
or 23-fid, up to 1.5 cm diameter; (2)520-flowered in males; flowers pubescent with
persistent scale-like hairs up to 0.1 mm long; perianth at sutures up to 1 mm thick.
Male flowers: pedicel 410 mm long; bracteole usually persistent, inserted on the pedicel
at or above the middle; buds subglobose, 2.53.5 mm diameter, inside reddish; perianth
at anthesis clefts to c. 4/5; staminal disc flat or faintly convex, including anthers circular,
1.52(2.5) mm diameter; anthers (8)1115, subsessile or stiped; staminal column 0.5
1 mm long. Female flower: pedicel 16 mm long; buds more or less ellipsoid, 4
6 mm long; perianth at anthesis clefts to about halfway. Fruits 14 per infructescence,
subglobose or ellipsoid, at base not saccate, 1.83(4?) 1.41.8(2.5?) cm, at first
pubescent with scale-like hairs up 0.1 mm long, early glabrescent, leaving a finely
granulate surface; dry pericarp 1.52.5 mm thick; stalk 312 mm long.
Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, Borneo, Java, and Bali.
Ecology. In primary and degraded lowland to lower montane evergreen forests on a great
variety of soils, up to 1500 m altitude. Flowering and fruiting throughout the year.
Key to varieties
Leaves membranous or chartaceous, elliptic to lanceolate. Fruit stalk 37 mm long; fruit
drying greyish brown.....................
var. glauca
Widespread, occurring in S Thailand, Sumatra, Peninsular Malaysia, Java, Bali, and
Borneo.
Leaves membranous, oblong to lanceolate. Fruit stalk 812 mm long; fruit drying bright
brown..............
var. riparia W.J. de Wilde
(Latin, riparia = frequenting riversides)
Blumea 32 (1987) 126. Type: Chai S. 18949, Borneo, Sarawak (holotype L; isotype SAR).
Tree up to 12 m. Leaves 1122 25.5 cm. Inflorescences simple or forked, up

to 0.4 cm long, sessile or with peduncle of less than 1 mm long, in males 26(
10)-flowered. Male flower: pedicel 810 mm long; buds subglobose, 3(3.5) mm
diameter; perianth at anthesis clefts to c. 5/6; anthers 913, distinctly stiped, well
spaced. Fruits solitary or 2 together, ellipsoid, 22.5 cm long, apex sometimes somewhat
acute.
Endemic to Borneo (Sarawak). Mainly in primary riverine forest, on steep slopes,
up to 500 m altitude. Vernacular name: kumpang ensluai (Iban).
8. Knema glaucescens Jack

(Latin, glaucescens = becoming somewhat sea-green; the lower leaf surface)
Mal. Misc. 7 (1821) 35; Warburg l.c. (1897) 616; J. Sinclair l.c. (1958) 302, p.p.; W.J. de Wilde l.c.
(1979) 462; Whitmore, Tantra & Sutisna l.c. 269; Turner l.c. 360; Coode et al. (eds.) l.c. 224; Argent et
411
al. (eds.) l.c. 455. Type: Jack s.n., Sumatra, Bengkulen (L). Synonyms: Myristica glaucescens (Jack)
Hook. f. & Thomson, Fl. Ind. 1 (1855) 157, Fl. Brit. Ind. 5 (1889) 111 (for the type only); M. geminata
Miq., Fl. Ind. Bat., Suppl. 1 (1861) 385; K. geminata (Miq.) Warb. l.c. (1897) 604, p.p., Gamble l.c.
(1912) 247, p.p., Ridley l.c. 72, p.p., Merrill l.c. (1921) 270, Masamune l.c. 302; K. cinerea (Poir.)
Warb. var. sumatrana auct. non Blume: J. Sinclair l.c. (1961) 185, p.p.
Tree up to 15 m tall. Bark ochrebrown, nearly smooth, not or only slightly flaky. Twigs
towards apex terete or somewhat angular, 12 mm diameter, tomentose with hairs c.
0.1 mm long, glabrescent; bark drying brown or blackish, not tending to crack or flake.
Leaves chartaceous or subcoriaceous, upper surface drying greenish to brown, lower
surface glaucescent or greyish, without blackish dots, subpersistently dense tomentose
with touching or interwoven hairs 0.10.2 mm long mixed with larger stellate-dendroid
hairs, when shed leaving scars; ovate to lanceolate, 620 26.5(7) cm, base rounded
to subattenuate; midrib flat to raised above; lateral veins 1320 pairs, usually raised
above; intercostal venation coarse above; petiole 0.51.5 cm long. Inflorescences up
to 0.5 cm diameter; peduncle up to 2 mm long; 520-flowered in males; flowers tomentose
with peristent hairs 0.10.2 mm long. Male flowers: pedicel 611 mm long; bracteole
early to late-caducous, inserted on the pedicel at or slightly above the middle; buds globose,
2.53.5 mm diameter, inside cream to yellowish; perianth at sutures 0.81 mm thick,
at anthesis clefts to c. 4/5; staminal disc flat to low-mammillate, including anthers
subcircular, 1.52 mm diameter; anthers 913(15), stiped; staminal column 0.51 mm
long. Female flowers: pedicel 36 mm long; bracteole inserted on the pedicel at or above
the middle; buds ellipsoid-obovoid, 45(5.5) mm long; style up to 0.5 mm long. Fruits
1 or 2 per infructescence, ellipsoid to obovoid, 1.82.2 1.21.8 cm, sometimes ridged,
mealy pubescent with persistent hairs 0.10.3 mm long; dry pericarp c. 1 mm thick; stalk
(3)411 mm long.
Ecology. In lowland and hill forests on sandy, clayey and granite-derived soils, up to
500 m altitude. Flowering and fruiting throughout the year.
9. Knema glomerata (Blanco) Merr.

(Latin, glomeratus = clustered closely together into a head; the flowers)
J. Str. Br. Roy. As. Soc. 76 (1917) 81, Sp. Blanc. (1918) 151, l.c. (1921) 270, Enum. Philip. Flow. Pl. 2
(1923) 183; Masamune l.c. 302; J. Sinclair l.c. (1961) 219 (p.p., excl. syn. K. stenocarpa Warb.); W.J.
de Wilde l.c. (1979) 437, l.c. (1987) 125; Whitmore, Tantra & Sutisna l.c. 269. Basionym: Sterculia
glomerata Blanco, Fl. Filip. (1837) 764; Merrill, Publ. Gov. Lab. Philip. n. 27 (1905) 24, 73. Type:
Merrill Spec. Blanc. 504, the Philippines, Luzon, Batangas Prov. (holotype A; isotypes BM, BO, CAL,
K, L, NY, NSW, P, US). Synonyms: Sterculia decandra Blanco, Fl. Filip. (1837) 766; Myristica
heterophylla Fern.-Vill., Novis. Append. (1880) 178; K. heterophylla (Fern.-Vill.) Warb. l.c. (1897)
573; K. heterophylla (Fern.-Vill.) Warb. var. pubescens Warb. l.c. (1897) 576; K. vidalii Warb. l.c.
(1897) 615; K. gitingensis Elmer, Leafl. Philip. Bot. 3 (1911) 1065; K. acuminata Merr., Philip. J. Sc. 17
(1920) 256.
Tree up to 20 m tall. Twigs slender, towards apex 1.52.5(3) mm diameter, tomentose

with woolly hairs 0.20.5 mm long, glabrescent; bark drying brown or blackish, not tending
412
to crack or flake. Leaves chartaceous, upper surface drying greenish brown, lower surface
at first tomentose with greyish soft stellate hairs 0.10.3 mm long, glabrescent, often
the tomentum somewhat long persistent on midrib and lateral veins, without blackish
dots; elliptic to lanceolate, 1023(35) 37(10) cm, base rounded to attenuate; midrib
flattish to slightly raised above; lateral veins (12)1520(22) pairs, flattish above, often
brownish on lower leaf surface; intercostal venation reticulate, faint above, fine but visible
beneath; petiole 0.51.5 cm long. Inflorescences sessile, up to 0.8 0.6 cm; 420-flowered
in males; flowers greyish to rusty tomentose with persistent hairs 0.20.5 mm long. Male
flowers: pedicel 411 mm long; bracteole caducous, minute, inserted on the pedicel at
about the middle; buds depressed globose, 35 mm diameter, inside pink or red; perianth
at sutures 0.30.7 mm thick, at anthesis clefts nearly to the base; staminal disc flattish
or with a low mammilla, including anthers circular, 22.5 mm diameter; anthers 913
(15), almost sessile to stiped; staminal column slender, 12 mm long. Female flowers:
pedicel 24 mm long; bracteole almost median on the pedicel; buds ellipsoid-obovoid,
c. 5 mm long. Fruits 13 per infructescence, globose-ellipsoid, 1.52.5(3) 1.22.3
cm, pubescent with scurfy hairs 0.10.2 mm long; dry pericarp c.1.5 mm thick; stalk
46(10) mm long.
Distribution. Borneo (one collection, S. 24419, from Sarawak), the Philippines (common)
and Seram (rare).
Ecology. In primary and degraded lowland and hill forests, up to 600 m altitude. Flowering
throughout the year; fruiting in JuneDecember.
10. Knema hirtella W.J. de Wilde

(Latin, hirtellus = finely hairy; the staminal column)
Blumea 25 (1979) 459, l.c. (1987) 130; Whitmore, Tantra & Sutisna l.c. 269. Type: Mikil SAN 30235,
Borneo, Sabah, Beaufort district (holotype L; isotype SAN).
Tree up to 25 m tall. Twigs towards apex 23.5 mm diameter, tomentose with scurfy hairs c.
0.1 mm (sometimes with a few emergents up to 0.3 mm) long, early glabrescent; bark drying
brownish or blackish, not tending to crack or flake. Leaves chartaceous, upper surface drying
olivaceous to brown, lower surface without blackish dots, either with persistent tomentum
comprising scattered hairs c. 0.1 mm long or at first tomentose with weak greyish scattered
hairs 0.10.2(0.3) mm long, mixed with scars of shed coarser hairs, glabrescent; oblonglanceolate, (7)1023 2.58.5 cm, base rounded to attenuate, apex acute-acuminate or
subobtuse; midrib faintly raised above; lateral veins 1022 pairs, flattish above; intercostal
venation reticulate, distinct above; petiole 0.81.2 cm long. Inflorescences up to 0.5 cm
diameter; 26(10)-flowered in males; flowers pubescent with persistent hairs up to 0.1 mm
long. Male flowers (only known in var. hirtella): pedicel 4.57 mm long; bracteole caducous
or persistent, inserted on the pedicel at 12 mm below the perianth; buds subglobose, 33.5
mm diameter, inside yellowish (?); perianth at sutures 0.50.8 mm thick, at anthesis clefts to
4/54/6; staminal disc more or less flat, including anthers circular, 1.51.7 mm diameter;
anthers 913(15); staminal column 11.5 mm long (in var. hirtella at base minutely hairy).
Female flowers: pedicel 47.5 mm long; buds long-obovoid, 4.56 mm long; style up to 0.5
mm long. Fruits 1 or 2 per infructescence, ellipsoid or ovoid, 2.53(3.5) 1.72.6 cm, apex
413
obtuse to acute, often with persistent style, persistently pubescent with hairs up to 0.5 mm
long; dry pericarp c. 1.5(2) mm thick; stalk 410(15) mm long.
Ecology. In lowland and hill mixed dipterocarp forests, often on slopes and ridges, on
sandy, sandstone-derived, lateritic, and clayey soils, up to 1000 m altitude. Flowering
in MayJuly; fruiting throughout the year.
Notes. Two largely sympatric varieties can be recognized, mainly based on the tomentum
and the length of the style. Male flowers are not known in var. pilocarpa.
Key to varieties
Twigs towards apex and terminal leaf buds tomentose with minute hairs c. 0.1 mm long.
Fruits pubescent with stellate-dendroid hairs up to 0.1 mm long. Staminal column minutely
pubescent with greyish stellate hairs towards the base...
var. hirtella
W.J. de Wilde l.c. (1987) 131.
Leaves beneath with persistent tomentum composed of pale yellowish stellate hairs
c. 0.1 mm long, intermixed with early shed coarser hairs which leave distinct hair
scars. Style 00.5 mm long, persistent in fruit. Fruits obtuse at apex.
Endemic to Sabah and Sarawak.
Twigs towards apex and terminal leaf buds tomentose with rusty stellate-dendroid hairs
0.10.2(0.3) mm long. Fruits, especially towards the base, tomentose with rather rough
dendroid hairs c. 0.5 mm long...............................
var. pilocarpa W.J. de Wilde
(Latin, pilus = hair; Greek, karpos = fruit; the hairy fruit)
Blumea 25 (1979) 461, l.c. (1987) 131. Type: Cockburn SAN 85084, Borneo, Sabah, Lahad Datu
district (holotype L; isotypes K, KLU, SAN, SAR).
Leaves beneath with persistent tomentum consisting of yellowish hairs 0.1(0.2) mm

long, intermixed with early shed coarser hairs which leave distinct hair scars. Style
persistent in fruit, up to 0.5 mm long. Fruits obtuse to acute-acuminate at apex.
Endemic to Borneo (Sabah, Sarawak and E Kalimantan)
11. Knema intermedia (Blume) Warb.

(Latin, intermedius = in between; inference unknown)
Monogr. Myrist. (1897) 564; Gamble l.c. (1912) 239; Merrill l.c. (1921) 270; Ridley l.c. 68; Masamune
l.c. 303; Corner l.c. 565; J. Sinclair l.c. (1958) 315, l.c. (1961) 227; Backer & Bakhuizen f. l.c. 140; W.J.
de Wilde l.c. (1979) 427; Anderson l.c. 267; Whitmore, Tantra & Sutisna l.c. 269; Turner l.c. 360.
Basionym: Myristica intermedia Blume, Rumphia 1 (1835) 187. Type: Blume s.n., Java (holotype L;
isotype K). Synonym: M. corticosa var. decipiens Miq., Ann. Mus. Lugd.-Bat. 2 (1865) 51.
Tree up to 30 m tall, sometimes with stilt roots. Bark smooth, or scaly, or flaky. Sapwood
white or somewhat reddish. Twigs slender to stout, terete or more or less angled or flattened,
414
towards apex 1.54(5) mm diameter, rusty tomentose with hairs 0.30.5 mm long,
glabrescent; bark brown or blackish, not tending to crack or flake. Leaves chartaceous
or coriaceous, upper surface drying greenish brown, lower surface at first densely
tomentose, early glabrescent, without brownish or blackish dots; oblong to lanceolate,
930(35) 1.57(8) cm, base rounded or attenuate; midrib raised above; lateral veins
1225 pairs, strongly raised above; intercostal venation reticulate, raised and distinct
above; petiole 12.5 cm long. Inflorescences simple or 23-furcate, up to 1.5 cm diameter;
peduncle up to 2 mm long; 525-flowered in males; flowers pubescent with woolly hairs
0.30.5 mm long. Male flowers: pedicel 513 mm long; bracteole subpersistent, median
to apically inserted on the pedicel; buds subglobose, subtrigonous, (3)45 (3)45
(5.5) mm, inside cream (or pinkish?); perianth at sutures 0.30.4 mm thick, at anthesis
clefts to 2/33/4; staminal disc subtriangular, including anthers 22.5 mm diameter,
provided with a conspicuous mammilla 11.5 mm long; anthers 1115. Female flowers:
pedicel 28 mm long; buds obovoid, c. 5(6) mm long. Fruits 14 per infructescence,
ellipsoid, 2.54 1.52.3 cm, pubescent with hairs 0.30.5 mm long which are more
or less easily rubbed off; dry pericarp 23 mm thick; stalk (3)514 mm long.
Vernacular name. Sarawakkumpang daun panjang (Iban, preferred name).
Distribution. Sumatra, Peninsular Malaysia, W Java, and Borneo (Sabah, Sarawak, S
and SE Kalimantan).
Ecology. In lowland, hill and peatswamp forests, up to 1000 m altitude. Flowering and
12. Knema kinabaluensis J. Sinclair

Gard. Bull. Sing. 18 (1961) 229; W.J. de Wilde l.c. (1979) 450; Anderson l.c. 267; Whitmore, Tantra &
Sutisna l.c. 269. Type: Sinclair 9224, Borneo, Sabah, Mt. Kinabalu (holotype K; isotypes B, E, L, M,
SAN, SING).
Tree up to 25 m tall. Bark smooth or finely and shallowly fissured; inner bark pale
brown to reddish. Sapwood whitish, cambium reddish. Twigs towards apex more or less
bluntly triangular, 1.52.5 mm diameter, drying blackish with yellowish rusty tomentum
consisting of hairs c. 0.1 mm long, glabrescent; bark drying brownish or blackish, not
tending to crack or flake. Leaves rigidly coriaceous, upper surface drying dark brown,
lower surface greyish, without blackish dots, at first rather densely tomentose with mixed
hairs up to 0.1 mm long, glabrescent, usually leaving hair scars; oblong-lanceolate, 9
21 2.55 cm, base rounded or attenuate; midrib raised above; lateral veins 1225
pairs, raised above; intercostal venation reticulate, very distinct above; petiole 12 cm
long. Inflorescences simple or bifid, up to 1.5 cm diameter; 410-flowered in males;
flowers tomentose with mealy hairs 0.10.2 mm long. Male flowers: pedicel 36 mm
long; bracteole caducous, attached to the pedicel at about the middle; buds subglobose,
2.53.5 mm diameter, inside pale pink to reddish; perianth 3(5)-lobed, at anthesis clefts
to (3/4)4/5, at sutures up to 1 mm thick; staminal disc flattish, including anthers circular,
c. 1.5(2) mm diameter; anthers 78(10?), almost sessile; staminal column c. 1 mm
long. Female flowers: pedicel 34 mm long; buds obovoid-oblong, c. 5 mm long. Fruits
415
1 or 2 per infructescence, ovoid to oblong, 24.5 1.53 cm, apex usually acute-apiculate,
sometimes ridged along the line of suture, pubescent with persistent hairs 0.10.2 mm
long; dry pericarp 24 mm thick; stalk 310 mm long.
Distribution. Endemic to Borneo (confined to the Mt. Kinabalu area, Sabah).
Ecology. In hill mixed dipterocarp to upper montane forests, on ridges and slopes, along
rivers, also in degraded forest, on sandstone-derived soils, at 10002300 m altitude.
13. Knema korthalsii Warb.

(P.W. Korthals, 18071892, sometime botanist and plant collector in Indonesia)
Monogr. Myrist. (1897) 557; J. Sinclair l.c. (1961) 232; W.J. de Wilde l.c. (1979) 428; Argent et al.
(eds.) l.c. 455. Type: Korthals s.n., Borneo, Kalimantan (holotype L; isotypes CAL, S, U). Synonyms:
Myristica korthalsii (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 90; K. insularis Merr., Philip. J. Sc.
30 (1926) 394; K. cenabrei Merr. & Quisumbing, Philip. J. Sc. 37 (1928) 144.
Tree up to 30 m tall. Bark smooth or scaly, grey; inner bark c. 10 mm thick, soft, reddish.
Sapwood red or white; cambium reddish or yellow; heartwood hard, brown. Twigs towards
apex faintly angular, 25 mm diameter, at first tomentose with hairs 0.30.5 mm long,
glabrescent; bark brown or blackish, lower down smooth or striate, or more or less tending
to crack or thinly flaky. Leaves chartaceous or thinly coriaceous, upper surface drying
greenish brown to dark brown, lower surface glabrescent with sparse soft stellate hairs
sometimes remaining on midrib and lateral veins, without blackish dots, greyish; oblong
to (ob)lanceolate, 1235 2.59(12) cm, base cuneate or rarely rounded; midrib slightly
raised above; lateral veins 1240 pairs, slightly raised above; intercostal venation reticulate,
slightly raised above; petiole (0.8)12.5 cm long. Inflorescences simple or more or less
warted, up to 0.8 cm diameter, (1)520-flowered umbels in males; flowers tomentose
with hairs 0.20.7 mm long. Male flowers: pedicel (4)513 mm long; bracteole caducous,
attached to the pedicel at about the middle; buds depressed globose, more or less obtusely
trigonous in transverse section, 46(7) mm diameter, inside red; perianth at anthesis
clefts to c. 4/5; staminal disc (including anthers) circular, flattish or convex or mammillate,
22.8 mm diameter; anthers 1020; staminal column 0.71 mm long, more or less clasped
by the thickened disk-like base of the perianth. Female flowers: pedicel 1.52.5 mm
long; buds obovoid-oblong, 56 mm long. Fruits 13 per infructescence, (ob)ovoid,
ellipsoid or oblong, 1.54 12 cm, tomentose with hairs 0.51 mm long; dry pericarp
c. 3 mm thick; stalk (1)26 mm long.
Distribution. Borneo and southern parts of the Philippines.
Key to subspecies
Twig towards apex 35 mm diameter, older bark not flaky. Lateral veins (14)2040
pairs, usually clearly interarching near the leaf margin. Fruit 24 cm long; stalk 26
mm long; persistent style and stigma c. 1 mm long..............
416
subsp. korthalsii
Borneo and the southern parts of the Philippines. In primary mixed dipterocarp forests
on level lands or hilly sides, on a variety of soils, up to 400 m altitude.
Flowering in Borneo in JulyOctober.
Twig towards apex 23 mm diameter, older bark thinly flaky. Lateral veins 1222 pairs,
faintly interarching near the leaf margin. Fruit c. 2 cm long; stalk 12 mm long; persistent
stigma sessile..................
subsp. rimosa W.J. de Wilde
(Latin, rimosus = cracked; the older twig)
Blumea 43 (1998) 242. Type: Wood SAN A 1996, Borneo, Sabah (holotype L; isotype SAN).
Endemic to Borneo (Sabah, Sarawak). In primary lowland and hill mixed dipterocarp
forest on sandy soils or leached loam over sandy soils, up to 900 m altitude. Flowering
MarchApril, fruiting AprilJune.
14. Knema kostermansiana W.J. de Wilde

(A.J.G.H. Kostermans, 19061994, formerly prominent botanist at the Indonesian Forest
Research Institute and Herbarium Bogoriense, Bogor, Indonesia)
Blumea 25 (1979) 455, l.c. (1987) 127; Whitmore, Tantra & Sutisna l.c. 269. Type: James SAN 19047,
Borneo, Sabah (holotype L; isotypes K, SAN, SING).
Tree up to 20 m tall. Bark fissured, chocolate-brown to dark-brown, or dark grey; inner

bark whitish brown; sap red. Sapwood yellowish white or brownish. Twigs towards apex
terete, 24 mm diameter, at first tomentose with greyish hairs less than 0.1 mm long,
early glabrescent; bark drying brown or blackish, not tending to crack or flake. Leaves
membranous to chartaceous, upper surface drying olivaceous, lower surface greyish, without
blackish dots, inconspicuously tomentose with weak hairs up to 0.1 mm long, early
glabrescent; elliptic to oblong, 1230 5.512 cm, base shallowly cordate to broadly
rounded; midrib raised above; lateral veins 1522 pairs, sunken to flattish above; intercostal
venation very finely reticulate, distinct above; petiole 11.5 cm long. Inflorescences 2
3-furcate, up to 0.5 cm diameter; 510-flowered in males; flowers tomentose with persistent,
scale-like hairs less than 0.1 mm long. Male flowers: pedicel 2.57 mm long; bracteole
minute, caducous, attached to the apical part of the pedicel; buds depressed globose,
33.2 mm diameter, inside pink; perianth at sutures up to 1 mm thick, at anthesis clefts
to c. 4/5; staminal disc flat to faintly mammillate, including anthers circular, 1.52 mm
diameter; anthers 812; staminal column 0.60.7 mm long. Female flowers: pedicel 3
4 mm long; buds ellipsoid-oblong, more or less narrowed in the middle, (4)56 mm
long. Fruits 13 per infructescence, oblong, ellipsoid-oblong to almost fusiform, (5)7
8 2 cm, base tapering, apex long-tapering or beaked, scaly pubescent with persistent
hairs less than 0.1 mm long, appearing as if the fruits are subglabrous; dry pericarp
c. 2 mm thick; stalk rather stout, 810 mm long, with the bracteole scars located at
slightly above the middle.
Distribution. Endemic to Borneo (Sabah and NE Kalimantan).
Ecology. In primary and logged-over riverine forests on sandstone, up to 700 m altitude.
Flowering in JuneOctober; fruiting in JulyOctober.
417
15. Knema kunstleri (King) Warb.

(H.H. Kunstler, 18371887, German explorer and Kings plant collector in Peninsular Malaysia)
Monogr. Myrist. (1897) 568; Gamble l.c. (1912) 241; Ridley l.c. 69; J. Sinclair l.c. (1958) 291, l.c.
(1961) 236; Whitmore l.c. 338; W.J. de Wilde l.c. (1979) 468, l.c. (1996) 386; Anderson l.c. 267;
Whitmore, Tantra & Sutisna l.c. 269; Turner l.c. 360. Basionym: Myristica kunstleri King, Ann. Roy.
Bot. Gard. Calc. 3 (1891) 314, pl. 149. Lectotype: Kings Collector 4216, Peninsular Malaysia
(hololectotype CAL; isolectotypes BM, K, L).
Tree up to 30 m tall. Bark smooth, finely scaly, rough, or sometimes fissured, grey or
reddish to dark brown; inner bark pink or reddish. Sapwood cream, whitish, yellowbrown or pink. Twigs towards apex 1.53 mm diameter, at first tomentose with hairs
0.10.2 mm long, glabrescent; bark lower down not tending to crack or flake. Leaves
membranous to coriaceous, upper surface drying greenish to brown, lower surface greyish,
papillose, with dense or sparse dark dots, especially on the veinlets, subpersistently
pubescent with scattered stellate hairs c. 0.1 mm long; ovate, elliptic-oblong to oblonglanceolate, 520(28) (1.5)28.5(10) cm, base attenuate to broadly rounded, apex
subobtuse or usually acute-acuminate; midrib raised above; lateral veins 618 pairs, raised
above; petiole 0.52 cm long. Inflorescences sometimes supra-axillary, simple or more
or less forked, 0.20.4 cm diameter; 315-flowered in males, 15-flowered in females;
peduncle up to 5 mm long; flowers pubescent with scale-like hairs 0.10.2 mm long.
Male flowers: pedicel 511 mm long; bracteole subpersistent, attached to the pedicel
at or above the middle; buds depressed globose, sometimes more or less trigonous, 2.5
3(4) mm diameter, inside reddish or (in Sabah and Sarawak) cream; perianth at anthesis
clefts nearly to the base; staminal disc flat or slightly convex, including anthers circular
or somewhat triangular, 1.32(3) mm diameter; anthers 814; staminal column 0.51
mm long. Female flowers: pedicel 29 mm long; buds almost obovoid, 45 mm long.
Fruits 14 per infructescence, broadly ellipsoid, 1.54.5 12.8 cm, pubescent with
minute scale-like hairs up to 0.1 mm long; dry pericarp 12 mm thick; stalk 315 mm
long.
Distribution. A variable species with 5 subspecies, distributed in Sumatra, Peninsular
Malaysia, Borneo, and the Philippines. In Borneo 2 subspecies occur, both are found
Key to subspecies
Reticulation of intercostal venation on the upper leaf surface usually prominent and distinct.
Fruits 34 cm long..........
subsp. alpina (J. Sinclair) W.J. de Wilde
(Latin, alpinus = growing in alpine zone of mountains; growing at high altitude)
Blumea 25 (1979) 471; Whitmore, Tantra & Sutisna l.c. 269; Coode et al. (eds.) l.c. 224. Basionym:
K. cinerea (Poir.) Warb. var. alpina J. Sinclair l.c. (1961) 287. Type: Anderson S. 4514, Borneo,
Sarawak, Marudi district, G. Mulu, path from Melinau Paku (holotype SAR; isotypes A, K, L,
SAN, SING).
Stilt roots absent. Twig towards apex (1.5)23 mm diameter, sometimes angular.
Leaves thickly coriaceous, upper surface drying brown, lower surface glabrescent;
418
elliptic-oblong to oblong-lanceolate, 421 1.57 cm, base rounded to attenuate;

lateral veins 1018 pairs. Inflorescences sessile. Male flower: pedicel 58 mm long;
buds c. 3 34 mm, inside pale yellowish; staminal disc flat or rarely shallowly
convex, anthers (7)1014. Female flowers with pedicel 23 mm long. Fruits 34.2
(4.5) 22.8 cm; stalk (3)59(12) mm long.
Endemic to Borneo (Sabah, Sarawak, Brunei, W and S Kalimantan). In lowland mixed
dipterocarp to upper montane forests on andeside-derived soils, at (100)9002000
m altitude. Flowering in February, June, and OctoberNovember; fruiting throughout
the year.
Reticulation of intercostal venation on the upper leaf surface usually slightly prominent,
distinct or indistinct. Fruits 1.72 cm long..........................
subsp. coriacea (Warb.) W.J. de Wilde
(Latin, coriaceous = leathery; the leaves)
Blumea 25 (1979) 470; Whitmore, Tantra & Sutisna l.c. 269; Coode et al. (eds.) l.c. 225. Basionym:
K. coriacea Warb. l.c. (1897) 614. Type: Beccari 670, Borneo, Sarawak (FI). Synonym: Myristica
coriacea (Warb.) Boerl. l.c. 92.
Stilt roots often present. Twig towards apex 23 mm diameter, sometimes coarsely
striate, more or less angular. Leaves coriaceous, upper surface drying light brown;
elliptic to oblong or ovate-oblong, 621 37(7.5) cm, base usually rounded or
sometimes cuneate; lateral veins 818 pairs. Inflorescences sessile or with stout
peduncle up to 5 mm long. Male flower: pedicel 610.5 mm long; buds 2.22.5
2.53 mm; perianth inside cream (always?); anthers 1012. Female flowers not seen.
Fruits 1.72 1.41.6 cm; stalk 37(9) mm long.
Endemic to Borneo (Sabah, Sarawak and Brunei). In primary peatswamp and kerangas
forests, up to 100 m altitude. Flowering in AprilJune; fruiting in JulyNovember.
16. Knema latericia Elmer

(Latin, latericeus = brick-red; the perianth inside)
Leafl. Philip. Bot. 5 (1913) 1815; J. Sinclair l.c. (1958) 278, l.c. (1961) 238, excl. var. lunduensis; W.J.
de Wilde l.c. (1979) 395, l.c. 32 (1987) 115, l.c. 41 (1996) 388, l.c. 43 (1998) 245; Whitmore, Tantra &
Sutisna l.c. 270. Type: Elmer 12757, the Philippines, Palawan (holotype A; isotypes BM, BRSL, CAL,
E, G, K, L).
Shrub or tree up to 20 m tall. Bark brown, reddish brown, or chocolate, scaly or flaking
in fairly large elongated portions, not furrowed; inner bark pink or reddish. Sapwood
white or pale yellow, or pale brown, sometimes with streaks. Twigs slender, towards
apex (1)24(8) mm diameter, terete or more or less angular, roughly tomentose with
hairs 0.52 mm long, glabrescent; bark longitudinally fissured and cracked, later on usually
flaky. Leaves thinly coriaceous, upper surface drying pale brown or greenish, lower surface
glabrous, grey or whitish, not papillose; oblong to lanceolate, 630 19 cm, base attenuate
to rounded; midrib raised above; lateral veins 925(30) pairs, raised above; petiole
(0.5) 11.8 cm long. Inflorescences sessile, warted or furcate, up to 1 cm diameter,
320-flowered in males; flowers pubescent with persistent hairs 0.20.7 mm long. Male
flowers: pedicel 36 mm long; bracteole caducous, attached halfway to subapically on
the pedicel; buds depressed obovoid to depressed globose, 33.5 (3)44.5 mm, inside
brick-red; perianth at anthesis clefts to 2/34/5; staminal disc circular, flat or more or
less convex, or low-mammillate, including anthers c. 2 mm diameter; anthers 912; staminal
419
column c. 1 mm long. Female flowers: pedicel up to 1.5 mm long; bracteole subapically

inserted on the pedicel; buds much larger than in males, obovoid, (4)58 mm long.
Fruits 14 per infructescence, broadly ellipsoid to obovoid, sometimes subglobose,
(1) 1.52.5(3) 12(2.5) cm, tomentose with rough hairs 12 mm long; dry pericarp
(1)23 mm thick; stalk 25 mm long.
Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, Singapore, Borneo, and
the Philippines (Palawan).
Ecology. In primary and degraded forests, on hilly sides and ridges, also in kerangas
forest, on a variety of soils, up to 1500 m altitude. Flowering and fruiting mainly in
MayOctober.
Notes. A variable species with three recognized subspecies, two of which are found in
Sabah and Sarawak.
Key to subspecies
Tomentum of twig towards apex and of flowers dark brown or rusty. Male flower buds
more or less depressed globose, often somewhat broader than long. Staminal disc usually
convex or low-mammillate. Leaves 410 cm broad, upper surface drying dark brown to
greenish brown, lower surface pale brown to grey-glaucous. Fruits variable in size, with
bright brown to rusty-reddish tomentum.......................
subsp. ridleyi (Gandoger) W.J. de Wilde
(H.N. Ridley, 18551956, sometime botanist and former Director of the Singapore Botanic
Gardens)
Blumea 25 (1979) 396, l.c. (1981) 223; Turner l.c. 360; Coode et al. (eds.) l.c. 225. Basionym:
Myristica ridleyi Gandoger, Bull. Soc. Bot. Fr. 66 (1919) 226, in clavi. Type: Ridley s.n., Singapore
(Herb. Gandoger, Lyon). Synonyms: K. conferta (King) Warb. var. borneensis Warb. l.c. 580, p.p.;
M. conferta King var. borneensis (Warb.) Boerl. l.c. 1.
Twig towards apex up to 8 mm diameter, variably rusty tomentose; bark of older

twig usually flaky. Female flower buds 58 mm long; pedicel up to 1.5 mm long.
Fruits (1)1.53 cm long.
Peninsular Thailand, Sumatra, Peninsular Malaysia, Singapore, and Borneo (Sabah,
Sarawak and Kalimantan).
Forma nana W.J. de Wilde l.c. (1998) 246, type: Anderson K 10 (L), from Sabah,
Sarawak and Brunei, is distinct by the slender, non-flaky twigs and small, lanceolate
leaves. It may be confused with K. minima W.J. de Wilde (l.c. 249) from Brunei
which is distinct by the male flower buds measuring only c. 2 mm diameter.
Tomentum of twig towards apex and of flowers pale brown or yellowish brown. Male
flower buds obovoid. Staminal disc flat or slightly convex. Leaves 25 cm wide, upper
surface drying pale brown or pale greenish brown, lower surface usually whitish. Fruits
rather small, with pale brown or yellowish brown tomentum......................
subsp. albifolia (J. Sinclair) W.J. de Wilde
(Latin, alba = white, folium = leaf; the pale colour of the lower leaf surface)
Blumea 25 (1979) 397; Anderson l.c. 267; Coode et al. (eds.) l.c. 225. Basionym: K. latericia
Elmer var. albifolia J. Sinclair l.c. (1961) 243. Type: Sinclair 9269, Borneo, Sabah (holotype K;
isotypes A, B, E, L, M, P, PNH, SAN, SING).
Twig towards apex with fine pale brown tomentum; bark later on flaky. Female flower
420
buds c. 5 mm long. Fruits subsessile, subglobose to ellipsoid, or obovoid, 1.42(3.5)

cm long.
Endemic to Borneo (Sabah, Sarawak and Kalimantan, including Nunukan Is.).
17. Knema latifolia Warb.
Fig. 8.
(Latin, latus = broad, folium = leaf; broad-leaved)

Monogr. Myrist. (1897) 610; Merrill l.c. (1921) 271; Masamune l.c. 303; J. Sinclair l.c. (1961) 244; W.J.
de Wilde l.c. (1979) 414; Anderson l.c. 267; Whitmore, Tantra & Sutisna l.c. 270; Coode et al. (eds.) l.c.
225; Argent et al. (eds.) l.c. 456. Syntypes: Forbes 2997, 3188, Sumatra (B, lost; K, L); Beccari 1991,
Borneo, Sarawak (FI, K). Synonyms: K. umbellata Warb. l.c. (1897) 609; Myristica umbellata (Warb.)
Boerl. l.c. 92, nom. inval.; K. nitida Merr., J. As. Soc. Str. Br. 85 (1922) 190; K. winkleri Merr. l.c.
(1922) 189.
Tree up to 20 m tall. Bark almost smooth, peeling off in narrow strips; inner bark brittle,
pinkish or red, or light brown. Sapwood off-white, or pale yellow, or pink. Twigs towards
apex (1)23 mm diameter, tomentose with hairs c. 0.1 mm long, early glabrescent; bark
drying brown or blackish, not flaky. Leaves thinly coriaceous, upper surface drying often
blackish and shining, lower surface often tinged reddish brown or purplish, at first sparsely
tomentose with minute, stellate, scale-like hairs c. 0.1 mm long, usually early glabrescent,
without blackish dots; elliptic-oblong to obovate-oblong, (7)1030 (2.5)4.512.5 cm,
base attenuate to rounded; midrib slightly raised above; lateral veins 816 pairs, rather
well-spaced, slightly raised above; intercostal venation coarsely reticulate, distinct above;
petiole 1.53 cm long. Inflorescences simple or 24-furcate, up to 0.8 cm diameter;
peduncle up to 5 mm long; 515-flowered in males; flowers pubescent with inconspicuous
persistent hairs up to 0.1 mm long, subglabrescent. Male flowers: pedicel 510 mm
long; bracteole minute, subpersistent, subapically attached on the pedicel; buds subglobose
or ovoid-trigonous, 2.53.5 3.55 mm, inside pale yellowish (always?); perianth at
anthesis clefts to c. 4/5(5/6); staminal disc conspicuously and sharply 3(4)-angled, rather
flat or slightly convex, including anthers 22.5 mm diameter; anthers 3(4); staminal
column slender, 0.51 mm long. Female flowers: pedicel (4)510 mm long; buds ellipsoidobovoid, 57 mm long. Fruits 12 per infructescence, globose-ellipsoid to ellipsoid-oblong,
23.5 23 cm, usually somewhat flattened, often broadly ridged, and somewhat saccate
at the base, tomentose with hairs 0.10.2 mm long, subglabrescent; dry pericarp 23
mm thick; stalk 720 mm long.
Distribution. Sumatra and Borneo (Sabah, Sarawak, Brunei and Kalimantan, including
Nunukan Is.).
Ecology. In primary and degraded lowland and hill mixed dipterocarp forests on loamy,
clayey, sandy, acidic, and lime-containing soils, up to 500 m altitude. Flowering throughout
the year; fruiting mainly in AugustDecember.
18. Knema laurina (Blume) Warb.

(Latin, laurinus = resembling laurel trees; the leaves)
421
vi
303; J. Sinclair l.c. (1958) 329, l.c. (1961) 248, p.p., excl. most synonyms; Backer & Bakhuizen f. l.c.
140; Whitmore l.c. 338; W.J. de Wilde l.c. (1979) 379; Anderson l.c. 267; Whitmore, Tantra & Sutisna
l.c. 270; Turner l.c. 360; Coode et al. (eds.) l.c. 225; Argent et al. (eds.) l.c. 456. Basionym: Myristica
laurina Blume, Rumphia 1 (1835) 189. Type: Blume s.n., Java (L). Synonyms: M. laurina Blume var.
borneensis Miq. l.c. (1865) 51; M. cantleyi Hook. f., Fl. Brit. Ind. 5 (1886) 110.
Shrub or tree up to 30 m tall. Bark grey, reddish brown, or brown, smooth, finely scaly,
slightly cracked, or furrowed; inner bark reddish. Sapwood white, yellowish, red with
brown rays, light brown, or yellow with brown stripes. Twigs towards apex 25(6) mm
diameter, rough tomentose with hairs 0.52 mm long, late-glabrescent; bark drying brownish
or blackish, lower down not tending to crack or flake. Leaves membranous to coriaceous,
upper surface drying greenish to brown, lower surface without dark dots, finely papillose,
persistently tomentose with stiff, regularly set, equally long-stalked dendroid hairs c. 1
mm long; obovate-elliptic to oblong-lanceolate, 930 2.510.5 cm, base attenuate to
rounded; midrib more or less raised above, late-glabrescent; lateral veins 1228 pairs,
flat or sunken above; petiole late-glabrescent, 12 cm long. Inflorescences subsessile,
simple or 2-fid, up to 0.7 cm diameter, 520-flowered in males; flowers pubescent with
dense hairs 0.51.5(2) mm long. Male flowers: pedicel (0.5)25 mm long; bracteole
subpersistent or caduous, inserted on the apical part of the pedicel or nearly so; buds
obovoid-oblong, 35(6) 24 mm, inside pink or red; perianth at anthesis clefts to
c. 1/2; staminal disc flat or slightly concave, including anthers circular, 11.5 mm diameter;
anthers 69, sessile, suberect; staminal column somewhat tapering, 1.52.5 mm long,
glabrous or rarely minutely pubescent at the base. Female flowers: pedicel up to 1 mm
long; buds ovoid-oblong, 46 mm long. Fruits 15 per infructescence, ovoid or ellipsoidoblong, 1.53 12 cm, rusty tomentose with hairs 13 mm long; dry pericarp 1.5
2 mm thick; stalk 01(5) mm long.
Vernacular name. Sabahdarah-darah kerantu (preferred name).
Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, W and C Java, and
Borneo. Common in Sabah and Sarawak. Also in Brunei and Kalimantan.
Ecology. In primary and degraded lowland to lower montane forests, on a variety of
soils, up to 1500 m altitude. Flowering and fruiting throughout the year.
19. Knema linguiformis (J. Sinclair) W.J. de Wilde
Fig. 9.
(Latin, linguiformis = tongue-shaped; the leaves)

Blumea 25 (1979) 416, l.c. (1981) 227, l.c. (1987) 121; Whitmore, Tantra & Sutisna l.c. 270; Coode et
al. (eds.) l.c. 225; Argent et al. (eds.) l.c. 456. Basionym: K. curtisii (King) Warb. var. linguiformis J.
Sinclair l.c. (1961) 200. Type: Sinclair (Kadim & Kapis) 9294, Borneo, Sabah, Sepilok FR, Compt. 16
(holotype K; isotypes A, B, BM, E, L, M, SAN, SING).
Tree up to 25 m tall. Bark smooth or almost so, or slightly scaly; inner bark soft, c.
5 mm thick, reddish or brown. Sapwood dirty white. Twigs towards apex 12 mm diameter,
at first tomentose with woolly hairs 0.20.5 mm long, late-glabrescent; bark lower down
yellowish, not tending to crack or flake. Leaves chartaceous, upper surface drying greenish,
lower surface minutely papillose, without reddish brown dots, early glabrescent; oblong
423
vi
to lanceolate, 515 1.55(6) cm, base attenuate to rounded, apex acute to obtuse;
midrib raised above; lateral veins 820 pairs, raised above; petiole late-glabrescent, 0.6
1.5 cm long. Inflorescences sessile, simple or forked, 0.20.5 cm diameter; 520-flowered
in males; flowers tomentose with hairs (0.1)0.20.3(0.4) mm long, glabrescent. Male
flowers: pedicel slender 5.513 mm long; bracteole persistent, attached to the pedicel
at about the middle; buds depressed globose to broadly obovoid, in transverse section
often obtusely trigonous, 3.56 mm diameter, inside pink; perianth at anthesis clefts to
c. 3/4; staminal disc subcircular or obtusely trigonous, flat, including anthers 1.53 mm
diameter; anthers 1925, closely appressed, sessile, suberect, hidden under the staminal
disc; staminal column tapering, 0.52 mm long. Female flowers: pedicels slender, 5
8 mm long; buds depressed globose or obovoid, 46 mm long; stigma subcircular, 8
21-lobulate. Fruits 13 per infructescence, ellipsoid to ovoid, 3.54 22.5 cm, at base
often contracted into a short stipe, at first tomentose with minute hairs, glabrescent; dry
pericarp c. 2 mm thick; stalk (2)58 mm long.
Ecology. In primary and degraded lowland mixed dipterocarp to lower montane forests
on hillsides and ridges, on various types of soils, up to 1650 m altitude. Flowering and
20. Knema longepilosa (W.J. de Wilde) W.J. de Wilde

(Latin, longus = long, pilosus = covered with hairs; the long-hairy flowers)
Blumea 41, 2 (1996) 389. Basionym: K. percoriacea J. Sinclair f. longepilosa W.J. de Wilde l.c. (1979)
394. Type: Sibat S. 22985, Borneo, Sarawak, Limbang district, Mt. Murud (holotype L; isotypes A, K,
SAN, SAR, SING).
Tree up to 10 m tall. Bark reddish brown; sap red. Twigs towards apex 24 mm diameter,
looking much thicker (46 mm) by the dense yellow-brown or dark rusty, shaggy hairs
23 mm long, late-glabrescent; bark finely flaky. Terminal leaf buds short, 5(10) 4
5 mm, pubescent with hairs 24 mm long similar to those of the twig apex and petiole.
Leaves thinly coriaceous, upper surface drying olivaceous, lower surface glabrous, greyish,
with midrib and veins brown-yellow, contrasting with the leaf surface; elliptic-oblong
to lanceolate, 1325 36 cm, base narrowly rounded or cuneate; midrib pale and
prominently raised above; lateral veins 1015 per side, raised; petiole 11.5 cm long,
at first densely yellow-brown or rusty pubescent with hairs 24 mm long, late-glabrescent,
in colour contrasting with the early glabrescent leaf blades. Inflorescences simple or forked,
up to 0.5(1) cm diameter, late-glabrescent; 26-flowered in males; flowers densely and
persistently pubescent with hairs 24 mm long. Male flowers densely yellow-brown or
rusty pubescent with hairs (2)4 mm long, appearing as mops of 68 mm diameter; pedicel
34 mm long; bracteole minute; buds subglobose, 3.54 mm diameter, perianth at anthesis
clefts to c. 1/2; staminal disc flat, bluntly subtriangular, 1.5(2) mm diameter; staminal
column short, tapering, c. 1 mm long, glabrous; anthers 1518, sessile, contiguous. Female
flowers and fruits not known.
Distribution. Endemic to Borneo (C Sarawak, W and C Kalimantan).
425
Ecology. In lowland mixed dipterocarp forest on basaltic or clay soils, up to 250 m altitude.
Flowering in January and April.
21. Knema lunduensis (J. Sinclair) W.J. de Wilde

(of Lundu area, Sarawak)
Blumea 25 (1979) 395, l.c. (1996) 391; Whitmore, Tantra & Sutisna l.c. 270; Coode et al. (eds.) l.c. 225.
Basionym: K. latericia Elmer var. lunduensis J. Sinclair l.c. (1961) 244. Type: Sinclair (& Kadim)
10381, Borneo, Sarawak, Lundu district, G. Gading (holotype K; isotypes A, E, L, SAR, SING).
Tree up to 20 m tall. Bark flaky, or fissured, or smooth; inner bark light red. Sapwood
white. Twigs stout, towards apex 46(10) mm diameter, tomentose with hairs (0.5)1
1.5 mm long, glabrescent; bark longitudinally cracked and flaky. Leaves chartaceous to
thinly coriaceous, upper surface drying brown, lower surface glabrous; obovate-elliptic
to oblong-oblanceolate, 1550 4.516(20) cm, base obtuse to rounded, or subtruncate,
or rarely subcordate; midrib raised above; lateral veins 1230 pairs, raised above; petiole
13 cm long. Inflorescences sessile, simple or forked, up to 1 cm diameter, 515-flowered
in males; flowers tomentose with persistent hairs (0.4)0.51.5(2) mm long. Male flowers:
pedicel 34 mm long; bracteole more or less persistent, attached to the pedicel slightly
above the middle; buds depressed globose, 33.5(7) 45(6) mm, inside flesh-pink;
perianth at anthesis clefts to c. 4/5; staminal disc flat or slightly convex, subcircular,
including anthers 1.82 mm diameter; anthers 1012(14); staminal column tapering, 0.6
1 mm long. Female flowers not seen. Fruits 16 per infructescence, ellipsoid, (2)2.5
3.5(4) 1.52.5(3.5) cm, tomentose with coarse hairs 12 mm long; dry pericarp 2
4 mm thick; stalk 03 mm long.
Ecology. In primary lowland mixed dipterocarp to lower montane forests, on granitic
soils, at 1001500 m altitude. Flowering in June; fruiting in JuneNovember.
22. Knema luteola W.J. de Wilde

(Latin, luteolus = yellowish; the pale twig)
Blumea 25 (1979) 451; Whitmore, Tantra & Sutisna l.c. 270. Type: Endert 2822, Borneo, central E
Kalimantan (holotype L).
Tree up to 20 m tall. Bark brownish, somewhat fissured, strongly peeling off; inner bark
c. 3 mm thick, reddish, inside yellow. Sapwood pale yellowish. Twig towards apex
somewhat flattened or bluntly triangular, yellow-brown, smooth, 1.52.5(3) mm diameter,
at first tomentose with minute greyish hairs up to 0.1 mm long, early glabrescent; bark
not tending to crack or flake. Leaves chartaceous, upper surface drying greenish, lower
surface greyish, without blackish dots, at first tomentose with minute, weak, grey hairs
and scales, early glabrescent; oblong to lanceolate, 1230 39.5 cm, base rounded
to cuneate; midrib slender, raised above; lateral veins 1625 pairs, not or only faintly
426
raised above; intercostal venation forming a very fine network; petiole 11.5 cm long.
Inflorescences simple or 24-fid, up to 0.5 cm diameter; 512-flowered in males; flowers
persistently pubescent with minute yellowish scale-like hairs up to 0.1 mm long. Male
flowers: pedicel 47.5 mm long; bracteole caducous, 12.5 mm long, attached on the
pedicel below the perianth; buds globose, not depressed, 2.54 mm diameter, inside yellow
or pale yellow; perianth at anthesis clefts to 3/44/5, at the sutures up to 1 mm thick;
staminal disc flat, including anthers circular, 1.72.3 mm diameter; anthers 910; staminal
column c. 0.5 mm long. Female flowers not seen. Fruits 14 per infructescence, ellipsoid
to ellipsoid-oblong, 37 22.5 cm, usually beaked at the apex, pubescent with minute
scale-like hairs less than 0.1 mm long; dry pericarp c. 2 mm thick; stalk stout, 1520
mm long, bracteole scar situated at or below the middle.
Distribution. Endemic to Borneo (Sarawak, W and E Kalimantan).
Ecology. In lowland and hill mixed dipterocarp forests, often on stream banks, at 100
700 m altitude. Flowering in June, August, and December; fruiting in FebruaryMarch.
23. Knema membranifolia Winkler

(Latin, membranus = thin, folium = leaf; thin-leaved)
Engl. Bot. Jahrb. 49 (1913) 368; J. Sinclair l.c. (1961) 262; W.J. de Wilde l.c. (1979) 422; Anderson l.c.
267; Whitmore, Tantra & Sutisna l.c. 270; Coode et al. (eds.) l.c. 225. Lectotype: Winkler 2460, Borneo,
SE Kalimantan (hololectotype BRSL; isolectotypes BM, BO, G, K, L, P, SING).
Tree up to 25 m tall. Bark thin, smooth, peeling off in thin strips, or flaky; inner bark
35 mm thick, brown. Sapwood pale brown or white. Twigs towards apex 1.53 mm
diameter, subterete or bluntly 23-angular, at first densely tomentose with hairs c. 1 mm
long, glabrescent; bark yellowish, that of older twig smooth or usually tending to crack
or flake. Leaves membranous to thinly coriaceous, upper surface drying olivaceous, lower
surface without blackish dots, early glabrescent; oblong to lanceolate, 1228 3.58.5
cm, base attenuate to subobtuse, apex sometimes obtuse; midrib prominent above; lateral
veins 1217 pairs, distinct but hardly raised above; petiole 1.52.5 cm long. Inflorescences
sessile, simple or 24-tubercled, up to 1 cm diameter; 530-flowered in males; flowers
at first tomentose with hairs 0.51 mm long, early glabrescent. Male flowers: pedicel
69 mm long; bracteole caducous, attached to the pedicel at about the middle; buds
depressed globose, obtusely trigonous, 3.55 57(8) mm; perianth at anthesis clefts
to 3/44/5; staminal disc flat or convex, but not mammillate, including anthers circular,
33.5 mm diameter; anthers 1920(23?); staminal column 0.71 mm long. Female
flowers: pedicel 14 mm long; buds broadly obovoid, 3.54.5 mm long. Fruits 13 per
infructescence, ellipsoid or obovoid, 2.54.5 23.5 cm, at first tomentose with very
short hairs, early glabrescent; dry pericarp 1.52 mm thick; stalk 15 mm long.
Ecology. In lowland and hill mixed dipterocarp forests on loamy and basalt-derived soils,
up to 500 m altitude. Flowering mainly in AprilJune; fruiting throughout the year.
427
24. Knema mogeana W.J. de Wilde

(J. Mogea, botanist of the Herbarium Bogoriense, Bogor, Indonesia)
Blumea 32 (1987) 136; Whitmore, Tantra & Sutisna l.c. 270. Type: Mogea (& de Wilde) 4446, Borneo,
C Kalimantan (holotype L; isotype BO).
Tree up to 15 m tall. Bark dark brown, smooth or rough when old, with some pustules,
not furrowed or striate; sap copious, red. Twigs slender, towards apex terete, 11.5
(2) mm diameter, minutely pubescent or glabrescent with dull brown hairs 0.10.2 mm
long; bark lower down blackish brown, not flaky. Terminal leaf buds slender, 812 mm
long, tomentose with hairs 0.10.2 mm long. Leaves membranous, upper surface drying
olivaceous or dark brown, lower surface greyish, without blackish dots, persistently
tomentose with widely scattered, pale stellate hairs 0.10.2 mm across; oblong to lanceolate,
817(24) 1.54(7) cm, base narrowly rounded to attenuate; midrib raised above;
lateral veins 1320 pairs, sunken or flatfish, sometimes raised above; reticulation of
intercostal venation faint above; petiole 0.81.2(1.5) cm long. Inflorescences simple
or forked, up to 0.4 cm diameter, not stalked, in males 48-flowered; flowers persistently
tomentose. Male flowers densely tomentose with hairs 0.10.2 mm long; pedicel slender,
24 mm long; bracteole persistent, 0.51 mm long, located at c. 0.5 mm below the perianth;
buds broadly obovoid, c. (2)2.5 mm diameter; perianth at anthesis clefts to 4/55/6,
at sutures up to 1 mm thick; staminal disc circular, flat above, including anthers 11.5
mm diameter; anthers 68; staminal column c. 1(1.4) mm long, glabrous. Female flowers:
pedicel 36(8) mm long; buds long-obovoid, c. 4 mm long. Fruits solitary or up to
6 per infructescence, ellipsoid to ellipsoid-oblong, 1.52 11.2 cm, tomentose with
persistent hairs c. 0.1 mm long; dry pericarp c. 1 mm thick; stalk 410 mm long.
Distribution. Endemic to Borneo (Sabah, Sarawak, C and E Kalimantan).
Ecology. In primary mixed dipterocarp forest on sandy loam soils, on hill slopes and
ridges, also in kerangas forest on sandy soils, and on limestone, up to 300 m altitude.
Flowering in DecemberJanuary, and MayJune; fruiting in January and JuneSeptember.
25. Knema muscosa J. Sinclair

(Latin, muscosus = moss-like; the twig apices)
Sutisna l.c. 270. Type: Clemens 22120, Borneo, Sarawak, Kapit district, G. Majau, Gat, upper Rejang
R. (holotype K; isotypes A, BM, BO, L, NY, PNH, SAR, SING).
Tree c. 8 m tall. Twigs slender, towards apex 11.5 mm diameter, initially tomentose
with hairs 0.20.7 mm long, glabrescent; bark greyish black, not tending to crack or
flake. Leaves chartaceous, upper surface drying brown, lower surface brownish grey, with
scattered dark brown dots, at first tomentose with conspicuous sessile stellate and dendroid
hairs 0.20.7 mm long, glabrescent; oblong to oblong-lanceolate, 5.512(15) 1.5
3.5 cm, base subattenuate; midrib slender, raised above; lateral veins 714 pairs, slender,
raised and distinct above; petiole 0.71 cm long. Inflorescences simple or 23-fid, up
to 0.3 cm diameter, (1)28-flowered in males; flowers woolly tomentose with hairs 0.2
0.5 mm long; peduncle up to 2 mm long. Male flowers: pedicel 23 mm long; bracteole
428
minute, more or less persistent, located on the pedicel at about the middle; buds subglobose
or broadly obovoid, 22.5 mm diameter; perianth at anthesis clefts to c. 3/4; staminal
disc flat, including anthers circular, 11.3 mm diameter; anthers 68; staminal column
0.71 mm long. Female flowers not known. Fruits 1 per infructescence, obovoid-oblong,
1.71.8 1 cm, apex shortly acute-mucronate, tomentose with hairs c. 0.3 mm long;
dry pericarp c. 0.5 mm thick; stalk 67 mm long, with the bracteole scar more or less
at median position.
Ecology. In montane mossy forest.
26. Knema oblongata Merr.
Fig. 10.
(Latin, oblongus = oblong; the leaves)

J. As. Soc. Str. Br. 85 (1922) 190; Masamune l.c. 303; W.J. de Wilde l.c. (1979) 400; Whitmore, Tantra
& Sutisna l.c. 271. Type: Ramos 1663, Borneo, Sabah, Sandakan (holotype PNH, probably destroyed;
isotypes A, BM, BO, BRI, K, L, UC, US).
Shrub or tree up to 20 m tall. Bark smooth, sometimes scaly or slightly fissured; inner
bark red-brown, beef-red, or pinkish. Sapwood pale yellowish, brownish, or whitish. Twigs
towards apex 28 mm diameter, at first tomentose with hairs 0.20.6 mm long, glabrescent;
bark drying brownish or blackish, lower down not cracked or flaky. Leaves chartaceous
to coriaceous, lower surface finely papillose, without reddish brown dots, with persistent
or partially late-glabrescent tomentum consisting of mixed sessile and stalked stellate
and dendroid hairs; elliptic to oblong or lanceolate, 1145 2.514 cm, base subattenuate
to rounded, or shallowly cordate; midrib raised above; lateral veins (10)1826 pairs,
raised to slightly sunken above; petiole late-glabrescent, 12.5 cm long. Inflorescences
simple or forked, up to 0.5 cm diameter, 525-flowered in males; peduncle 04 mm long;
flowers pubescent with hairs 0.11(2) mm long. Male flowers: pedicel 28 mm long;
bracteole usually caducous, attached to the pedicel at the middle or subapically; buds
depressed globose-obovoid, 2.55 2.55 mm, tapering in the lower half; perianth at
sutures 0.51 mm thick, at anthesis clefts to 1/22/3(3/4); staminal disc including anthers
subcircular, 1.42.5 mm diameter; anthers 79 or 1318(21), sessile to slightly stiped;
staminal column 12 mm long. Female flowers: pedicel 2.510 mm long; bracteole attached
to the pedicel at or above the middle; buds ellipsoid to obovoid, 67.5 mm long. Fruits
13 per infructescence, ovoid to obovoid, 23.8 1.52 cm, pubescent with hairs 1
1.5 mm long; dry pericarp 24 mm thick; stalk 310 mm long.
Ecology. In primary and degraded lowland mixed dipterocarp to lower montane forests,
on various types of soils, up to 1200 m altitude. Flowering mainly in JuneAugust; fruiting
mainly in OctoberDecember.
Notes. A variable species with three subspecies, with subsp. oblongata the most common.
All three subspecies occur in Sabah and Sarawak.
429
Key to subspecies
1.
Tomentum of twig towards apex composed of hairs c. 0.2(0.3) mm long. Lateral

veins c. 10 pairs. Mature male flower buds c. 2.5 mm diameter, tomentose with hairs
c. 0.1(0.2) mm long...........
subsp. parviflora W.J. de Wilde
(Latin, parvus = small, florum = flower)
Blumea 25 (1979) 404. Type: Agam SAN 31488, Borneo, Sabah, Lahad Datu district (holotype
L; isotypes BO, K, SAN).
Twigs slender, towards apex 23 mm diameter, tomentose with hairs 0.10.3 mm

long. Leaves 817 36.5 cm, base rounded; lateral veins c. 10 pairs. Inflorescences
sessile. Male flowers: pedicel c. 2 mm long; bracteole attached to
the pedicel apically; buds broadly obovoid; perianth lobes at the sutures c. 0.5
mm thick, at anthesis clefts to c. 2/3; anthers 8, shortly stiped, 0.30.4 mm long.
Tomentum of flowers consists of hairs c. 0.1 mm long.
Endemic to Borneo: Sabah, Lahad Datu district. In primary mixed dipterocarp
forest on hill sides, up to 150 m altitude. Flowering in May.
Tomentum of twig towards apex composed of hairs 0.30.6 mm long. Lateral veins
1826 pairs. Mature male flower buds 3.55 mm diameter, tomentose with hairs 0.4
1(2) mm long.................................2
2.
Leaves rather narrow, oblong to lanceolate, 25.5 cm wide, base attenuate to obtuse.
Inflorescences with peduncle 14 mm long. Male flower buds obovoid, c. 4 mm
diameter; pedicel 23 mm long. Anthers 79, almost entirely sessile.............................
subsp. pedunculata W.J. de Wilde
(Latin, pedunculatus = provided with a peduncle; the inflorescence)
Blumea 25 (1979) 403. Type: Aban SAN 49195, Borneo, Sabah, Kota Kinabalu district, Sapangar
Is. (holotype L; isotypes K, SAN).
Twigs slender, towards apex 23.5 mm diameter, tomentose with hairs c. 0.5 mm
long. Leaves 1132 25.5 cm; lateral veins c. 20 pairs. Male flowers obovoid,
34 34.2 mm; perianth at the sutures 0.31.5 mm thick, at anthesis clefts
to 1/2 or slightly more. Tomentum of flowers consists of hairs 0.42 mm long.
Fruits c. 2 cm long; stalk c. 6 mm long.
Endemic to Borneo: Sabah (Beaufort, Tenom and Kota Kinabalu districts). In
primary mixed dipterocarp forest on ridges and hilly sides, on yellow or brownish
soils, up to 600 m altitude. Flowering in September.
Leaves usually broader, elliptic to oblong-lanceolate, 414 cm wide, base usually
broadly rounded to shallowly cordate, rarely subattenuate. Inflorescences sessile or
with peduncle up to 1 mm long. Male flower buds 3.55 mm diameter; pedicel 2
8 mm long. Anthers 1321, subsessile or shortly stiped..........................
subsp. oblongata
Synonym: K. obovoidea Merr. l.c. (1929) 75; Masamune l.c. 303.
Twig usually stout, towards apex 2.58 mm diameter, tomentose with hairs 0.3
0.6 mm long. Leaves 1145 414 cm, base sometimes shallowly cordate; lateral
veins c. 20 pairs. Male flower buds broadly obconical, 45 3.55 mm; perianth
lobes at the sutures 0.51 mm thick, at anthesis clefts to 1/22/3 (rarely to c.
3/4). Tomentum of flowers consists of hairs 0.41 mm long. Fruits 23.8 cm
long; stalk 310 mm long.
Distribution as the species.
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27. Knema pallens W.J. de Wilde
Fig. 11.
(Latin, pallens = pale; the whole plant)

Blumea 25 (1979) 391; Whitmore, Tantra & Sutisna l.c. 271; Argent et al. (eds.) l.c. 456. Type: Saikeh
SAN 73312, Borneo, Sabah, Keningau district (holotype L; isotype SAN).
Tree up to 30 m tall. Bark scaly; inner bark light red or whitish. Sapwood yellow or
yellow-ochre; heartwood dark brown. Twigs stout, subterete, towards apex 512 mm
diameter, tomentose with light brown hairs 0.22(2.5) mm long, glabrescent; bark
conspicuously longitudinally cracked and flaky. Leaves subcoriaceous, upper surface drying
greenish brown, lower surface early glabrescent, papillose; elliptic to lanceolate, 2555
(5)623 cm, base usually cordate, apex obtuse or rounded; midrib raised above; lateral
veins 1832 pairs, raised above; petiole stout, late-glabrescent, 12 cm long. Inflorescences
sessile, simple or warted, 0.52 cm diameter, (5)1030-flowered in males; flowers
pubescent with persistent yellowish hairs 0.10.8 mm long. Male flowers: pedicel 12
20 mm long; bracteole caducous or subpersistent, attached to the pedicel at or above
the middle, or almost apically; buds depressed globose or broadly obovoid, 610 6.5
10 mm, inside reddish, at the basal part much thickened into a disc; perianth at anthesis
clefts to 4/5 or more; staminal disc circular, flattish, including anthers 2.54.5 mm diameter;
anthers 1523; staminal column 11.5 mm long. Female flowers: pedicel (7)1020 mm
long; buds obovoid, c. 11 89 mm. Fruits 16 per infructescence, ellipsoid, 3.57
2.53.5(4.5) cm, tomentose with hairs 12 mm long; dry pericarp 47 mm thick;
stalk (10)1430 mm long.
Ecology. In lowland mixed dipterocarp forest, also in kerangas forest, and forest behind
mangrove, up to 300 m altitude. Flowering mainly in JanuaryMay; fruiting throughout
the year.
28. Knema pectinata Warb.

(Latin, pectinatus = with comb-like divisions; the veins of the leaves)
Monogr. Myrist. (1897) 556; Masamune l.c. 303; J. Sinclair l.c. (1961) 266; W.J. de Wilde l.c. (1979)
430, l.c. (1981) 227; Anderson l.c. 268; Whitmore, Tantra & Sutisna l.c. 271. Type: Beccari 1607,
Borneo, Sarawak, Kuching district (holotype FI; isotypes G, K, M, P, S).
Tree up to 25 m tall. Bark hard, with a few large flakes and dents where the flakes
have fallen out, otherwise smooth. Twigs towards apex subterete, grey-brown, 36 mm
diameter, at first mealy tomentose with hairs 0.1(0.2) mm long, early glabrescent; lower
down bark not tending to crack or flake. Leaves coriaceous or rigidly coriaceous, upper
surface drying olivaceous to brown, lower surface without blackish dots, early glabrescent
or with grey-brown, subpersistent tomentum (especially near the veins) consisting of hairs
0.10.3 mm long; oblong to lanceolate, often more or less parallel-sided, (15)2045
3.510 cm, base truncate or obtuse, to attenuate, apex obtuse or acute-acuminate; midrib
slightly raised above; lateral veins 2550 pairs, flat to slightly raised and distinct above;
petiole 1.52.5 cm long. Inflorescences simple or warted, 0.30.5 cm diameter, 26 (
10)-flowered in males; flowers pubescent with persistent hairs up to 0.1 mm long. Male
432
vi
flowers: pedicel 515 mm long; bracteole caducous, attached to pedicel at about the
middle; buds broadly obovoid or depressed globose, (3.5)47 mm diameter, inside pale
pink; perianth at anthesis clefts to 3/44/5, at the sutures 12 mm thick; staminal disc
flattish, including anthers circular to subtrigonous, 24 mm diameter; anthers 1115;
staminal column slender, 12 mm long, clasped by the fleshy base of the perianth. Female
flowers not seen. Fruits 12 per infructescence, ellipsoid to subglobose-obovoid, 34.5
(5) 1.73 cm; dry pericarp 24 mm thick, tomentose with hairs c. 0.2 mm long; stalk
210 mm long.
Ecology. In hill mixed dipterocarp to lower montane forests, at (400)5001600 m altitude.
Flowering and fruiting in JuneOctober.
Key to subspecies
Leaves coriaceous; tomentum of lower surface inconspicuous, subpersistent to glabrescent,
consisting of rather weak scattered minute stellate hairs of various size; intercostal venation
fine and distinct above. Fruit flanged along the line of suture or not..............
subsp. pectinata
Common in Sabah and Sarawak, at 4001500 m altitude.
Leaves rigidly coriaceous; tomentum of lower surface largely persistent, consisting of
densely set, yellowish rusty, minute stellate hairs 0.10.2 mm long, mixed with fewer
emergent ones; intercostal venation distinct on both surfaces. Fruit not
flanged..............................................................................................................................
subsp. vestita W.J. de Wilde
(Latin, vestitus = clothed)
Blumea 27 (1981) 228. Type: Ilias S. 26349, Borneo, Sarawak, Lawas district (holotype L; isotypes
K, SAR, SING).
Twig stout, towards apex 35 mm diameter, rusty puberulous with minute stellate,
scale-like hairs up to 0.1 mm long. Leaves elliptic-oblong to lanceolate, (10)20
30 (3)3.510.5 cm, base rounded to cuneate, apex obtuse to acute-acuminate; lateral
veins 2530 pairs. Fruits 1 per infructescence, 34 2.73 cm, tomentose with hairs
0.10.2 mm long; stalk c. 3 mm long.
Endemic to Borneo: Sarawak (Lawas, Ulu Limbang), at 14001600 m altitude.
29. Knema pedicellata W.J. de Wilde
Fig. 12.
(Latin, pedicellatus = provided with a pedicel; flowers with long pedicels)

Blumea 25 (1979) 474; Whitmore, Tantra & Sutisna l.c. 271. Type: Ilias S. 16626, Borneo, Sarawak,
Miri district, G. Lambir (holotype L; isotypes A, K, SAR).
Tree up to 30 m tall; buttresses up to 50 cm tall, 30 cm wide. Bark smooth or flaky;

inner bark brownish; sap pinkish to dark red. Sapwood whitish. Twigs rather stout, towards
apex terete, 46 mm diameter, at first tomentose with hairs 0.51 mm long, glabrescent;
bark not or only rarely tending to crack or flake. Leaves chartaceous, upper surface
glabrescent except part of the midrib, drying olivaceous-brown, lower surface greenish
434
vi
grey, papillose, with reddish brown dark dots predominantly on the finer veins, subpersistently
tomentose with stout stellate and stellate-dendroid hairs 0.31.2 mm long; elliptic to oblong,
(12)1736 (5)713 cm, base cordate to subcuneate; midrib slightly raised above;
lateral veins 1422 pairs, slightly raised above; petiole late-glabrescent, 0.61.8 cm long.
Inflorescences simple or forked, 0.40.8 cm diameter; 2050-flowered in males, 1040flowered in females; flowers pubescent with hairs 0.10.5(0.8) mm long. Male flowers:
pedicel slender, 2030 mm long; bracteole 1 (or 2), persistent, attached to the pedicel
at about the middle; buds depressed globose, more or less sagged at the base, 34
45 mm, inside glabrous, reddish?; perianth (3)45-lobed, at anthesis clefts to 3/4
4/5, at the base inside with a conspicuous disc surrounding the staminal column; staminal
disc distinctly mammillate, including anthers circular, 23 mm diameter; anthers 1418;
staminal column 0.61 mm long. Female flowers: pedicel 1012 mm long; buds ellipsoid
obovoid, c. 6 mm long, inside with a disc at the base; perianth lobes 45. Fruits 1
4 per infructescence, subglobose, 23 2.52.8 cm, usually somewhat broader than long,
tomentose with hairs 0.30.5 mm long; dry pericarp 45 mm thick; stalk 1830 mm long.
Ecology. In lowland and hill mixed dipterocarp forests on sandstone-derived, brownish
soils, sandy soils, or sandy clay soils derived from granite, up to 700 m altitude. Flowering
in MayJuly; fruiting in NovemberDecember.
30. Knema percoriacea J. Sinclair

(Latin, per- = exceedingly, coriaceous = leathery; the thick-leathery leaves)
Gard. Bull. Sing 18 (1961) 268; W.J. de Wilde l.c. (1979) 392, l.c. (1996) 392; Anderson l.c. 268;
Whitmore, Tantra & Sutisna l.c. 271; Argent et al. (eds.) l.c. 456. Type: Kostermans 9945, Borneo, E
Kalimantan, Loa Djanan, W Samarinda (holotype K; isotypes BO, L, SING).
Tree up to 25 m tall. Bark smooth, fissured, scaly or flaky; inner bark inside red-brown
or pinkish. Sapwood usually white, sometimes reddish or red-brown. Twigs stout, subterete,
towards apex 36 mm diameter, at first tomentose with hairs 0.51.5(2) mm long, lateglabrescent; bark greyish brown, slightly to distincly longitudinally cracked and flaky.
Leaves chartaceous to coriaceous, upper surface drying olivaceous-brown, lower surface
glabrous, without blackish dots; oblong-lanceolate, 1240 2.59(15) cm, base attenuate
to rounded, apex sometimes subobtuse; midrib prominently raised above; lateral veins15
25 pairs, raised above; petiole sometimes somewhat pulvinate, late-glabrescent, 0.72.5
cm long. Inflorescences simple or 2-furcate, up to 1.5 cm diameter, 1020-flowered in
males; peduncle up to 2 mm long; flowers tomentose with persistent hairs 0.52 mm
long, sometimes partially glabrescent. Male flowers: pedicel 34.5 mm long; bracteole
usually persistent, attached to the pedicel at or above the middle; buds depressed globoseobovoid, (3.5)45.5 (3.5)45.5 mm, inside red; perianth at anthesis clefts to 2/3
3/4; staminal disc circular to subtriangular, flat, including anthers 22.5 mm diameter;
anthers 1018, subsessile. Female flowers: pedicel 24 mm long; buds ovoid-ellipsoid,
6.58(9) mm long. Fruits 12 per infructescence, broadly ellipsoid, 34.5 2.53.5
cm, tomentose with hairs 12 mm long; dry pericarp c. 4 mm thick; stalk 23 mm long.
Distribution. Endemic to Borneo (Sabah, Sarawak, Brunei, NE and E Kalimantan).
436
Ecology. In primary and degraded mixed dipterocarp forests, on hill sides and ridges,
on sandy, clayey and sandy loam soils, up to 700 m altitude. Flowering and fruiting
mainly in JulyNovember.
Notes. A variable species, in which three forms are distinguished. Two forms occur in
Sabah and Sarawak, mostly found in kerangas forest.
Key to forms
Leaves chartaceous; reticulation of intercostal venation distinct on both surfaces. Tomentum
of twig towards apex pale brown or yellowish brown, comprising hairs 0.31 mm long.
Anthers 1114................................
forma sarawakensis W.J. de Wilde
(of Sarawak)
Blumea 25 (1979) 393; Coode et al. (eds.) l.c. 226. Type: Wright S. 23889, Borneo, Sarawak, Kapit
district (holotype L; isotypes A, BO, K, KEP, MEL, SAN, SAR, SING).
Twigs towards apex 46 mm diameter, coarsely striate; bark lower down distinctly
and coarsely flaky.
Endemic to Borneo: Sarawak, Brunei, Sabah, NE and C Kalimantan (Nunukan Is.).
Leaves coriaceous; reticulation of intercostal venation on both surfaces indistinct. Tomentum
of twig towards apex chocolate to dark brown, comprising hairs 12 mm long. Anthers
15.........................................................................................................
forma fusca W.J. de Wilde
(Latin, fuscus = dark brown; the indumentum)
Blumea 25 (1979) 394; Coode et al. (eds.) l.c. 225. Type: Ashton BRUN 5164, Borneo, Brunei
(holotype L; isotypes BO, K).
Twig towards apex c. 6 mm diameter, flaky, dark brown tomentose with hairs c. 1.5 mm
long.
Endemic to Borneo: Sarawak and Brunei.
31. Knema piriformis W.J. de Wilde

(Latin, pirus = pear; forma = shape; the pear-shaped male flower buds)
Blumea 25 (1979) 400; Whitmore, Tantra & Sutisna l.c. 271. Type: Meijer SAN 34619, Borneo, Sabah,
Ranau district (holotype L; isotypes K, SAN).
Tree up to 12 m tall. Twigs somewhat angular, towards apex 1.53 mm diameter, drying
brown or blackish, tomentose with hairs c. 0.1(0.2) mm long, glabrescent; bark not tending
to crack or flake. Leaves thinly coriaceous, upper surface drying olivaceous, with scattered
hair scars, lower surface without blackish dots, tomentose with soft greyish hairs c. 0.1
mm long, early to rather late-glabrescent, sometimes mixed with larger dendroid hairs
which are early shed and leaving scars; oblong to lanceolate, 1525 37 cm, base
rounded to attenuate; midrib sunken to slightly raised above; lateral veins 2025 pairs,
sunken to flattish above; petiole 0.81.5 cm long. Inflorescences sessile, simple or forked,
up to 0.6 cm diameter, 510-flowered in males; flowers persistently pubescent with hairs
0.10.2 mm long. Male flowers: pedicel 25 mm long; bracteole minute, caducous, situated
on the pedicel at or above the middle; buds pear-shaped, 45 34 mm, tapering in
437
the lower half, inside yellow; perianth at the sutures up to 1 mm thick, at anthesis clefts
to 1/22/3; staminal disc flat, including anthers circular, 1.52 mm diameter; anthers
1112; staminal column slender, 1.52.2 mm long. Female flowers not seen. Fruits 1
2 per infructescence, ovoid to ellipsoid, 2.42.6 1.51.7 cm, pubescent with persistent
grey hairs c. 0.1 mm long; dry pericarp c. 2 mm thick; stalk c. 6 mm long.
Distribution. Endemic to Borneo (Sabah and Sarawak; mainly found in Mt. Kinabalu
and its vicinity).
Ecology. In hill mixed dipterocarp to montane forests, at (400?)10002000 m altitude.
Flowering in AprilJuly; fruiting in January.
32. Knema psilantha W.J. de Wilde

(Greek, psilo = bare or smooth, anthos = flower)
Blumea 25 (1979) 391, l.c. (1996) 392; Whitmore, Tantra & Sutisna l.c. 271. Type: Anderson S. 14373,
Borneo, Sarawak, Kapit district (holotype L; isotypes K, SAR).
Tree up to 10 m tall. Bark smooth, finely lenticellate, brittle; inner bark white mottled
with reddish. Twigs fairly stout, sometimes angled, towards apex 35 mm diameter,
tomentose with woolly yellowish brown hairs 0.51 mm long, soon glabrescent; bark
cracked and flaky. Leaves thinly coriaceous, upper surface drying greenish brown, lower
surface glabrous, finely papillose, without blackish dots; lanceolate, 2048 3.59.5
cm, base obtuse to broadly rounded; midrib raised above; lateral veins 3035 pairs, raised
above; petiole late-glabrescent, 0.51.5 cm long. Inflorescences sessile, simple or warted,
0.20.5 cm diameter, 525-flowered in males, 15-flowered in females; flowers tomentose
with pale hairs 0.50.7 mm long, early glabrescent in the upper half or more. Male
flowers: pedicel 23 mm long; bracteole subpersistent, attached about halfway or on
the upper part of the pedicel; buds ellipsoid-obovoid, 4.55 33.5 mm, inside red;
perianth at anthesis clefts to 3/44/5; staminal disc flattish, including anthers circular,
c. 1.5 mm diameter; anthers 67, stiped; staminal column 11.2 mm long. Female flowers:
pedicel 12 mm long; buds larger than that of male flowers, ovoid, 89 mm long. Fruits
13 per infructescence, subellipsoid, 2.53 2 cm, pubescent with persistent rough
yellowish hairs c. 1 mm long; dry pericarp c. 3 mm thick; stalk c. 2 mm long.
Ecology. In mixed dipterocarp forest on ridges and hill sides, at 50300 m altitude.
Flowering in AugustSeptember; fruiting in June.
33. Knema pubiflora W.J. de Wilde

(Latin, pubeus = soft-hairy, florus = flower; flowers hairy inside)
Blumea 25 (1979) 466; Whitmore, Tantra & Sutisna l.c. 271; Argent et al. (eds.) l.c. 456. Type: Meijer
SAN 25105, Borneo, Sabah, Beluran district (holotype L; isotypes BO, L, SAN, SING).
438
Tree up to 40 m tall; buttresses up to 2 m high, extending to c. 30 cm. Bark smooth,

or when old peeling off in soft irregular small scales; inner bark chocolate, or brown,
or reddish; sap reddish. Sapwood reddish white, or pale brown, or pale pinkish. Twigs
towards apex 1.53 mm diameter, at first tomentose with scurfy hairs 0.10.2 mm long,
glabrescent; bark not tending to crack or flake. Leaves chartaceous to thinly coriaceous,
upper surface drying dark brown, lower surface grey-brown, papillose, with scattered
dark dots especially on the veinlets, at first tomentose with mixed sessile and dendroid
hairs 0.20.5 mm long, late-glabrescent or persistent; ovate-oblong to oblong, 415
1.57 cm, base rounded to subattenuate; midrib slightly raised above; lateral veins 7
14 pairs, slightly raised above; petiole late-glabrescent, 0.51 cm long. Inflorescences
simple or forked, up to 0.4 cm diameter; 520-flowered in males, 310-flowered in females;
peduncle up to 2 mm long; flowers outside brown tomentose with hairs c. 0.1 mm long.
Male flowers: pedicel 1015 mm long; bracteole persistent or late-caducous, attached
to the pedicel at above the middle; buds depressed globose, 22.5 2.53 mm, inside
densely hairy; perianth at anthesis clefts to 3/4 or 4/5; staminal disc convex, including
anthers circular, 1.21.5 mm diameter; anthers 1113; staminal column 0.30.6 mm long.
Female flowers: pedicel 710 mm long; buds obovoid, 34 mm long. Fruits 15 per
infructescence, subglobose to shortly ellipsoid, 1.52.2(2.5) 1.52.3 cm, often ridged
at one side, tomentose with hairs 0.10.2 mm long; dry pericarp c. 1.5 mm thick; stalk
1014 mm long.
Distribution. Endemic to Borneo (Sabah, Sarawak and Kalimantan, including Nunukan
Is.).
Ecology. In mixed dipterocarp forests on sandy, sandy clay, or sandy loam soils, on hill
sides and ridges, up to 300 m altitude, scattered but locally common. Flowering and
34. Knema pulchra (Miq.) Warb.

(Latin, pulcher = beautiful; inference unclear)
Monogr. Myrist. (1897) 600; Merrill l.c. (1921) 271; Masamune l.c. 303; W.J. de Wilde l.c. (1979) 399,
l.c. (1981) 224; Whitmore, Tantra & Sutisna l.c. 271; Turner l.c. 361; Coode et al. (eds.) l.c. 226.
Basionym: Myristica pulchra Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 51. Type: Korthals s.n., Borneo,
Kalimantan (L). Synonyms: K. glaucescens Jack var. cordata J. Sinclair l.c. (1958) 310; K. cinerea
(Poir.) Warb. var. cordata (J. Sinclair) J. Sinclair l.c. (1961) 181.
Tree up to 15 m tall; stilt roots sometimes present. Bark with a few lenticels, not flaking
or pustulate, or finely fissured, or smooth; inner bark orange-brown. Sapwood pale brown
or white. Twigs towards apex 36(8) mm diameter, tomentose with short hairs c. 0.1
mm long, glabrescent; bark not tending to crack or flake, drying brownish or blackish.
Leaves chartaceous to thinly coriaceous, upper surface drying olivaceous-brown, often
glossy, lower surface greyish, without blackish dots, with persistent or deciduous tomentum
of scattered pale stellate hairs 0.10.2 mm long; elliptic-oblong to oblong-lanceolate,
(20)3050 (6)1017 cm, base rounded to cordate; midrib raised above; lateral veins
1825 pairs, flat or hardly raised above; petiole stout, 1.54 cm long. Inflorescences
sessile, simple or 23-furcate, up to 1 cm diameter, 110-flowered in males, 15-flowered
in females; flowers tomentose with greyish hairs up to 0.1 mm long; perianth at the
439
sutures up to 1 mm thick. Male flowers: pedicel 48 mm long; bracteole caducous or

subpersistent, situated at about or above the middle of the pedicel; buds pear-shaped,
(3)45.5 35 mm, inside cream or pale brown; perianth at anthesis clefts to 1/2
3/4; staminal disc flat to concave, including anthers circular to faintly triangular, 1.8
3 mm diameter; anthers 1225; staminal column 0.81.5 mm long. Female flowers: pedicel
(2)58 mm long; buds obovoid-oblong, 56 mm long. Fruits 12 per infructescence,
subglobose to ellipsoid-oblong, (3)46 23.5 cm, apex rounded, glabrescent or
tomentose with hairs (0.1)0.2 mm long; dry pericarp 34 mm thick; stalk (2)512 mm
long.
Distribution. Peninsular Malaysia and Borneo (Sabah, Sarawak, Brunei, and Kalimantan).
Ecology. In primary lowland and hill mixed dipterocarp forests, also in swamp forest,
on sandy, loamy and basalt-derived soils, up to 1000 m altitude. Flowering in May
September; fruiting in MarchAugust.
35. Knema riangensis W.J. de Wilde

(of the Riang R., C Kalimantan)
Blumea 32 (1987) 138; Whitmore, Tantra & Sutisna l.c. 272. Type: Veldkamp 7941, Borneo, C
Kalimantan, Upper Samba R., Bt. Raya area, estuary of Riang R. (holotype L; isotype BO).
Tree up to 12 m tall. Twigs towards apex terete, 1.53(4) mm diameter, more or less
angular, drying brownish black, tomentose with minute, yellowish or greyish white hairs
c. 0.1 mm long; bark not flaky. Leaf buds slender, 1020 mm long, tomentose with hairs
c. 0.1 mm long. Leaves membranous, upper surface early glabrescent, drying olivaceous,
lower surface greyish, without blackish dots, with subpersistent tomentum consisting of
densely set, almost touching, pale stellate, yellowish or greyish white hairs c. 0.1(0.2)
mm long; elliptic-oblong to oblong-lanceolate, 1430 510.5 cm, base short-attenuate
to rounded; midrib raised above; lateral veins 1523 pairs, flattish to raised above; petiole
late-glabrescent, 12 cm long. Inflorescences simple or forked, up to 0.5 cm diameter,
stalks up to 1 mm long, in males 26-flowered, in females 24-flowered; perianth at
the sutures up to 1 mm thick. Male flowers tomentose with yellowish or greyish white
hairs c. 0.1(0.2) mm long; pedicel 34 mm long; bracteole persistent, c. 0.5 mm long,
attached to the pedicel below the perianth; buds broadly obovoid, c. 2 1.7 mm, inside
pink; perianth at anthesis clefts to 5/6; staminal disc subcircular to faintly triangular,
flat above, including anthers 0.81 mm diameter; anthers 9; staminal column tapering
(obconical), c. 0.8 mm long, glabrous or with a few minute hairs at the base. Female
flowers rather stout, pedicel 512 mm long; buds almost obovoid, 47 mm long. Fruits
1 or 2 per infructescence, ellipsoid, 2.44 1.72.5 cm, tomentose with persistent yellowish
or greyish white hairs c. 0.1 mm long; dry pericarp 12 mm thick; stalk 812 mm long.
Seeds ellipsoid, 22.5 cm long.
Distribution. Endemic to Borneo (Sarawak and C Kalimantan).
Ecology. In primary riverine mixed dipterocarp forest on yellow sandy clay soils, at 100
600 m altitude. Flowering in JanuaryFebruary; fruiting in May.
440
36. Knema rufa Warb.

(Latin, rufus = reddish; the indumentum)
Monogr. Myrist. (1897) 556; Merrill l.c. (1921) 271; Masamune l.c. 304; J. Sinclair l.c. (1961) 274,
288; W.J. de Wilde l.c. (1979) 409; Anderson l.c. 268; Coode et al. (eds.) l.c. 226. Type: Beccari 1798,
Borneo, Sarawak, Mt. Matang (holotype FI; isotypes G, K, P).
Tree up to 20 m tall. Twigs towards apex 23 mm diameter, slightly angular, tomentose

with hairs c. 0.3 mm long, glabrescent; bark not or faintly cracked or flaky. Leaves
thinly coriaceous, on both surfaces at first densely rusty pubescent with hairs c. 0.5 mm
long, early glabrescent, upper surface drying olivaceous to blackish brown, lower surface
without blackish dots; oblong to lanceolate, 1530 1.54(5.5) cm, base attenuate; midrib
raised above; lateral veins 1525(30) pairs, prominently raised above, usually with lesser
side-veins in between; petiole 12 cm long. Inflorescences simple, up to 1 cm diameter;
210-flowered in males; peduncle up to 2 mm long; flowers pubescent with hairs c. 0.3
mm long. Male flowers: pedicel 1015 mm long; bracteole caducous, attached to the
pedicel above the middle; buds depressed obovoid, 67 56 mm, often somewhat
furrowed, inside red (always?); perianth at anthesis clefts to c. 2/3; staminal disc circular,
concave, including anthers 22.5 mm diameter; anthers (9) 1015, connectives protrude
slightly beyond the thecae; staminal column tapering, c. 1 mm long. Female flowers
not seen. Fruits 1(3) per infructescence, ellipsoid, c. 2.5 1.5 cm, tomentose with shaggy
hairs c. 3 mm long; pericarp 2(3) mm thick; stalk 1020 mm long.
Ecology. In primary lowland mixed dipterocarp forest on sandy clay soils, up to 200
m altitude.
37. Knema sericea W.J. de Wilde

(Latin, sericeus = silky; the indumentum on the lower leaf surface)
Blumea 25 (1979) 383, l.c. (1996) 394; Whitmore, Tantra & Sutisna l.c. 272; Coode et al. (eds.) l.c. 226.
Type: Jaheri 611, Borneo, NE Kalimantan (holotype L; isotypes BO, SING).
Tree up to 20 m tall. Bark smooth. Twigs towards apex 45 mm diameter, pubescent

with dense scale-like hairs up to 0.1 mm long, glabrescent; bark not tending to crack
or flake. Leaves membranous to chartaceous, upper surface drying greenish to blackish,
lower surface silky to the touch, tomentose with short, dense, greyish or pale cinnamon,
interwoven hairs c. 0.2 mm long, mixed with sparse stouter dendroid hairs, without blackish
dots; oblong to oblong-lanceolate, 3045 915.5 cm, base rounded to shallowly cordate,
apex rounded or acute-acuminate; midrib raised above; lateral veins 3035 pairs, raised
above, basal veins somewhat crowded; intercostal venation reticulate, coarse above; petiole
stout, 1.22 cm long. Inflorescences c. 0.5 cm diameter, with stout peduncle up to 7
mm long; 410-flowered in males; flowers densely tomentose with hairs c. 0.1 mm long.
Male flowers: pedicel 1218 mm long; bracteole caducous, attached at the top of pedicel;
buds subglobose, 610 mm diameter; perianth at anthesis clefts to c. 4/5; staminal disc
convex to mammillate, including anthers circular, c. 3 mm diameter; anthers 1015, stiped;
staminal column c. 0.5 mm long. Female flowers not known. Fruits 12 per infructescence,
441
subglobose to broadly ovoid, 3.55 3.53.8 cm, pubescent with scale-like hairs c. 0.1
mm long; dry pericarp c. 5 mm thick; stalk 1520 mm long, with bracteole scars attached
at or somewhat above the middle.
Distribution. Endemic to Borneo (Sarawak, Brunei and NE Kalimantan).
Ecology. In forest at the foot of limestone hill, at 50350 m altitude. Fruiting in January.
38. Knema stenophylla (Warb.) J. Sinclair

(Greek, stenos = narrow, phullon = leaf; narrow-leaved)
Gard. Bull. Sing. 16 (1958) 300, l.c. (1961) 276; W.J. de Wilde l.c. (1979) 457, p.p., excl. stouter forms
from Borneo, now in K. subhirtella, l.c. (1987) 129; Anderson l.c. 268; Whitmore, Tantra & Sutisna l.c.
272; Turner l.c. 361. Basionym: Gymnacranthera stenophylla Warb. l.c. (1897) 364. Type: Kehding
150, Peninsular Malaysia, Klang (in Herb. Beccari 7714; FI). Synonym: Myristica stenophylla (Warb.)
Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 88.
There are two subspecies; subsp. stenophylla occurs in Peninsular Malaysia and Sumatra,
and subsp. longipedicellata in Borneo. The latter differs from the former by its longer
pedicel of the male flowers (710 mm long) and fruits stalk (1020 mm long).
subsp. longipedicellata (Sinclair) W.J. de Wilde
(flowers with long pedicels)
Blumea 32 (1987) 130; Coode et al. (eds.) l.c. 226. Basionym: K. cinerea (Poir.) Warb. var. patentinervia
J. Sinclair forma longipedicellata J. Sinclair l.c. (1961) 182. Type: Clemens 20345, Borneo, Sarawak,
Mukah district, G. Puai (holotype K; isotypes A, BO, G, L, NY, SAR).
Tree up to 25 m tall. Bark smooth or rough with some pustules or flakes when old,
not striate or furrowed; inner bark reddish, granular. Sapwood white or cream. Twigs
towards apex terete or subangular, slender, up to 1 mm diameter, drying brown or blackish,
at first tomentose with hairs up to 0.1 mm long, sometimes mixed with longer hairs
0.20.3 mm long, glabrescent; bark not tending to crack or flake. Leaves membranous
to chartaceous, upper surface drying olivaceous to brown, lower surface greyish, without
dark dots, with subpersistent tomentum of scattered mainly stellate, scale-like hairs up
to 0.1 mm long; elliptic to lanceolate, (3)518 15 cm, base cuneate; midrib flattish
to raised above; lateral veins 1021 pairs, sunken or slightly raised, usually indistinct
above; intercostal venation coarse above; petiole 0.61.6 cm long. Inflorescences simple
or 23-fid, 0.10.3(0.7) cm diameter; (1)310-flowered in males, 16-flowered in females;
peduncle up to 1 mm long; flowers tomentose with persistent hairs up to 0.1(0.3) mm
long; perianth at the sutures up to 1 mm thick. Male flowers: pedicel 710 mm long;
bracteole persistent or caducous, attached to the pedicel at about the middle to near
the apical part; buds broadly obovoid, (1.5)23(4?) mm diameter, inside cream or
yellowish; perianth at anthesis clefts to 3/44/5; staminal disc flattish, including anthers
circular to subtriangular, 12.2 mm diameter; anthers 611; staminal column c. 1 mm
long. Female flowers: pedicel 812 mm long; buds ellipsoid to obovoid, 3.55 mm long.
Fruits 15 per infructescence, subglobose or obovoid, (1.2)1.5 0.71 cm, apex and
442
base rounded or sometimes somewhat attenuate, pubescent with persistent scale-like hairs
up to 0.1 mm long; dry pericarp c. 1 mm thick; stalk 1020 mm long.
Ecology. In primary and degraded lowland mixed dipterocarp to lower montane forests,
often found on hill sides, crests, ridges, occasionally in swamp and kerangas forest, on
various types of soils including, loamy, sandy, clayey, and sandstone and shale-derived
soils, up to 1200 m altitude. Flowering mainly in MarchJuly; fruiting mainly in July
December.
39. Knema stylosa (W.J. de Wilde) W.J. de Wilde

(Latin, stylosus = with a persistent style; the fruit)
Blumea 32 (1987) 119; Whitmore, Tantra & Sutisna l.c 272. Basionym: K. hirtella W.J. de Wilde var.
stylosa W.J. de Wilde l.c. (1979) 460. Type: Mikil SAN 30179, Borneo, Sabah, Beaufort district (holotype
L; isotype SAN).
Tree up to 25 m tall. Bark rather smooth; inner bark pinkish; sap dark red. Sapwood
whitish. Twigs towards apex subterete, 23 mm diameter, at first tomentose with hairs
up to 0.1 mm long, glabrescent; bark drying dark brown, not tending to crack or flake.
Leaves thinly chartaceous, upper surface drying olivaceous, lower surface without dark
dots, greyish, early glabrescent or with subpersistent tomentum of scattered, minute hairs
less than 0.1 mm long, sparsely mixed with scars of coarser caducous hairs c. 0.2 mm
long; elliptic-oblong to oblong, 1224 3.57 cm, base subcordate or rounded to shortattenuate; midrib convex above, but as a whole generally partly sunken under surface
level; lateral veins 1520 pairs, flattish or sunken above; intercostal venation coarse above;
petiole 12 cm long, late-glabrescent. Inflorescences sessile, wart-like, simple or forked,
up to 0.4 cm diameter; 36(10)-flowered in males and females; flowers tomentose with
hairs c. 0.1 mm long, perianth at the sutures up to 1 mm thick. Male flowers: pedicel
45 mm long; bracteole apical, persistent; buds obovoid, 3.54 33.5 mm; perianth
at anthesis clefts to c. 2/3; staminal disc flattish, bluntly triangular, including anthers
c. 1.4 mm diameter; staminal column tapering, c. 1.2 mm long, glabrous; anthers 11.
Female flowers: pedicel 35 mm long; buds obovoid-ellipsoid, c. 4.5 mm long. Fruits
solitary or 23 per infructescence, ovoid, 2.53.5 1.82.5 cm, tomentose with persistent
hairs up to 0.1 mm long, sometimes glabrescent; style 0.51 mm long, persistent or leaving
a distinct scar; dry pericarp 1.52.5 mm thick; stalk 36 mm long.
Distribution. Endemic to Borneo (Sabah and NE Sarawak).
Ecology. In primary mixed dipterocarp forest on hill sides, often on blackish sandstonederived soils, up to 800 m altitude. Flowering in June; fruiting throughout the year.
40. Knema subhirtella W.J. de Wilde

(Latin, sub- = somewhat, hirtellus = finely hairy; the twigs and leaves)
Blumea 32 (1987) 131; Whitmore, Tantra & Sutisna l.c. 272; Coode et al. (eds.) l.c. 226. Type: Yii et al.
443
Tree up to 25 m tall. Bark dark brown or chocolate-brown, smooth or dippled, not furrowed,
brittle; inner bark pink-cream, fibrous; sap pink to blood-red. Sapwood cream. Twigs
bluntly triangular, towards apex 1.52(2.5) mm diameter, pubescent with hairs c. 0.1
mm long, mixed with sparse hairs up to 0.3 mm long, glabrescent; bark drying brown
or blackish, not flaky. Leaf buds tomentose with hairs 0.1(0.3) mm long. Leaves
membranous, upper surface drying olivaceous, lower surface greyish, without dark dots,
with persistent tomentum of scattered stellate-dendroid hairs 0.10.2(0.3) mm long;
elliptic-oblong to lanceolate, 920 (1.5)25.5 cm, base cuneate; midrib raised above;
lateral veins 1525 pairs, flattish to raised above; intercostal venation coarse and distinct
above; petiole 0.81.2 cm long. Inflorescences simple or forked, up to 0.4 cm diameter;
peduncle up to 1 mm long; in males 47-flowered, in females 14-flowered; perianth
at the sutures up to 1 mm thick. Male flowers tomentose with hairs 0.10.2(0.3) mm
long; pedicel 25 mm long; bracteole late-caducous, attached to the pedicel at 0.51.5
mm below the perianth; buds broadly obovoid, 2.53.5(4) 2.53.5 mm; perianth at
anthesis clefts to 1/23/4; staminal disc circular, flat or somewhat convex, including anthers
1.21.5(2) mm diameter; anthers (7)913(15); staminal column 11.5 mm long,
(almost)glabrous. Female flowers: pedicel 47(10) mm long; buds obovoid, c. 5 mm
long. Fruits solitary or 2 per infructescence, broadly ellipsoid, 22.7 1.52 cm, tomentose
with persistent hairs c. 0.1 mm long; dry pericarp c. 1.5 mm thick; stalk 410 mm long.
Ecology. In mixed hill dipterocarp forest on slopes and ridges, on sandy clay, loamy,
and basalt- or shales-derived soils, up to 800(1200) m altitude. Flowering in March
May(August); fruiting mainly in SeptemberOctober.
41. Knema tridactyla Airy Shaw
Fig. 13.
(Greek, tris = three, daktylos = finger; with 3-forked inflorescences)

Kew Bull. 1939, no. 10 (1940) 543; Masamune l.c. 304 (K. tridactyia); W.J. de Wilde l.c. (1979) 434,
l.c. (1998) 250; Coode et al. (eds.) l.c. 226. Type: Richards 1220, Borneo, Sarawak, Mt. Dulit (holotype
K).
Tree up to 10 m tall. Bark brown, smooth. Twigs towards apex terete or faintly angular,
1.54 mm diameter, pubescent with woolly, dendroid hairs 0.51(1.5) mm long, lateglabrescent, or with inconspicuous scale-like hairs up to 0.1 mm long and rather early
glabrescent; bark drying brown or blackish, not tending to crack or flake. Leaves
chartaceous to thinly coriaceous, upper surface drying olivaceous or brown, often shining
and with hair scars, lower surface greyish, usually with scars of coarse hairs, without
dark dots, and either with subpersistent tomentum of sessile-stellate hairs 0.30.5 mm
long, intermixed with stronger stellate-dendroid hairs, or at first tomentose with remote
stellate, scale-like hairs up to 0.1 mm long, early glabrescent; oblong to lanceolate, 7
28 25.5(7.5) cm, base cuneate to rounded; midrib prominently raised above, in subsp.
tridactyla on the lower side with characteristic appressed scale-like hairs; lateral veins
1023 pairs, flat or sunken above; petiole 0.51.5 cm long, tomentose with scale-like
hairs, late-glabrescent. Inflorescences slender, up to 1.7 cm diameter, sessile or with
peduncles up to 3 mm long, simple or 23-fid, branches 12(2.5) mm thick; (2)4
444
vi
10-flowered in males (with in addition numerous scars of pedicels), 15-flowered in females;

flowers pubescent with dendroid hairs 0.30.6 mm long or scale-like hairs c. 0.1 mm
long. Male flowers: pedicel 1.53.5 mm long; bracteole subpersistent, attached to the
pedicel at or above the middle; buds globose to broadly obovoid, 1.52(2.3) 1.5
2.5 mm, inside pinkish (?); perianth at anthesis clefts to 3/44/5; staminal disc flat, including
anthers circular or faintly triangular, 0.61.2 mm diameter; anthers 511; staminal column
0.30.7 mm long. Female flowers: pedicel c. 1.5 mm long; buds obovoid, 22.5 mm
long. Fruits 1(2) per infructescence, ellipsoid or obovoid to ovoid-oblong, 1.42 0.8
1.6 cm, pubescent either with dendroid hairs 0.20.5 mm long or scale-like hairs c. 0.1
mm long; dry pericarp 11.5 mm thick; stalk 23 mm long.
Ecology. In primary lowland and hill mixed dipterocarp forests on clay and yellow sandy
soils, up to 1000 m altitude. Flowering throughout the year; fruiting in October, December,
and February.
Key to subspecies
1.
Tomentum of twig towards apex, flowers, and fruits consisting of scale-like hairs
c. 0.1 mm long...............
subsp. sublaevis W.J. de Wilde
(Latin, sub- = somewhat, laevis = smooth; the plant nearly devoid of hairs)
Blumea 25 (1979) 434, l.c. (1996) 395, l.c. (1998) 251; Coode et al. (eds.) l.c. 226. Type: Ilias
Sarawak and Brunei.

Tomentum of twig towards apex, flowers, and fruits consists of dendroid hairs 0.5
1(1.5) mm long...........2
2.
Leaves 712 1.52.5 cm; lateral veins 812 pairs. Anthers 6.......
subsp. salicifolia W.J. de Wilde
(Latin, with leaves like those of Salix)
Blumea 43 (1998) 251. Type: Wood SAN A 4237, Borneo, Sabah (holotype L; isotype SAN).
Sabah, Kota Belud district.

Leaves larger; lateral veins 1223 pairs. Anthers 711....3
3.
Hairs 1(1.5) mm long. Branches of male inflorescences 22.5 mm thick. Anthers

1011...............................
subsp. pachydactyla W.J. de Wilde
(Greek, pachus- = thick; dactylos = finger; the thick branches of the inflorescence)
Blumea 43 (1998) 251. Type: Leopold et al. SAN 134086, Borneo, Sabah (holotype L; isotype
SAN).
Sabah, Penampang district.

Hairs 0.51 mm long. Branches of male inflorescences (1)1.52 mm thick. Anthers
78............
subsp. tridactyla
W.J. de Wilde l.c. (1979) 434; Coode et al. (eds.) l.c. 226.
Sabah, Sarawak and Brunei.
446
42. Knema uliginosa J. Sinclair

(Latin, uliginosus = growing in marshes)
Sutisna l.c. 272. Type: Main 2098 (exp. Polak), Borneo, W Kalimantan, Pulau Nibung, Silimban, Kapuas
(holotype K; isotypes A, L, SING).
Shrub or tree up to 10 m tall. Twigs bluntly 23-angular, towards apex 1.53(4) mm

diameter, tomentose with hairs 0.51 mm long, glabrescent; bark drying brown or blackish,
not tending to crack or flake. Leaves thinly coriaceous, upper surface drying olivaceousbrown, lower surface at first pubescent with dense hairs 0.51 mm long, early glabrescent,
without dark dots; oblong to lanceolate, 935 27.5 cm, base rounded to attenuate;
midrib raised above; lateral veins 815 pairs, prominently raised above; intercostal venation
raised and distinct above; petiole c. 1 cm long. Inflorescences sessile, simple or forked,
up to 0.5 cm diameter, 520-flowered in males, 15-flowered in females; flowers pubescent
with persistent hairs 0.30.4 mm long; perianth at the sutures up to 1 mm thick. Male
flowers: pedicel 510 mm long; bracteole subpersistent, attached near the apex of pedicel;
buds depressed globose, faintly trigonous in transverse section, 34 45 mm, inside
reddish (?); perianth at anthesis clefts to c. 4/5; staminal disc convex or low-mammillate,
including anthers circular, 22.5 mm diameter; anthers 1013, stiped; staminal column
0.51 mm long. Female flowers: pedicel 45 mm long; buds obovoid, c. 5 mm long.
Fruits 13 per infructescence, broadly obovoid, c. 1.8 1.5 cm, pubescent with hairs
c. 0.3 mm long; stalk 46 mm long.
Vernacular name. Sarawakkumpang kabang (Iban, preferred name).
Distribution. Endemic to Borneo (Sarawak and W Kalimantan).
Ecology. In freshwater and peatswamp forests, up to 300 m altitude. Flowering in October.
43. Knema viridis W.J. de Wilde
Fig. 14.
(Latin, viridis = green; leaves drying green)

Blumea 32 (1987) 121; Whitmore, Tantra & Sutisna l.c. 272. Type: Yii S. 37882, Borneo, Sarawak
(holotype L; isotype SAR).
Tree up to 30 m tall. Bark smooth, or slightly scaly, or with lenticels, brown or blackish;
inner bark soft; sap red. Twigs towards apex (2)2.54.5 mm diameter, bluntly 3-angular,
tomentose with pale hairs 0.20.3 mm long, early glabrescent; bark drying pale brown,
neither cracked or flaky. Leaves chartaceous, upper surface drying greenish, lower surface
early glabrescent, without dark dots, greyish green; oblong to oblong-lanceolate, 1230
3.510.5 cm, base attenuate to rounded, apex blunt to acute; midrib raised on both
surfaces; lateral veins 1423 pairs, raised on both surfaces; intercostal venation very distinct
on both surfaces; petiole 1.21.8 cm long. Inflorescences wart-like, simple or bifurcate,
0.20.5 cm diameter, peduncle up to 2 mm long; in males 812-flowered, in females
47-flowered. Male flowers tomentose with hairs c. 0.1(0.2) mm long; pedicel stout,
710 mm long, at above the bracteole scar distinctly thicker than the lower portion; bracteole
caducous, leaving a distinct scar on the pedicel at about halfway; buds depressed globose,
447
vi
4.55 67 mm, faintly trigonous in transverse section; perianth at anthesis clefts to

3/44/5; staminal disc flattish or convex in the centre, bluntly trigonous in outline, 2.5
3 mm diameter; anthers 2530, closely set; staminal column tapering, 11.5 mm long.
Female flowers (slightly immature): pedicel stout, 1517 mm long; buds depressed globose,
c. 3.5 5 mm. Fruits 13 per infructescence, ellipsoid, 56 44.5 cm, early glabrescent;
dry pericarp c. 10 mm thick; stalk c. 15 mm long.
Ecology. In lowland and hill mixed dipterocarp forests on gentle and wet slopes, at 50
600 m altitude. Flowering in AugustNovember; fruiting in September.
44. Knema woodii J. Sinclair

(G.H.S. Wood, 19271957, formerly forest botanist at the Forestry Department, Sandakan,
Sabah)
Gard. Bull. Sing 18 (1961) 283; W.J. de Wilde l.c. (1979) 429, l.c. (1987) 125; Anderson l.c. 268;
Whitmore, Tantra & Sutisna l.c. 272. Type: Sinclair & Kadim 9298, Borneo, Sabah, Sepilok FR, Compt.
15 (holotype SING; isotypes A, B, BM, E, K, L, M, SAN).
Tree up to 30 m tall. Bark grey or brown, flaking in thin strips or pieces; inner bark
810 mm thick, red or pale red. Sapwood reddish or dirty white. Twigs towards apex
2.54 mm diameter, more or less 3-angled or ridged, often blackish, pubescent with hairs
0.10.2 mm long, early glabrescent; bark not tending to crack or flake. Leaves thinly
coriaceous, upper surface drying olivaceous-brown, lower surface greyish, without blackish
dots, tomentose with minute hairs 0.10.2 mm long, early glabrescent; oblong to
(ob)lanceolate, 1535 (2)49 cm, base rounded to cuneate; midrib raised or not above;
lateral veins 2230 pairs, sunken above; petiole 12 cm long. Inflorescences sessile,
simple or wart-like, 0.20.5(1) cm diameter, 110-flowered in males and females; flowers
persistently tomentose with minute, pale brown hairs 0.10.3 mm long. Male flowers:
pedicel 815 mm long; bracteole caducous, attached to the pedicel at about or below
the middle; buds subglobose, (5)68 mm diameter, inside red; perianth at the sutures
12 mm thick, at anthesis clefts to c. 3/4; staminal disc subtriangular, flat, including anthers
35 mm diameter; anthers 1823; staminal column c. 2(3) mm long, clasped by the
fleshy basal part of the hypanthium. Female flowers: pedicel 34 mm long; buds obovoid,
57 mm long. Fruits 1 or 2 per infructescence, ellipsoid, 34 22.5 cm, mealy pubescent
with rather pale hairs c. 0.1 mm long, subglabrescent; dry pericarp c. 4 mm thick; stalk
c. 10 mm long, with the bracteole-scar located at about the middle.
Ecology. In mixed dipterocarp forest and periodically inundated forest along rivers, on
clay and sandstone-derived soils, up to 500 m altitude. Flowering throughout the year.
5. MYRISTICA Gronov., nom. cons.

(Greek, muron = a fragrant vegetable ointment; the nutmeg tree)
449
Fl. Orient. (1755) 141; Warburg, Monogr. Myrist. (1897) 374; Gamble, J. As. Soc. Beng. 75, 2 (1912)
226; Ridley, FMP 3 (1924) 62; J. Sinclair Gard. Bull. Sing. 26 (1958) 333, ibid. 28 (1968) 1; Backer &
Bakhuizen f. FJ 1 (1964) 138; Whitmore, TFM 1 (1972) 340; Anderson, CLTS (1980) 268; Corner,
WSTM 3rd. edition 2 (1988) 565; Whitmore, Tantra & Sutisna CLK 2, 1 (1990) 273; W.J. de Wilde,
Blumea 38 (1994) 349, ibid. 40 (1995) 237, ibid. 42 (1997) 111, ibid. 43 (1998) 165; PROSEA 5, 2
(1995) 346; Coode et al. (eds.), CBDS (1996) 226; Argent et al. (eds.), MNDT-CK 2 (1997) 459.
Dioecious, rarely monoecious or paroecious (having male and female organs in the same
inflorescence but separate, the male ones in the axils of lower bracts) trees; stilt roots
sometimes present. Bark rough, sometimes longitudinally fissured, greyish brown to coalblack; sap watery, red or pink. Twigs striate, older bark sometimes tending to crack
or flake. Leaves membranous, chartaceous or coriaceous, upper surface usually shining
when dry, lower surface often whitish or glaucous, glabrous or with tomentum of scalelike hairs, in Borneo without dark dots; base acute, rounded or less often cordate, apex
acute-acuminate or rarely obtuse; lateral veins nearly always impressed above, prominent
beneath and interarching near the leaf margin; intercostal venation usually laxly reticulate.
Inflorescences axillary branched panicles with the flowers arranged in cymes or subumbels,
or fascicles of flowers borne on unbranched or short-branched brachyblasts, or less often
a combination of the two types; bracts small and falling off early; bracteoles mostly
persistent, suborbicular and mostly embracing the base of the flowers. Flowers urceolate
or campanulate, less often tubular, pedicelled, often fragrant, glabrous, puberulous, pubescent
or tomentose outside, glabrous inside, rusty-brown or cream-coloured; perianth (2
)3(4)-lobed, lobes at anthesis generally reflexed; female flowers more swollen than
the males. Male flowers: androecium consists of an elongated synandrium which is made
up of a thickish central column to which 830 linear anthers are completely adnate by
their backsides and connate to each other by their edges, a rarely shallowly excavated
sterile apex, and a stalk (androphore). Female flowers: ovary subglobose, glabrous or
tomentose, style absent, stigma minutely bilobed. Infructescence either simple or furcate,
short, subsessile or panicle-like with peduncle, without scars of cataphylls at the base.
Fruits usually large, subglobose, ovoid, pyriform or oblong, orange, yellow or rusty brown,
glabrous, tomentose or less often lanose; pericarp thick, coriaceous, or fleshy but firm;
bracteoles persistent. Seeds shining, large and conform to the fruits; testa hard, brown
or brownish black; endosperm ruminate with a fixed oil and starch; aril red (or orange
or yellow), laciniate and clefts to the base or nearly to the base.
Distribution. A genus comprising about 175 species, distributed from S India and IndoChina (rare) through Malesia eastwards to NE Australia, the Pacifics and Fiji. Most species
are found in New Guinea, with many endemics. In Borneo 20 species occur, of which
19 are recorded in Sabah and Sarawak.
Ecology. All species are confined to undisturbed and disturbed, lowland to lower montane
rainforests. Many have a restricted distribution area, and/or are confined to a restricted
altitudinal range, sometimes to soils derived from specific bedrock, e.g. limestone or
ultrabasic.
Taxonomy. Sinclair in his monograph (l.c. 1968) distinguished arbitrarily two sections,
(1) sect. Myristica, with inflorescences generally more or less panicle-like and branched,
and with a smooth, non-scar covered (often more or less flattened) common peduncle,
450
generally lasting for one flowering season, and (2) sect. Fatua, with subsessile, simple
or forked, scar-covered, wart-like brachyblasts, lasting generally for several flowering
seasons. Within each section, 9 and 10 series respectively, are recognized. Though the
differences in the general shape and architecture of the inflorescences are useful in the
keys, a monophyletic division of the genus into two sections is unwarranted. In the present
treatment we have refrained from recognizing any sections or series.
Key to Myristica species

(Myristica fragrans, the nutmeg of commerce, widely cultivated and occasionally escaped
or remained in abandoned plantations, is not included in the key)
1. Tomentum of terminal leaf buds short, composed of minute appressed hairs 0.10.2
(0.3) mm long..........................2
Tomentum of terminal leaf buds consisting of hairs more than 0.2 mm long.............14
2. Lower leaf surface with persistent or subpersistent dense tomentum (hairs may be
very small), older leaves sometimes late-glabrescent. Fruits with minute scurfy tomentum
comprising hairs c. 0.1(0.2) mm long...................3
Lower leaf surface glabrous or early or late-glabrescent or with scattered minute hairs.
Fruits with various tomentum or glabrous or glabrescent..............5
3. Flower buds (best seen in staminate flowers) usually angular in the upper portion.
Twig slender, towards apex 1.52.5(4) mm diameter. Leaves (8)1022 cm long;
lateral veins 1017 pairs ...............3. M. cinnamomea
Flower buds not angular in the upper portion. Twig generally stouter. Leaves larger,
with more lateral veins per side..............................4
4. Twig stout, towards apex (4)58 mm diameter. Leaf upper surface drying blackish
brown; lateral veins (20)2330(33) pairs.............15. M. maxima (in part)
Twig slender, towards apex 23 mm diameter. Leaf upper surface drying olivaceous;
lateral veins 1623 pairs.......................6. M. depressa
5. Fruit glabrous or glabrescent or with minute remote hairs; in glabrescent fruit sometimes
some minute tomentum remaining in sheltered places near the insertion of the fruit
stalk. Inflorescences peduncled; flowers glabrous or pubescent...............................6
Fruit with persistent dense scurfy tomentum or partly glabrescent but with dense
tomentum remaining in sheltered places of the fruit. Inflorescences sessile brachyblasts
or peduncled; flowers pubescent............................11
6. Flower buds (especially in staminate flowers) sharply angular in the apical portion.
Twig drying conspicuously yellowish. Lateral veins 813(15) pairs.........7. M. elliptica
Flower buds not angular in the apical portion. Twig drying yellowish brown, brown,
or blackish. Lateral veins (10)1220(28) pairs.......7
7. Marginal interarching veins of lateral veins on the lower leaf surface indistinct..............8
Marginal interarching veins of lateral veins on the lower leaf surface generally raised
and distinct.....................10
451
8.
Bracteole (in staminate flowers) caducous. Leaf apex acute or rounded; lower leaf
surface distinctly papillose. Fruits 58 cm long, dry pericarp 1520 mm thick..
..............10. M. gigantea (in part)
Bracteole (in staminate flowers) persistent, subpersistent or late-caducous. Leaf apex
subacute-acuminate; lower leaf surface not papillose. Fruits 36(8) cm long, dry
pericarp 310 mm thick............................9
9.
Leaves membranous or chartaceous, or subcoriaceous, 724 cm long. Common

peduncle of female inflorescences and infructescences up to 10 mm long. Fruits 3.5
6(8) cm long, late or early glabrescent; dry pericarp 410 mm thick. Male flower
buds 46(7) mm long......12. M. iners (in part)
Leaves thinly membranous, 714 cm long. Common peduncle of female inflorescence
and infructescence slender, 1025(30) mm long. Fruits 3.54.5 cm long, early
glabrescent; dry pericarp 34 mm thick. Male flower buds c. 3 mm long......9. M. fallax
10. Twig slender, towards apex 24 mm diameter. Leaves smaller and narrower, 1030
49.5 cm; apex acute-acuminate, base rounded or acute; lateral veins 1420 pairs.
Male flower buds 33.5 mm long. Female flower buds 22.5 mm long. Fruits 4
6 cm long........14. M. malaccensis
Twig stout, towards apex 48 mm diameter. Leaves generally larger and broader,
1644 814.5(18) cm, apex blunt or acute, base rounded or subcordate; lateral
veins (16)2028 pairs. Male flower buds 68 mm long. Female flower buds c. 6
mm long. Fruits 79 cm long...........................16. M. papyracea
11. Inflorescences short, scar-covered, sessile brachyblast..............5. M. crassa
Inflorescences (and infructescences) with common peduncle, and branched.........12
12. Leaves 1130 3.510 cm; lateral veins 1320 pairs. Inflorescences 26 cm long
in males, 1.52.5 cm in females...........2. M. borneensis
Leaves (16)2435(40) (5)814(20) cm; lateral veins 1633 pairs. Inflorescences
large, widely or narrowly long-paniculate, 618 cm long in males, (1.5)59 cm long
in females..................................13
13. Twig stout, towards apex (4)58 mm diameter. Lateral veins (20)2330(33) pairs,
marginal interarching veins distinct. Male flower buds 57 mm long...........
...........15. M. maxima (in part)
Twig slender, towards apex 2.55 mm diameter. Lateral veins 1624 pairs, marginal
interarching veins indistinct. Male flower buds c. 4(4.5) mm long.............8. M. extensa
14. Twig stout, towards apex 510 mm diameter. Terminal leaf buds with very conspicuous
villous tomentum comprising hairs (1)23(4) mm long.......................19. M. villosa
Twig slender to rather stout, towards apex up to 6 mm diameter. Tomentum of terminal
leaf buds consisting of hairs up to 1.5 mm long......................15
15. Leaves pubescent beneath. Fruits 2.54.5 cm long.............16
Leaves glabrous or early glabrescent beneath. Fruits 5 cm long or more...........19
16. Inflorescences (best seen in male specimens) paniculate, i.e. branched and with distinct
more or less flattened main peduncles, with the flowers in subumbellate or spicate
452
clusters. Fruits 2.54.5 cm long..................................17

Inflorescences sessile or subsessile brachyblasts. Fruits 2.53.5 cm long...............18
17. Twig rather stout, towards apex 35(10) mm diameter. Leaves (12)1535 cm long;
lateral veins 1519(25) pairs. Flower buds not angular in the upper portion. Fruits
2.54.5 cm long, densely shaggy pubescent with hairs 0.51(2) mm long..............
........11. M. guatteriifolia
Twig slender, towards apex 1.53(3.5) mm diameter. Leaves 1118 cm long; lateral
veins (10)1214 pairs. Flower buds sharply angular in the upper portion. Fruits 3.5
4 cm long, glabrescent or thinly tomentose with scattered pale brown appressed hairs
up to 0.5 mm long.....................17. M. simiarum
18. Twig towards apex 23 mm diameter. Leaves thinly coriaceous, base rounded or shortacute; lateral veins not distinct above. Fruits tomentose with scurfy hairs 0.20.3(
0.5) mm long.........................18. M. smythiesii
Twig towards apex 34(5) mm diameter. Leaves coriaceous, base mostly rounded;
lateral veins distinct above. Fruits pubescent with woolly hairs 0.51 mm long........
......1. M. beccarii
19. Fruits glabrous (early glabrescent). Lower leaf surface not or indistinctly papillose,
concolourous with the upper surface................20
Fruits either with persistent tomentum, or late-glabrescent, or early glabrescent (in
M. gigantea). Lower leaf surface distinctly papillose, generally paler than the upper
surface......21
20. Leaves usually membranous, up to 24 cm long, base usually attenuate. Twig towards
apex up to 3 mm diameter, smooth or striate, bark lower down cracked or not. Terminal
leaf buds slender. Dry pericarp (4)510 mm thick.............12. M. iners (in part)
Leaves chartaceous or thinly coriaceous, up to 35 cm long, base broadly rounded
or subcordate. Twig towards apex (4)56 mm diameter, bark lower down conspicuously
coarsely flaky. Terminal leaf buds stout. Dry pericarp 1520 mm thick....
........4. M. corticata
21. Twig slender, towards apex 1.52.5(3) mm diameter. Leaves 712(15) cm long,
apex acute or rounded; lateral veins 1015(18) pairs. Fruits at first with scurfy
tomentum comprising hairs 0.20.3 mm long, late-glabrescent....
......10. M. gigantea (in part)
Twig generally stouter, towards apex 36 mm diameter. Leaves 1432(35) cm long,
apex acute-acuminate; lateral veins 1522 pairs. Fruits with conspicuous persistent
tomentum composed of shaggy hairs 12.5(3) mm long.........13. M. lowiana
1. Myristica beccarii Warb.

Whitmore, Tantra & Sutisna l.c. 273; W.J. de Wilde l.c. (1997) 140; Argent et al. (eds.) l.c. 460. Type:
Beccari 247, Borneo, Sarawak (holotype FI; isotypes G, K, M, P).
453
Tree up to 24 m tall, with low buttresses or a few stilt roots. Bark rough, fissured,
with narrow strips of 13 mm thick, or flaky, brownish black; inner bark orange-brownish;
sap yellowish or red. Sapwood white. Twigs towards apex 34(5) mm diameter, at first
tomentose with hairs c. 0.5(1) mm long; bark lower down blackish, somewhat flaky;
lenticels absent. Terminal leaf buds tomentose with hairs 0.51 mm long. Leaves coriaceous,
upper surface drying olivaceous or brown, lower surface with persistent dense tomentum
of pale cinnamon hairs (0.1)0.20.3 mm long, mixed with scattered, caducous, emergent
hairs c. 0.4 mm long, not papillose; elliptic-oblong to oblong-lanceolate, 920(25)
3.56.5(9) cm, base mostly rounded; midrib slightly raised above, late-glabrescent; lateral
veins (11)1419 pairs, sunken but distinct above, interarching veins indistinct on both
surfaces; intercostal venation indistinct on both surfaces; petiole 0.51(1.5) cm long.
Inflorescence a sessile or subsessile brachyblast, tomentose with hairs c. 0.5 mm long;
males simple or forked, woody, up to 1.2 cm long, few-flowered; bracts minute; females
small, 1few-flowered. Male flowers: pedicel 34 mm long; bracteole subpersistent, 2
4.5 mm long; buds ovoid-ellipsoid, 45.5 2.54.5 mm, woolly pubescent with hairs
0.30.8 mm long; perianth 3-lobed, at anthesis clefts to nearly 1/2; androecium 2.23
mm long; synandrium 23 mm long; anthers 68, sterile apex irregularly somewhat blunt,
0.20.3 mm long, pubescent with minute pale hairs up to 0.1 mm long; androphore 0.2
0.4 mm long, at base with a dense collar of hairs 0.50.8(1) mm long. Female flowers
pubescent with rufous hairs 0.51 mm long; buds ellipsoid, c. 5 3 mm long; ovary
ovoid (immature), 23 mm long. Fruits usually solitary, subsessile, ellipsoid-obovoid,
2.53.5 1.52 cm, rufous pubescent with woolly hairs 0.51 mm long, longest towards
the base; dry pericarp 12 mm thick; stalk rather stout, 24 mm long. Seeds ellipsoid,
c. 2(2.5) cm long.
Distribution. Sumatra and Borneo (Sabah, Sarawak, and Kalimantan, including Nunukan
Is.).
Ecology. In lowland and hill mixed dipterocarp forests on sandy or leached sandy clay
soils, and in ridge kerangas forest, up to 600 m altitudes. Flowering and fruiting throughout
the year.
2. Myristica borneensis Warb.
Fig. 15.
(of Borneo)
Monogr. Myrist. (1897) 401; Merrill l.c. (1921) 269; Masamune l.c. 304; Coode et al. (eds.) l.c. 226;
W.J. de Wilde l.c. 42 (1997) 139. Lectotype: Beccari 1574, Borneo, Sarawak, Mt. Matang (hololectotype
FI; isolectotypes G, K, P).
Tree up to 20 m tall; buttresses small or absent. Bark dark brown or blackish, smooth
or rough, shallowly flaky and minutely and narrowly fissured, or longitudinally striate,
not furrowed, thin, brittle; inner bark c. 25 mm thick, soft, red-brown; sap red, copious
or not, watery. Sapwood soft, white. Twigs towards apex 2.55 mm diameter, sometimes
yellowish, glabrescent, at first tomentose with pale hairs 0.1(0.3) mm long; bark lower
down frequently flaky; lenticels absent. Terminal leaf buds tomentose with greyish to
pale brown appressed hairs up to 0.1(0.3) mm long. Leaves membranous, chartaceous,
or thinly coriaceous, upper surface drying olivaceous, lower surface early glabrescent
454
vi
455
or with subpersistent sparse greyish hairs less than 0.1(0.3) mm long, drying grey or
pale cinnamon, usually distinctly papillose; elliptic-oblong or oblong-lanceolate, 1130
3.510 cm, base rounded or short-acute, apex acute-acuminate; midrib flat or slightly
raised above; lateral veins 1320 pairs, sunken above, interarching veins usually indistinct;
intercostal venation indistinct; petiole 1.22.8 cm long. Inflorescences pedunculate,
branched, mealy tomentose with greyish hairs 0.1(0.3) mm long; males 26 cm long,
peduncle (5)1020 mm long, flowers of different stages of development in subumbellate
clusters of 510; bracts minute; females 1.52.5 cm long, few-flowered, peduncle 510
mm long. Flowers drying yellow-brown or orange, thinly pubescent with grey-brown hairs
up to 0.1(0.3 mm) long. Male flowers: pedicel 36 mm long; bracteole rounded or
irregularly 3-lobed, 1.52(3) mm long; buds broadly ellipsoid, 34 2.53.5 mm, apex
broadly rounded, slightly 3-angular; perianth at anthesis clefts to 1/22/3; androecium
ellipsoid-cylindrical, 1.53 mm long; synandrium 1.52 mm long; anthers 710, sterile
apex broad, flat or irregularly truncate, 0.10.3 mm long, or absent; androphore about
as broad as the androecium, 0.20.8(1) mm long, pubescent with minute hairs up to
0.1 mm long. Female flowers: pedicel 23 mm long; buds 4(4.5) 3.54 mm; perianth
at anthesis clefts to c. 1/2; ovary ovoid, densely pubescent with minute hairs. Fruits
1 or 2 per infructescence, more or less ellipsoid, (4.5)57.5 34.5 cm; dry pericarp
5(8) mm thick, cinnamon puberulent with hairs 0.1(0.3) mm long, persistent or lateglabrescent; stalk smooth, tapering, 59 mm long, with the scar of the bracteole located
more or less immediately below the fruit. Seeds more or less ellipsoid, (3.5)44.5 cm
long.
Ecology. In primary lowland and hill mixed dipterocarp forests, and heath forest, mostly
on sandy clay and clay-rich yellow soils, up to 1000 m altitude. Flowering in April
June; fruiting mainly in AugustNovember.
3. Myristica cinnamomea King

(Latin, cinnamomeus = yellow-brown; the tomentum of lower leaf surface)
Ann. Roy. Bot. Gard. Calc. 3 (1891) 292; Warburg l.c. (1897) 445; Gamble l.c. 232; Ridley l.c. 65; J.
Sinclair l.c. (1958) 358, l.c. (1968) 210; Whitmore l.c. 342; Anderson l.c. 268; Corner l.c. 565; Whitmore,
Tantra & Sutisna l.c. 273; Turner, Gard. Bull. Sing. 47 (1995) 362; Coode et al. (eds.) l.c. 227; W.J. de
Wilde l.c. 42 (1997) 138; Argent et al. (eds.) l.c. 460. Lectotype: Kings Collector 7474, Peninsular
Malaysia, Perak (hololectotype CAL; isolectotypes BM, FI, K, L, P).
Tree up to 30 m tall; crown dense; trunk at base often somewhat fluted or with low
buttresses, or often with few small stilt roots. Bark blackish brown, or chocolate, or
dark grey, britle, finely fissured, or sometimes with fine horizontal cracks or thinly flaking
with appressed flakes; inner bark c. 5 mm thick, usually reddish brown to pinkish, soft,
more or less laminated; sap red, watery, slow to appear. Sapwood soft, pale yellowish
to pale brownish, often reddish speckled. Twigs slender, towards apex 1.52.5(4) mm
diameter, at first tomentose with hairs c. 0.1 mm long, glabrescent; bark lower down
not or rarely flaky; lenticels absent. Terminal leaf buds tomentose with minute appressed
hairs c. 0.1 mm long. Leaves chartaceous to thinly coriaceous, upper surface drying
456
olivaceous, more or less glossy, lower surface densely grey-brown or silvery-brown

tomentose with inconspicuous hairs 0.1(0.2) mm long, sometimes late-glabrescent,
indistinctly papillose, without brownish or blackish dots; elliptic to lanceolate, (8)10
22 (2)2.58 cm, base attenuate, apex acute or acute-acuminate; midrib flat or somewhat
raised but lying in a groove above; lateral veins 1017 pairs, flat or sunken above,
interarching veins indistinct; intercostal venation indistinct; petiole 1.22.2 cm long.
Inflorescences short or distinctly pedunculate, branched, pubescent with mealy hairs 0.1
(0.2) mm long; males 12.5 12 cm, peduncle 512(15) mm long, flower buds of
different stages of development and sizes in subumbellate clusters of 58(10); bracts
minute; females with peduncle up to 3 mm long (rarely somewhat forked), flowers in
subumbellate clusters of 24; flower buds more or less angular in the upper portion,
pubescent with hairs 0.1(0.3) mm long; bracteole subpersistent or caducous. Male flowers:
pedicel 36 mm long; buds ellipsoid-oblong, 57 (2)2.53 mm, apex subacute, angular;
perianth 3(4)-lobed, at anthesis clefts to 1/31/2; androecium 3.54(4.5) mm long;
androphore 1.52(2.5) mm long, pubescent with pale hairs up to 0.1 mm long; synandrium
ellipsoid-oblong, 1.82 mm long; anthers 68, contiguous, sterile apex somewhat blunt,
0.20.3 mm long, minutely pubescent. Female flowers: pedicel 2.54 mm long; buds
ovoid-ellipsoid, apex subacute, angular, 56 2.53 mm; ovary ovoid-conical, c. 2.5
1.52 mm, tomentose with dense hairs up to 0.1 mm long. Infructescences 0.52 cm
long. Fruits solitary or 2(3) per infructescence, broadly ellipsoid or oblong, 59 2.5
5 cm, base rounded or subattenuate, densely scurfy pubescent with rusty hairs 0.1(0.2)
mm long; dry pericarp (5)815 mm thick; stalk rather stout, 510 mm long. Seeds ellipsoid,
3.55.5 cm long.
Distribution. Sumatra, Peninsular Malaysia, Singapore, Borneo, and possibly in the
Philippines (Mindanao). In Sabah and Sarawak, locally common or scattered in various
types of primary forests. Also in Brunei and Kalimantan.
Ecology. In primary lowland and hill mixed dipterocarp, peatswamp, kerangas and lower
montane forests, on a variety of soils, up to 1400 m altitude. Flowering and fruiting
throughout the year, but flowering predominantly in MarchJune.
4. Myristica corticata W.J. de Wilde

(Latin, corticatus = covered with bark; the coarsely flaked bark of twigs)
Blumea 42 (1997) 157. Type: Krispinus SAN 87293, Borneo, Sabah (holotype L; isotypes K, KEP
SAR, SING).
Tree up to 25 m tall; buttresses up to 1 m tall and wide; bole straight. Bark brown
or blackish, cracked, hard and brittle, c. 2.5 mm thick, peeling off in small longitudinal
thin flakes; inner bark c. 0.6 mm thick, hard, orange-brown, reddish, or yellowish. Sapwood
pink, red, or light brown. Twigs rather stout, towards apex angular, (4)56 mm diameter,
pubescent with hairs 0.51 mm long, early glabrescent; bark lower down coarsely flaky;
lenticels absent. Terminal leaf buds stout, tomentose with rough hairs c. 1 mm long.
Leaves chartaceous or thinly coriaceous, upper and lower surfaces drying olivaceousbrown, lower surface glabrous, indistinctly papillose; elliptic-oblong or oblong-lanceolate,
1535 611.5 cm, base broadly rounded or subcordate, apex acute-acuminate; midrib
457
slightly raised above; lateral veins 1724 pairs, generally sunken above, interarching veins
mostly distinct; intercostal venation generally indistinct on both surfaces; petiole 1.5
2.5 cm long. Inflorescences pedunculate, branched, woolly pubescent with hairs 1(1.5)
mm long; males 24.5 23 cm, peduncle flattened, 46 mm long, flowers of various
stages of development and sizes in clusters of 510, woolly pubescent with hairs 0.3
(0.5) mm long. Male flowers: pedicel 46 mm long; bracteole persistent, broadly ovate,
c. 3.5 mm long; buds ovoid, 55.5 44.5 mm, apical portion not angular; perianth
3-lobed, at anthesis clefts to c. 1/2; androecium slender, c. 4.5 0.8 mm; synandrium
c. 3 mm long; anthers 78, sterile apex somewhat blunt, c. 0.2 mm long, glabrous;
androphore c. 1.5 mm long, pubescent with hairs 0.20.3 mm long in the lower half.
Female flowers not seen. Infructescences little-branched, 1.52 cm long, glabrescent.
Fruits 1(2) per infructescence, elliptic-oblong, 7.59.5 45 cm, at first pubescent
with dense scurfy hairs less than 0.1 mm long, early glabrescent; dry pericarp 15(20)
mm thick; stalk (2)5 mm long. Seeds ellipsoid-oblong, 4.55.5 cm long.
Distribution. Endemic to Borneo (Sabah and Brunei).
Ecology. In primary and logged-over hill mixed dipterocarp forests, on low undulating
country, hilly sides, and sandstone ridges with sandy clay soils, up to 500 m altitude.
Flowering in July; fruiting in May, August and November.
5. Myristica crassa King

(Latin, crassus = thick; the thick pericarp)
Sinclair l.c. (1958) 366, l.c. (1968) 435; Anderson l.c. 268; Turner l.c. 362; W.J. de Wilde l.c. (1997)
159. Lectotype: Kings Collector 5065, Peninsular Malaysia (hololectotype CAL; isolectotypes FI, G,
K, L). Synonym: M. suavis King l.c. 295, Warburg l.c. (1897) 441.
Tree up to 25 m tall. Bark brown, smooth to slightly and shallowly fissured longitudinally
and horizontally; inner bark red. Sapwood white. Twigs towards apex 37 mm diameter,
at first tomentose with minute hairs, early glabrescent; bark lower down not flaky; lenticels
sometimes present, not distinct. Terminal leaf buds tomentose with minute appressed hairs
up to 0.1 mm long. Leaves chartaceous or thinly coriaceous, upper surface drying dark
olivaceous, lower surface greyish brown, glabrous or with scattered scale-like hairs up
to 0.1 mm long, distinctly papillose; elliptic-oblong to oblong-lanceolate, (13)1740
4.512(13) cm, base rounded to cuneate, margin often somewhat revolute when dry,
apex acute-acuminate; midrib slightly raised above; lateral veins 1318(22) pairs, sunken
above, interarching veins not distinct; intercostal venation indistinct; petiole 23 cm long.
Inflorescences: brachyblasts, 12 cm long, pubescent with hairs up to 0.1 mm long,
sessile or with peduncle up to 6 mm long; males 1030(many)-flowered with the flowers
in different stages of development and sizes; bracts minute; females 25 flowered. Flowers
tomentose with hairs less than 0.1 mm long, sometimes late-glabrescent. Male flowers:
pedicel (5)612(15) mm long; bracteole subpersistent, sometimes somewhat angular
at the apex, (2)34 mm long; buds ovoid, urceolate to oblong, 4.56 2.55 mm; perianth
3-lobed, at anthesis clefts to c. 1/3; androecium slender, 33.5 0.5 mm; synandrium
ellipsoid-oblong, c. 2 mm long; anthers (4)58, sterile apex narrow, somewhat blunt,
458
c. 0.3 mm long; androphore (0.7)12 mm long, minutely pubescent in the lower 2/3
or towards the base only. Female flowers: pedicel rather stout, c. 2 mm long; bracteole
caducous; buds subovoid, 45 34 mm; ovary 3(4) 22.5 mm, minutely pubescent.
Infructescences short. Fruits 1 or 2 per infructescence, broadly ellipsoid, (3.5)45.5
(6) 34 cm, rusty pubescent with scurfy hairs c. 0.1 mm long (the scurf rather easily
rubbed off); dry pericarp (7)10 mm thick; stalk stout, 5(10) mm long. Seeds ellipsoid,
33.5 cm long.
Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo (uncommon in Sarawak,
Brunei and Kalimantan).
Ecology. In freshwater swamp, peatswamp and kerangas forests, up to 600 m altitude.
Notes. Trees are occasionally paroecious, with male and female flowers in the same
inflorescence.
6. Myristica depressa W.J. de Wilde

(Latin, depressus = flattened, sunken; the androecium with cavity at the apex)
Blumea 42 (1997) 161. Type: S. 39108, Borneo, Sarawak (holotype L; isotypes K, SAR).
Tree up to 40 m tall, sometimes with buttresses or stilt roots. Bark brittle, fissured or
flaky, blackish; inner bark brownish. Sapwood white or brownish. Twigs slender, towards
apex 23 mm diameter, at first tomentose with hairs up to 0.1 mm long, early glabrescent,
sometimes yellowish; bark lower down coarsely striate, sometimes flaky; lenticels absent.
Terminal leaf buds pubescent with minute appressed hairs 0.1(0.2) mm long. Leaves
membranous or thinly chartaceous, upper surface drying olivaceous, lower surface greybrown, densely pubescent with persistent arachnoid hairs up to 0.1 mm long, not papillose;
oblong-lanceolate, 1530 49 cm, base short cuneate or rounded, apex acute-acuminate;
midrib flat to moderately raised above; lateral veins 1623 pairs, flat or sunken above,
sometimes indistinct, interarching veins distinct or not; intercostal venation usually faint
on both surfaces; petiole 12.5 cm long. Inflorescences pedunculate, branched, rather
slender, pubescent with woolly hairs 0.10.3 mm long; males 23.5 cm long, peduncle
510 mm long, flower buds of different stages of development and sizes in subumbellate
clusters of 48; bracts 1.52 mm long; females as the males but smaller. Male flowers
tomentose with woolly hairs (0.1)0.2(0.3) mm long; pedicel 33.5 mm long; bracteole
apically attached to the pedicel, often faintly 2-lobed, c. 2 mm long; buds not angular
in the upper portion, somewhat carnose, ovoid or ellipsoid, 34 2.53 mm; perianth
3-lobed, at anthesis clefts to 3/44/5; androecium short and broad, sometimes club-shaped,
c. 1.5 1.2 mm; synandrium broadly ellipsoid, 11.3 11.2 mm, sterile apex absent,
the apex truncate and with cavity up to 0.3 mm deep, more or less lobed corresponding
with the anthers; anthers 68(10); androphore broadly cylindrical, 0.30.5 mm long,
minutely pubescent. Mature female flowers not seen. Infructescences 0.52 cm long,
with 1 or 2 fruits. Fruits ellipsoid-oblong, 57 34.5 cm; dry pericarp (2)36(10)
mm thick, tomentose with dense scurfy hairs 0.1(0.2) mm long; stalk rather stout, roughly
fissured or cracked, 35 mm long. Seeds ellipsoid, 45 cm long; aril pink.
459
Distribution. Sumatra (1 collection), Peninsular Malaysia and Borneo (Sabah and Sarawak).
Ecology. In primary mixed dipterocarp forests, on various types of soils; occasionally
on inundated river banks, sandstone and diorite screes, and hill sides, up to 300 m altitude.
7. Myristica elliptica Wall. ex Hook. f. & Thomson

(Latin, ellipticus = shaped like an ellipse; the leaves)
Fl. Ind. 1 (1855) 162; King l.c. 295; Warburg l.c. (1897) 435; Gamble l.c. 231; Merrill l.c. (1921) 269;
Ridley l.c. 65; Masamune l.c. 304; J. Sinclair l.c. (1958) 353, l.c. (1968) 188; Whitmore l.c. 342; Anderson
l.c. 268; Corner l.c. 565; Whitmore, Tantra & Sutisna l.c. 273; Turner l.c. 362; W.J. de Wilde l.c. (1997)
164; Argent et al. (eds.) l.c. 460. Type: Porter in Wallich Cat. 6798a, Penang (holotype K; isotypes BM,
E, G, M, NY). Synonym: M. elliptica Hook. f. & Thomson var. elliptica J. Sinclair l.c. (1958) 353, l.c.
(1968) 188.
Tree up to 40 m tall; bole sometimes fluted or buttressed up to 1.2 m high. Bark rather
smooth or finely fissured or scaly, sometimes lenticellate, grey to dark brown or blackish;
inner bark soft, red-brown to pink; sap watery, pink or red. Sapwood white to pale yellow.
Twigs towards apex 24 mm diameter, drying yellowish, at first tomentose with greyish
hairs 0.10.3 mm long, glabrescent; bark lower down not cracked, straw-coloured,
sometimes with few lenticels. Terminal leaf buds pubescent with glossy light brown, minute,
appressed hairs 0.10.2(0.3) mm long. Leaves chartaceous, upper surface drying olivaceous,
lower surface greyish olivaceous, at first tomentose with appressed pale hairs up to 0.1
mm long, glabrescent, finely pustulate but indistinctly papillose, sometimes with faint
brownish dots; elliptic to oblong, 921(32) 39(12) cm, base rounded to acute, apex
acute-acuminate; midrib slightly raised above; lateral veins 813(15) pairs, faint, flat
or slightly raised above, interarching veins sometimes distinct; intercostal venation faint
on both surfaces; petiole 12(3) cm long. Inflorescences pedunculate, branched, sparsely
pubescent with pale appressed hairs up to 0.1 mm long, almost glabrescent; males 1
2.5 11.5 cm, rather few-flowered, peduncle slightly flattened, 510(12) mm long;
flower buds of different stages of development in loose subumbellate clusters of 35;
bracts minute; females much reduced, 13-flowered. Flowers angular in the upper portion,
thinly pubescent with appressed pale hairs c. 0.1 mm long; bracteole persistent, often
more or less truncate, 0.51(2) mm long. Male flowers: pedicel 35 mm long; buds
elliptic-oblong, sometimes slightly contracted below the perianth-lobes, 78(9) 2.5
3.5 mm, upper portion subacute, angular; perianth 3-lobed, at anthesis clefts to c. 1/
3; androecium cylindrical, 57 11.5 mm; androphore 2(2.5) mm long, glabrous or
at the base minutely pubescent; synandrium 44.5 mm long; anthers 5 or 6, sterile apex
somewhat blunt, c. 0.5 mm long. Female flowers: pedicel c. 3 mm long; buds ovoidoblong, upper portion acute, angular, 68 34 mm; ovary conical, including style c.
5 1 mm, tomentose with dense appressed hairs c. 0.1 mm long. Infructescences often
conspicuously lenticellate, with 1 or 2 fruits. Fruits ovoid or ellipsoid-oblong, 4.57.5
(2)2.54 cm, apex sometimes beaked, base often narrowed, at first tomentose with
scattered hairs c. 0.1 mm long, early glabrescent; dry pericarp dark brown, 5(10) mm
thick; stalk slender, 510 mm long. Seeds ellipsoid, 45 cm long.
Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, and Borneo (Sabah,
460
Ecology. In primary freshwater swamp and peatswamp forests but also in lowland and
hill forests on various soils, up to 500 m altitude. Flowering and fruiting throughout
the year.
8. Myristica extensa W.J. de Wilde

(Latin, extensus = stretched out; the extended inflorescences)
Blumea 42 (1997) 164. Type: Ilias S. 25678, Borneo, Sarawak (holotype L; isotype SAR).
Tree up to 25 m tall; bole fluted, with or without buttresses, or with stilt roots extending
up to 80 cm over the ground. Bark smooth or scaly, hard, c. 1 mm thick, cracked, at
base of bole peeling off in strips, brittle, grey to blackish or dark brown; inner bark
c. 5(10) mm thick, brown; sap blood-red or pink. Sapwood brown. Twigs slender, towards
apex subterete, 2.55 mm diameter, at first tomentose with pale hairs up to 0.1 mm long,
early glabrescent; bark lower down cracked; lenticels few or absent. Terminal leaf buds
tomentose with minute appressed hairs c. 0.1 mm long. Leaves membranous or thinly
chartaceous, upper surface drying olivaceous-brown, lower surface grey-brown, at first
tomentose with flat, open cobweb-like silvery indumentum, late-glabrescent, indistinctly
papillose; elliptic-oblong to oblong-lanceolate, (16)2435 (5)813.5 cm, base attenuate
or narrowly rounded, apex acute-acuminate; midrib flat above; lateral veins 1624 pairs,
indistinct and flat or sunken above, interarching veins not distinct; intercostal venation
usually indistinct on both surfaces; petiole 1.53 cm long. Inflorescences pedunculate,
branched, almost glabrous; males widely paniculate, 1015 610 cm, peduncle 20
45 mm long, flower buds of different stages of development in subumbellate clusters
of 510; bracts not known, bract scars usually considerably dislocated upwards; females
(1.5)58 cm long (including 3040 mm long peduncle). Male flowers: pedicel slender,
912 mm long; bracteole persistent, 22.5 mm long; buds ovoid, 4(4.5) 33.5 mm,
apex not angular; perianth membranous, thinly pubescent with hairs 0.1(0.2) mm long,
3-lobed, at anthesis clefts to nearly 2/3; androecium short, club-shaped, 33.5 11.2
mm long; synandrium ellipsoid-oblong, c. 2 mm long; anthers somewhat spaced, c. 8,
sterile apex short, 0.20.3 mm long, somewhat blunt; androphore c. 1 1 mm, glabrous.
Fruits 1 or 2 per infructescence, ellipsoid-oblong, 67 33.5 cm; pericarp dull brown,
drying 36 mm thick, densely pubescent with scurfy hairs up to 0.1 mm long; stalk 6
15 mm long. Seeds 55.5 cm long; aril orange-red to red.
Distribution. Endemic to Borneo (Sarawak, Brunei, C and E Kalimantan).
Ecology. In primary lowland mixed dipterocarp forest on undulating lands, spurs, steep
slopes, and hilly crests, on clay-rich, sandy yellow loam, yellow clayey, and sandy clay
soils, up to 800 m altitude. Flowering in June; fruiting mainly in March and August.
9. Myristica fallax Warb.

(Latin, fallax = deceptive; closely resembling another species)
Monogr. Myrist. (1897) 410; Merrill l.c. (1921) 269; Masamune l.c. 304; W.J. de Wilde l.c. (1997) 166.
Type: Beccari 1590, Borneo, Sarawak, Mt. Matang (FI).
461
Tree up to 20 m tall; buttresses up to 1 m out. Bark smooth or slightly flaky, chocolate,

with paler patches or whitish flakes; sap reddish, watery. Twigs slender, towards apex
11.5 mm diameter, drying yellowish brown, at first sparsely pubescent with scattered
hairs less than 0.1 mm long, early glabrescent; bark lower down brown, inconspicuously
longitudinally cracked; lenticels inconspicuous, or absent. Terminal leaf buds tomentose
with minute appressed hairs up to 0.1 mm long. Leaves thinly membranous, upper surface
drying olivaceous, lower surface paler, greyish, at first tomentose with minute, scattered,
pale grey-brown stellate hairs less than 0.1 mm long, early glabrescent, not papillose;
oblong-lanceolate or lanceolate, 714 24 cm, base cuneate, apex acute-acuminate;
midrib flattish or slightly raised above; lateral veins 1216 pairs, flat or sunken above,
faintly conspicuous below, interarching veins indistinct; intercostal venation indistinct;
petiole slender, 12 cm long. Male inflorescences c. 4 cm long, pedunculate, branched;
peduncle c. 10 mm long; bracts not known. Male flowers: pedicel c. 8 mm long; bracteole
c. 0.5 mm long, persistent; buds glabrous, ovoid, c. 3 2 mm, not angular in the upper
portion; androecium (including androphore) c. 2.3 mm long; androphore c. 1 mm long,
subpuberulous, sterile apex minute, obtuse; anthers 5. Female flowers not known.
Infructescences located at the apical portion of the twigs, axillary to leaves, c. 3 cm
long; peduncle slender, 1025 mm long, with a few scars of fallen flowers, glabrous
(early glabrescent). Fruits solitary, ovoid-ellipsoid, 3.54(4.5) 22.7 cm, glabrous (i.e.
when young tomentose with pale rusty scurfy scale-like hairs less than 0.1 mm long,
early glabrescent); dry pericarp 34 mm thick; stalk slender, 510 mm long. Seeds ellipsoid,
22.5 cm long.
Distribution. Endemic to Borneo (Sarawak and E Kalimantan).
Ecology. In hill mixed dipterocarp forest on yellow sandy loam soils, at 100600m altitude.
Fruiting in August and OctoberNovember.
10. Myristica gigantea King

(Latin, giganteus = very large; probably referring to the size of the trees)
Sinclair l.c. (1958) 343, l.c. (1968) 163; Whitmore l.c. 342; Whitmore, Tantra & Sutisna l.c. 273; Turner
l.c. 362; W.J. de Wilde l.c. (1997) 167; Argent et al. (eds.) l.c. 460. Lectotype: Kings Collector 6050,
Peninsular Malaysia, Perak (hololectotype CAL; isolectotypes E, FI, K, KEP, P, SING).
Tree up to 40 m tall; with low buttresses or a few stilt-roots up to 2 m high. Bark

generally blackish (charcoal colour), longitudinally fissured, or often flaking in strips or
pieces c. 1 cm wide, brittle; inner bark 1012 mm thick, brown or red; sap red, sometimes
recorded as sticky. Sapwood white-yellow, heartwood brown. Twigs drying brownish,
slender, towards apex 1.52.5(3) mm diameter, at first tomentose with short conspicuous
hairs 0.11 mm long mixed with emergent hairs 11.5 mm long, late-glabrescent; bark
lower down usually coarsely flaky; lenticels inconspicuous or absent. Terminal leaf buds
pubescent with hairs 0.11.5 mm long. Leaves chartaceous or thinly coriaceous, upper
surface drying olivaceous, lower surface greyish or pale brown, at first tomentose with
scattered hairs up to 0.1 mm long, early glabrescent, distinctly papillose; elliptic-oblong
to lanceolate, 715 24 cm, that of sapling shoots up to 22 8 cm, base rounded
462
or attenuate, apex rounded or acute, in sapling leaves acute-acuminate; midrib slightly

raised above; lateral veins 1015(18) pairs, sunken above, interarching veins generally
indistinct; intercostal venation indistinct; petiole 1.23.5 cm long. Inflorescences pedunculate,
branched, shaggy pubescent with hairs (0.2)0.51 mm long, sometimes glabrescent; males
2.53.5 2.53 cm, peduncle 610 mm long, flower buds of different stages of development
and sizes in subumbellate clusters of 612; bracts c. 4 mm long, caducous; females c.
1.5 cm long, few-flowered, flowers pubescent with hairs (0.2)0.5 mm long. Male flowers:
pedicel c. 4 0.5 mm long; bracteole c. 2 mm long, caducous; buds ovoid-ellipsoid,
44.5 3 mm, not angular in the upper portion; perianth 3(4)-lobed, at anthesis clefts
to 1/51/4; androecium slender, c. 3.5 0.81 mm long; androphore c. 1.5 mm long,
pubescent with hairs c. 0.3 mm long in the lower 2/3; synandrium cylindrical, acutish,
c. 2 0.8 mm, sterile apex absent; anthers 78. Infructescences 11.5 cm long, each
with 1 or 2 fruits. Fruits broadly ovoid, 58 4.55 cm, at first scurfy tomentose with
hairs 0.20.3 mm long, early or late-glabrescent (tomentum usually persistent near insertion
of the fruit stalk); dry pericarp 1520 mm thick; stalk stoutish, 59 mm long. Seeds
ellipsoid, 33.5 cm long.
Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo (Sabah, Sarawak and
Kalimantan).
Ecology. In lowland mixed dipterocarp forest on sandy and sandy loam soils, up to 450
m altitude. Flowering and fruiting possibly throughout the year.
11. Myristica guatteriifolia A. DC.

(Latin, guatteriifolius = with leaves like those of Guatteria, an American genus of the
Annonaceae)
Ann. Sc. Nat. 4, 4 (1855) 20 (guatteriaefolia); Warburg l.c. (1897) 412; Merrill l.c. (1921) 269; Masamune
l.c. 304; J. Sinclair l.c. (1958) 350, l.c. (1968) 213; Backer & Bakhuizen f. l.c. 139; Anderson l.c. 268;
Corner l.c. 566; Whitmore, Tantra & Sutisna l.c. 273; Turner l.c. 362; Coode et al. (eds.) l.c. 227; W.J.
de Wilde l.c. (1997) 168; Argent et al. (eds.) l.c. 460. Type: Cuming 1582, the Philippines, Luzon,
Batangas Prov. (G). Synonyms: M. littoralis Miq., Fl. Ind. Bat. 1, 1 (1858) 57; M. cookii Warb. l.c.
(1897) 414; M. riedelii Warb. l.c. (1897) 417; M. palawanensis Merr., Phil. J. Sc. C. Bot. 13, 5 (1918)
283.
Tree up to 25 m tall; buttresses up to 2 m high, 30(150) cm out. Bark chocolate-brown

or brown-black, 24 mm thick, smooth or rather rough, only rarely fissured, usually cracked
or scaly, flaking irregularly in more or less rectangular pieces c. 2 4 cm; inner bark
510 mm thick, soft, fibrous, pinkish to beef-red; sap watery, pale red to red. Sapwood
usually pale, whitish to yellow, streaked with red; heartwood reddish. Twigs rather stout,
subterete, towards apex 36 mm diameter, pubescent with hairs 0.20.5 mm long,
glabrescent, usually without lenticels; bark of older twigs not or indistinctly flaky. Terminal
leaf buds pubescent with hairs (0.1)0.20.5(1) mm long. Leaves chartaceous, upper
surface drying olivaceous, lower surface pubescent or late-glabrescent with dense, greybrown or cinnamon hairs 0.20.3 mm long, mixed with scattered emergent hairs c. 0.5
(2) mm long; elliptic-oblong to oblong-lanceolate, (12)1535 312 cm, base rounded
or attenuate, apex acute-acuminate; lateral veins 1519(25) pairs, slightly sunken above,
interarching veins distinct or faint; intercostal venation distinct or faint; petiole 1.53
463
cm long. Inflorescences paniculate, pedunculate, densely tomentose with hairs 0.20.6

mm long; bracts caducous, c. 2 mm long; males 28 cm long, with flowers of different
stages of development in subumbellate clusters of (1)210, peduncle somewhat flattened,
(0)420 mm long; females few-flowered, 0.52 cm long. Flowers pubescent with hairs
0.30.6 mm long, apex not angular; bracteole (2)2.53 mm long, caducous. Male flowers:
pedicel 36 mm long; buds ovoid to ellipsoid, (4)57 3.55 mm; perianth at anthesis
clefts to 1/41/3; androecium (3)3.55 0.81.2(1.8) mm; synandrium cylindrical, 2
3 mm long; anthers 812, sterile apex up to 0.2 mm long, or with shallow cavity,
subglabrous; androphore 1.52 mm long, towards the base with pale hairs 0.40.7 mm
long. Female flowers: pedicel 12(4) mm long; buds broadly ovoid, 46 3.56 mm;
ovary broadly ovoid, tapered, 2.53(3.5) mm diameter, densely pubescent with hairs
c. 0.5 mm long, stigma obliquely 2-lobed, each lobe faintly lobulate or not. Fruits solitary
or up to 3 per infructescence, short-ellipsoid or subglobose, 2.54(4.5) (2)2.53 cm;
dry pericarp 35(8) mm thick, pubescent with dense shaggy hairs 0.51(2) mm long;
stalk 1.55 mm long. Seeds 22.5(3) cm long.
Vernacular name. Sabahalanagni (Dusun, preferred name).
Distribution. Burma, possibly Thailand (no specimens seen), Indo-China (P. Condor),
Sumatra, Peninsular Malaysia, Borneo (throughout), Java, Bali, and the Philippines.
Ecology. Frequently found in coastal forest/vegetation but also in forests further inland
on various types of soils, including sandstone- and limestone-derived and ultrabasic soils,
up to 200 m altitude. Flowering and fruiting throughout the year.
12. Myristica iners Blume

(Latin, iners = inert or sluggish; the aroma of the seed)
Bijdr. Fl. Ned. Ind. 2, 11 (1826) 575; Gamble l.c. 230; Ridley l.c. 64; J. Sinclair l.c. (1958) 363, l.c.
(1968) 177; Backer & Bakhuizen f. l.c. 139; Whitmore l.c. 343; Anderson l.c. 268; Whitmore, Tantra &
Sutisna l.c. 273; Turner l.c. 362; Coode et al. (eds.) l.c. 227; W.J. de Wilde l.c. (1997) 168; Argent et al.
(eds.) l.c. 461. Type: Blume s.n., W Java (L). Synonyms: M. sublanceolata Miq. l.c. 58; M. vordermannii
Warb. l.c. (1897) 525; M. cumingii Warb. var. floribunda Airy Shaw, Kew Bull. 1939, no.10 (1940) 539.
Tree up to 40 m tall, sometimes with stilt roots. Bark rough, deeply fissured, 35 mm
thick; inner bark c. 10 mm thick, soft, laminated, pinkish or brown; sap clear, reddish,
sometimes copious. Sapwood white or pale coloured, sometimes red-streaked. Twigs
slender, towards apex more or less angular, 1.53 mm diameter, drying brown, at first
tomentose with greyish hairs up to 0.5 mm long, early glabrescent; bark lower down
usually flaky, lenticels sparse. Terminal leaf buds slender, pubescent with hairs c. 0.2
(0.5) mm long. Leaves membranous, chartaceous, or subcoriaceous, upper surface drying
olivaceous-brown, lower surface concolourous with the upper surface, at first tomentose
with scale-like hairs less than 0.1 mm long, early glabrescent, not papillose; ellipticoblong to oblong-lanceolate, 724 1.58(10) cm, base rounded or attenuate, apex
subacute-acuminate; midrib slightly raised above; lateral veins 1116 pairs, flat to sunken
above, interarching veins usually indistinct; intercostal venation indistinct; petiole 12.5
cm long. Inflorescences branched, glabrescent; males 28 cm long, peduncle slender or
464
stout, 230(40) mm long, flower buds of different stages of development and sizes in
subumbellate clusters of 515, not angular in the upper portion; females reduced, with
peduncle up to 10 mm long or sometimes sessile, 16-flowered; bracts caducous, 34
mm long. Flowers thinly pubescent with woolly hairs 0.10.3 mm long, usually early
glabrescent; bracteole 1.53 mm long, subpersistent or late caducous. Male flowers:
pedicel slender, 412 mm long; buds ovoid or ovoid-ellipsoid, 46(7) 24(6) mm;
perianth at anthesis clefts to (1/4)1/31/2; androecium 3(6) 0.61 mm long; synandrium
23 mm long; anthers 68, sterile apex 0.10.3 mm long, entire or shallowly lobed, glabrous;
androphore cylindrical or somewhat swollen, about as broad as the synandrium, 1.22(
3) mm long, pubescent with hairs 0.10.2 mm in the lower half. Female flowers: pedicel
17 mm long; buds ovoid, 44.5 3.54 mm; ovary long-ovoid, c. 3 (1)1.5 mm,
tomentose with dense hairs up to 0.1 mm long. Fruits usually 1 per infructescence, scurfy
pubescent with grey-brown hairs c. 0.2 mm long, usually early glabrescent, ellipsoid or
ovoid-ellipsoid, 3.56(8) 2.53.5(4.5) cm; dry pericarp (4)510 mm thick, stalk
rather slender, 515 mm long. Seeds ellipsoid, 24(5) 12(2.3) cm, brown or black;
aril orange to red.
Vernacular name. Sabahdarah-darah (preferred name).
Distribution. Cambodia, Thailand, Sumatra, Peninsular Malaysia, Java, Borneo (throughout),
and possibly in the Philippines.
Ecology. In lowland to lower montane forests on flat lands, slopes and ridges, on sandy,
sandstone-derived, sandy loam, and tuff soils, up to 1800 m altitude. Flowering and fruiting
13. Myristica lowiana King

(Sir H. Low, 18241905, officer of the British East India Company; plant collector)
Ann. Roy. Bot. Gard. Calc. 3 (1891) 293, p.p. (pl. 120, fig. 1, 5, 6, 7 = M. crassa); Warburg l.c. (1897)
496; Gamble l.c. 235; Ridley l.c. 66; J. Sinclair l.c. (1958) 345, l.c. (1968) 164; Whitmore l.c. 344;
Anderson l.c. 269; Whitmore, Tantra & Sutisna l.c. 273; Turner l.c. 362; Coode et al. (eds.) l.c. 227;
W.J. de Wilde l.c. (1997) 141, 174; Argent et al. (eds.) l.c. 461. Lectotype (J. Sinclair, 1968): Scortechini
1851, Peninsular Malaysia, Perak, Port Weld (CAL, K). Synonym: M. hackenbergii Diels, Bot. Jahrb.
60, 3 (1926) 308, Masamune l.c. 305.
Tree up to 25 m tall; stilt roots or buttresses often present. Bark chocolate to blackish,
hard, brittle, longitudinally fissured or ridged, or scaly; inner bark slightly laminated,
pink or brown; sap watery, red. Sapwood white, turning brownish on exposure. Twigs
rather stout, towards apex 36 mm diameter, at first pubescent with rough-woolly rufous
hairs 11.5 mm long, glabrescent; bark lower down blackish, coarsely flaky; lenticels
not distinct or absent. Terminal leaf buds pubescent with hairs 11.5 mm long. Leaves
coriaceous, upper surface glossy, drying olivaceous or brown, lower surface grey-brown,
at first rusty pubescent with mealy hairs 0.10.2 mm long, very early glabrescent, distinctly
papillose; oblong to oblong-lanceolate, 1432(35) 3.59(11.5) cm, base rounded or
short-acute, apex acute-acuminate; midrib raised above; lateral veins 1522 pairs, raised
or sunken above, interarching veins usually not distinct; intercostal venation sunken above
and rendering the leaf blade somewhat bullate; petiole 2.55 cm long. Inflorescences
465
pedunculate, branched, woolly pubescent with rough hairs 11.5 mm long; bracts 35
mm long, late-caducous; males 2.55.5 23 cm, peduncle (10)1530 mm long, flower
buds of different stages of development and sizes in subumbellate clusters of 510; females
11.5 cm long, hardly branched, few-flowered. Flowers pubescent, with hairs 0.51 mm
long; bracteole subpersistent. Male flowers: pedicel 23(5) mm long; bracteole c. 3
mm diameter, apex sometimes subtruncate, late-caducous; buds ellipsoid-oblong, 44.5
3 mm, apex narrowly rounded; perianth at anthesis clefts to c. 1/4(1/3); androecium
(3)3.54 mm long; androphore cylindrical, 1.52 mm long, pubescent with dense hairs
c. 0.5 mm long in the lower 4/5; synandrium oblong, 22.2 mm long; anthers 57, sometimes
with lobed sterile apex (0.1)0.20.3 mm long. Female flowers: buds ovoid, c. 5 4
mm; ovary rusty tomentose. Infructescences 11.5 cm long. Fruits mostly solitary, ellipsoid,
(5)67 3.54 cm, velvety pubescent with persistent shaggy hairs 12.5(3) mm long;
dry pericarp (5)1015 mm thick; stalk rather thick, 25 mm long. Seeds ellipsoid, 4
5 cm long, glossy dark brown; aril pale orange or red.
Vernacular name. Sarawakkumpang pendarahan (Iban, preferred name).
Distribution. Sumatra, Peninsular Malaysia, Singapore, and the whole of Borneo.
Ecology. In lowland and hill mixed dipterocarp forests on yellow sandy soils, in peatswamp
forest, and kerangas forest, up to 800 m altitude. Flowering and fruiting apparently
14. Myristica malaccensis Hook. f.

(of Malacca, Peninsular Malaysia)
Fl. Br. Ind. 5 (1886) 104; King l.c. 287; Warburg l.c. (1897) 411; Gamble l.c. 230; Ridley l.c. 64; J.
Sinclair l.c. (1958) 356, l.c. (1968) 184, p.p. excl. most of Bornean specimens; Whitmore l.c. 344;
Anderson l.c. 269; Whitmore, Tantra & Sutisna l.c. 273; Turner l.c. 362; W.J. de Wilde l.c. (1997) 177;
Argent et al. (eds.) l.c. 461. Type: Maingay 1305, Peninsular Malaysia (holotype K; isotype CAL).
Synonym: M. pandurifolia H. Winkler, Bot. Jahrb. 49, 3 & 4 (1913) 367, Merrill l.c. (1921) 269,
Masamune l.c. 305.
Tree up to 35 m tall; buttresses spreading or not, 0.53 m high. Bark smooth with narrow
furrows or scaly, greenish grey to dark brown; inner bark pale yellow, soft; sap slow
to appear, clear, watery, faint red. Sapwood white or brown. Twigs slender, towards apex
24 mm diameter, at first tomentose with hairs less than 0.1 mm long, early glabrescent;
bark lower down drying often blackish, moderately flaky; lenticels absent. Terminal leaf
buds tomentose with minute appressed hairs up to 0.1 mm long. Leaves thinly chartaceous,
upper surface drying olivaceous-brown, lower surface glabrous (early glabrescent), drying
greyish brown, not, indistinctly or distinctly (subsp. papillosa) papillose; elliptic-oblong
to oblong, 1030 49.5 cm, base rounded or acute, apex acute-acuminate; midrib raised
above; lateral veins 1420 pairs, flat or sunken above, interarching veins raised and
distinct; intercostal venation distinct; petiole 1.22.4 cm long. Inflorescences at first
tomentose with minute hairs less than 0.1 mm long, early glabrescent; bracts 23 mm
long, caducous; males narrowly paniculate, many-flowered, 710 34 cm, peduncle
2030 mm long, flower buds of different sizes and stages of development in subumbellate
clusters of 515, largely glabrous, not angular in the upper portion; females smaller,
466
1.52.5 cm long. Male flowers tomentose with hairs up to 0.1 mm long, early glabrescent;
bracteole subcircular, c. 1.5 mm across, caducous or persistent, located at the apex of
pedicel, early glabrescent with ciliate margin; pedicel 35 mm long; buds subglobose
to ovoid, 33.5 23(3.5) mm; perianth at anthesis clefts to 1/41/3, or slightly over;
androecium 1.5(2) 0.8(1) mm, consisting almost entirely of the synandrium; anthers
68, c. 1.5 mm long, with free apices 0.10.2 mm long, sterile apex absent or apex
with shallow cavity; androphore 0.20.5 mm long, glabrous. Female flowers small; pedicel
c. 2 mm long; bracteole (1)1.5 mm long, margin ciliate, persistent; buds broadly ellipsoid,
22.5 1.5 mm; perianth at anthesis breaks off, leaving a persistent, disc-like collar
around the base of developing young fruit; ovary conical-ovoid, c. 1.3 1 mm, almost
glabrous. Infructescences 12 cm long, with few flower scars. Fruits 1 or 2 per
infructescence, ellipsoid-oblong, 46 2.54 cm, glabrous, with vestigial minute tomentum
in sheltered places near the stalk; dry pericarp 510 mm thick; stalk 58 mm long. Seeds
3.54.5 cm long.
Distribution. Sumatra, Peninsular Malaysia and Borneo. Apparently a rare species, with
two subspecies, both occurring in Sabah and Sarawak.
Ecology. In primary and disturbed forests on hillside, up to 700 m altitude. Flowering
in June; fruiting in NovemberMarch.
Key to subspecies
Leaves 1528 710 cm; intercostal venation strongly reticulate; lower surface very distinctly,
regularly, finely papillose................................
subsp. papillosa W.J. de Wilde
(Latin, papillosus = covered with papillae; the lower leaf surface)
Blumea 42 (1997) 177. Type: Ilias S. 36528, Borneo, Sarawak (holotype L; isotype SAR).
Endemic to Borneo (Sarawak).

Leaves smaller; intercostal venation faintly reticulate; lower leaf surface not or very irregularly
and faintly papillose........................................
subsp. malaccensis
Sumatra, Peninsular Malaysia and Borneo.
15. Myristica maxima Warb.

(Latin, maximum = greatest; inference unclear)
Monogr. Myrist. (1897) 385; Gamble l.c. 228; Ridley l.c. 63; J. Sinclair l.c. (1958) 339, l.c. (1968) 131;
Whitmore l.c. 345; Anderson l.c. 269; Whitmore, Tantra & Sutisna l.c. 274; Turner l.c. 362; Coode et al.
(eds.) l.c. 227; W.J. de Wilde l.c. (1997) 139, 178; Argent et al. (eds.) l.c. 461. Lectotype: Kings
Collector 5513, Peninsular Malaysia, Perak, Larut (CAL, E, FI, G, K, L, MEL, SING, UPS).
Tree up to 35 m tall; crown deep and dense, pyramidal in outline, with numerous horizontal
and descending branches; buttresses up to 3 m out and high, narrow. Bark smooth or
rather rough, hard, shallowly scaly or dippled, or shallowly fissured, brown to dark brown,
grey-brown, or chocalate; inner bark 1020 mm thick, usually reddish brown, laminated,
467
soft; sap clear, light to dark red, copious or not. Sapwood pale brown to whitish, also
exuding red sap; heartwood brown. Twigs stout, towards apex (4)58 mm diameter,
at first tomentose with minute hairs, early glabrescent; bark lower down becoming rough
and scaly, not cracked; lenticels often present but not distinct. Terminal leaf buds tomentose
with minute appressed hairs up to 0.1 mm. Leaves membranous to thinly coriaceous,
upper surface drying blackish brown, sometimes bullate, lower surface grey, at first
pubescent with scale-like hairs up to 0.1(0.2) mm long, early or late-glabrescent, usually
distinctly papillose; elliptic or oblong-lanceolate, 1640 (6)1020 cm, base rounded
or shallowly cordate, sometimes attenuate, apex acute-acuminate; midrib flat or raised
above; lateral veins (20)2330(33) pairs, impressed above, interarching veins usually
distinct on both surfaces; intercostal venation distinct on both surfaces; petiole 2.53.5
cm long. Inflorescences at first tomentose with minute, grey, villous hairs, subglabrescent,
pedunculate, branched; males 612 (18) 38 cm, peduncle 2045 mm long, flattened,
flower buds of different sizes and stages of development in subumbellate clusters of 5
10, pubescent, not angular in the apical part, bracts small, caducous; females smaller,
59 cm long, flowers in fascicles of 15, perianth coriaceous, thinly pubescent with hairs
less than 0.1 mm long. Male flowers: pedicel 310 mm long; bracteole 1.52 mm long
with ciliate margin, broadly reniform, or shallowly 23-lobed, persistent; buds ovoid or
ovoid-ellipsoid, 57 45 mm; perianth c. 0.7 mm thick at the sutures, at anthesis clefts
to c. 1/2 or slightly over; androecium 34.5(5) (0.8)11.2 mm long; androphore 0.5
0.8(1) mm long, finely pubescent in the lower half; synandrium cylindrical, 22.5 mm
long; anthers 812, sterile apex 0.30.5 mm long. Female flowers: pedicel 13(4) mm
long; bracteole broadly rounded, 2(3) mm across, persistent; buds ovoid, 57 46
mm; ovary ovoid, 34 mm across, pubescent with appressed hairs 0.20.4 mm long, stigma
conspicuous, 2-lipped, (0.5)1 mm long. Fruits 14 per infructescence, ellipsoid-oblong,
(4.5)69 3.55 cm, scurvy pubescent with persistent minute hairs up to 0.1 mm long,
sometimes late-glabrescent; dry pericarp 1015 mm thick; stalk stout, 310 mm long.
Seeds ellipsoid-oblong, (4)4.56 cm long; aril deeply divided but lower 1/4 sometimes
undivided, yellow turning to red.
Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, Singapore, and the whole
of Borneo.
Ecology. In primary lowland to hill forests, along rocky Saraca streams on granite rock,
on slopes and ridges with various types of soils, up to 700 m altitude. Flowering and
16. Myristica papyracea J. Sinclair

(Latin, papyraceus = papery; the thin dry leaves)
Gard. Bull. Sing. 23 (1968) 133; W.J. de Wilde l.c. (1997) 139, 178; Argent et al. (eds.) l.c. 461. Type:
Wood SAN A 4775, Borneo, Sabah, Lahad Datu district, Bt. Kretam (holotype SING; isotypes A, KEP,
L, MEL, SAN).
Tree up to 40 m tall; often flanged and/or with conspicuous, laterally compressed stilt
roots up to 2 m high. Bark dark grey or blackish brown, cracked, in old trees flaking
in thin papery strips; inner bark 813 mm thick, red-brown; sap pink, not copious. Sapwood
468
pale brown or dirty white, red-brown or white-streaked with red. Twigs stout, towards
apex somewhat angular, 48 mm diameter, drying yellowish brown, at first tomentose
with hairs up to 0.1 mm long, early to late-glabrescent; bark lower down often coarsely
flaky, with or without lenticels. Terminal leaf buds tomentose with minute appressed hairs
up to 0.1 mm long. Leaves thinly coriaceous, upper surface drying olivaceous, lower
surface yellowish, glabrous, distinctly papillose; elliptic-oblong or oblong-lanceolate, 16
44 814.5(18) cm, base rounded or subcordate, apex somewhat blunt or acute; midrib
above slightly raised; lateral veins (16)2028 pairs, sunken or flat above, interarching
veins raised and distinct; intercostal venation more or less scalariform, distinct or not;
petiole 2.24(5.5) cm long. Inflorescences tomentose with scurfy hairs 0.10.2(0.3)
mm long, essentially paniculate, pedunculate; males 25 24 cm, peduncle 820 mm
long, flowers few in loose clusters, buds strongly varying in size and age, largely glabrous,
not angular in the upper portion, bracts 23(4) mm long, caducous; females smaller,
perianth coriaceous, pubescent with hairs c. 0.1 mm long, sometimes late-glabrescent,
bracteole broadly reniform, 1.52 mm long, caducous. Male flowers: pedicel 58 mm
long, with bracteole scar located up to 2 mm below the perianth; buds more or less
ellipsoid, 68 45 mm; perianth at anthesis clefts to 2/33/4; androecium subcylindrical,
(4)56 1.52 mm; androphore 12 mm long, densely pubescent with hairs c. 0.1 mm
long; synandrium oblong, (2)2.53 mm long; anthers 912, sterile apex 0.30.6 mm
long. Female flowers: pedicel 24 mm long; buds ovoid, c. 6 4 mm, ovary (3)4.5
33.5 mm, pubescent with hairs up to 0.1 mm long. Infructescences 13 cm long,
with 1 or 2 fruits. Fruits ellipsoid, 79 4.56 cm, the style remnant more or less
lateral (always?), at first tomentose with scurfy dark brown hairs c. 0.1 mm long, lateglabrescent; dry pericarp 1020 mm thick; stalk stout, 410 mm long. Seeds long-ellipsoid,
56 cm long; aril pale coral-red.
Distribution. Endemic to Borneo (Sarawak and Kalimantan).
Ecology. In mixed dipterocarp forest on various types of soils, up to 300 m altitude.
17. Myristica simiarum A. DC.

(Latin, simia = ape; fruit eaten by monkeys, or inflorescence resembling an ape)
Ann. Sci. Nat. 4, 4 (1855) 29; Prod. 14, 1 (1856) 192; W.J. de Wilde l.c. (1997) 184. Type: Callery 34,
the Philippines, Luzon, Manila (holotype G; isotype P). Synonym: M. elliptica Wall. ex Hook. f. &
Thomson var. simiarum (A. DC.) J. Sinclair l.c. (1958) 356, l.c. (1968) 190.
subsp. calcarea W.J. de Wilde

(Latin, calcareus = growing on limestone)
Blumea 42 (1997) 184. Type: Kostermans 13866, Borneo, E Kalimantan, Berau (holotype L; isotypes
BO, K, SING).
Tree up to 36 m tall; buttresses up to 1 m high. Bark superficially fissured or very

rough and deeply fissured, hard, dark brown or red-brown, or black, pitted, c. 5 mm
thick, peeling off in irregular strips of 23 cm wide and 23 mm thick; inner bark 10
469
15 mm thick, undulate, yellowish brown or red-brown or dark red; sap watery, pale red.
Sapwood dirty white or yellowish white; heartwood dark brown. Twigs terete, slender,
towards apex 1.53(3.5) mm diameter, pale brown, at first pubescent with yellow-brown
hairs 0.10.2(0.4) mm long, early glabrescent; bark lower down with or without small
lenticels. Terminal leaf buds tomentose with appressed hairs 0.51 mm long. Leaves thinly
chartaceous, very brittle when dry, upper surface drying olivaceous, lower surface drying
greyish, yellow-brown pubescent with persistent, appressed silky hairs (0.1)0.30.5 mm
long, not or indistinctly papillose; ovate-elliptic or oblong-lanceolate, 1118 37 cm,
base acute or nearly rounded, apex acute-acuminate; midrib flat or slightly raised above;
lateral veins (10)1214 pairs, flat and faint above, interarching veins distinct or indistinct;
intercostal venation distinct or indistinct; petiole 1.42.5 cm long. Inflorescences lateglabrescent, slender, peduncled, branched; males paniculate, 1.54 1.53.5 cm, peduncle
715 mm long, flowers buds of different stages and sizes in clusters of 510; bracts
small, subpersistent or caducous; females like the males, but smaller and with fewer flowers;
flowers pubescent with rusty hairs 0.20.3(0.5) mm long; bracteole located at the apex
of pedicel, persistent. Male flowers: pedicel (2)35 mm long; buds ellipsoid-oblong,
triangular and somewhat acute at apex, 78 23 mm; perianth at anthesis clefts to
nearly 1/2; androecium cylindrical, (4)4.55(5.5) 0.81 mm; androphore (0.5)1
1.5 mm long, pubescent with hairs 0.31 mm long; synandrium (2.5)33.5 mm long;
anthers 812, sterile apex up to 0.5 mm long. Fruits 13 per infructescence, broadly
ellipsoid, 3.5(4) 2.53 cm, base contracted into pseudo-stalk 24 mm long, 34 mm
broad, apex shortly acute, glabrescent or thinly pubescent with inconspicuous greyish
brown silky hairs c. (0.3)0.5 mm long; dry pericarp c. 3 mm thick; stalk slender or
stout, 36 mm long. Seeds conforming to the fruits; aril bright red.
Distribution. A species with 3 subspecies, distributed in Borneo, the Philippines, Sulawesi
and N Maluku. Subspecies calcarea is endemic to Borneo (W Sarawak and Kalimantan).
Ecology. In forests on old coral-limestone and sandstone with yellow sandy soils, on
hilly country, up to 700 m altitude. Flowering in September; fruiting in JuneSeptember.
18. Myristica smythiesii J. Sinclair

(B.E. Smythies; 19591964 Conservator of Forest in Sarawak, ornithologist, botanist)
Gard. Bull. Sing. 23 (1968) 316; Anderson l.c. 269; Whitmore, Tantra & Sutisna l.c. 274; Coode et al.
(eds.) l.c. 227; W.J. de Wilde l.c. (1997) 140, 185. Type: Ashton, Smythies & Wood SAN 17440, Borneo,
Brunei, Badas FR (holotype SING; isotypes A, BO, BRIS, KEP, K, L, SAN).
Tree up to 18 m tall; sometimes with small buttresses or stilt roots. Bark smooth, with
fine longitudinal striation or narrowly fissured or flaky, dark grey or brown-black; inner
bark soft, red-brown. Sapwood soft, orange-yellow; heartwood deep red. Twigs slender,
towards apex 23 mm diameter, pubescent with hairs 0.30.5 mm long, glabrescent; bark
lower down dark brown or blackish, longitudinally cracked, sometimes flaky; lenticels
absent. Terminal leaf buds pubescent with coarse hairs 0.5(1) mm long. Leaves thinly
coriaceous, upper surface drying olivaceous, late-glabrescent on midrib towards the base,
lower surface grey or cinnamon tomentose with persistent hairs 0.10.2 mm long, mixed
with scattered emergent hairs 0.30.4 mm long, indistinctly papillose; elliptic-oblong or
470
oblong-lanceolate, 820 26 cm, base rounded or shortly acute, apex acute-acuminate

(sometimes somewhat blunt); midrib raised above; lateral veins 1215(18) pairs, faint,
flat or sunken above, interarching veins indistinct; intercostal venation indistinct or
invisible; petiole 0.71.5 cm long. Inflorescences simple or forked brachyblasts, sessile
or with stalk 11.5 mm long, pubescent with brown-red hairs 0.5(1) mm long; males
up to 0.6(1) cm long, at apex with few flower buds of varying stages of development;
bracts minute, caducous; flowers pubescent with hairs 0.30.5(1) mm long. Male flowers
(immature): pedicel (2)3 mm long; bracteole subcircular, faintly 3-lobed, c. 1.5 2
mm, subpersistent; buds ovoid to ellipsoid-oblong, 33.5 2 mm, apex rounded; perianth
at anthesis clefts to 1/4(1/3); androecium c. 1.5 0.4 mm; synandrium 11.2 mm long,
anthers 56, sterile apex blunt, c. 0.2 mm long; androphore c. 0.3 mm long, pubescent
with dense hairs c. 0.2 mm long. Female flowers not seen. Fruits solitary, almost sessile,
broadly ellipsoid, 2.53.5(3.8) 22.4 cm, densely pubescent with scurfy hairs 0.2
0.3(0.5) mm long; dry pericarp 1(2) mm thick; stalk rather stout, 36 mm long. Seeds
broadly ellipsoid, c. 2.5 cm long, red-brown, shining; aril red.
Ecology. In primary mixed dipterocarp forest on hill slopes, also in kerangas and peatswamp
forests, on yellow sandy clay or loam soils and shallow peat overlying clay, up to 700 m
altitude. Flowering and fruiting in AprilJuly.
19. Myristica villosa Warb.
Fig. 16.
(Latin, villosus = long shaggy hairy; the terminal leaf buds, flowers, fruits)
Anderson l.c. 269; Whitmore, Tantra & Sutisna l.c. 274; Coode et al. (eds.) l.c. 227; W.J. de Wilde l.c.
(1997) 140, 188; Argent et al. (eds.) l.c. 462. Lectotype: Beccari 1526, Borneo, Sarawak, Mt. Matang
(hololectotype FI; isolectotype K).
Tree up to 40 m tall, occasionally with stilt roots. Bark hard, rough, sometimes smooth
or finely scaly, blackish, flaking into strips (1.5)23 mm wide and long, and c. 1 mm
thick; inner bark reddish brown. Sapwood cream or pale brown; sap red. Twigs stout,
towards apex 510 mm diameter, at first villose with hairs (1)23(4) mm long,
glabrescent; bark lower down coarsely and longitudinally cracked and flaky; lenticels
absent. Terminal leaf buds conspicuously pubescent with hairs (1)23(4) mm long. Leaves
chartaceous or thinly coriaceous, upper surface drying olivaceous or brown, lower surface
late-glabrescent (old leaves) or with persistent tomentum of dense, grey or cinnamon hairs
0.10.2 mm long mixed with spaced unbranched uniseriate emergent hairs 13 mm long,
indistinctly papillose; elliptic-oblong or oblong, (15)2038(40) 612.5(13) cm, base
acute or rounded, apex acute-acuminate; midrib raised above; lateral veins 2025 pairs,
sunken above, interarching veins usually distinct; intercostal venation distinct; petiole
(1.2)22.5 cm long, late-glabrescent. Inflorescence a sessile scar-covered brachyblast,
pubescent; males simple or 24-furcate, 0.51.3 0.51 cm, at apices with 13 subsessile
flowers; bracts minute, caducous; females similar, shorter, with 1 or 2 flowers more or
less hidden in coarse pubescence; perianth coriaceous, tomentose with villose hairs 1
2 mm long; bracteole late-caducous. Male flowers: pedicel rather thick, 34(5) mm
471
vi
long; buds ovoid or ovoid-ellipsoid, 68 45(6) mm, apex somewhat blunt; perianth
at anthesis clefts to c. 2/3; androecium slender, 45 mm long; synandrium narrowly
cylindrical, 3.54.5 0.6 mm; anthers c. 6, sterile apex somewhat blunt, (0.2)0.30.5
mm long; androphore slender, c. (0.5)1 mm long, at base with a collar of pale hairs
c. 1 mm long. Female flowers: pedicel c. 1 mm long; buds ovoid, c. 8(10) (5)6
8 mm; ovary ovoid, 56 4 mm, pubescent with hairs 11.5 mm long. Fruits 1 or
2 per infructescence, subsessile, broadly ovoid-ellipsoid, 46(8) 2.54(4.5) cm, apex
often gradually narrowed with the tip somewhat uncinate, pubescent with hispid hairs
23(4) mm long; dry pericarp 36(8) mm thick; stalk up to 5 mm long. Seeds broadly
ellipsoid, 35 cm long; aril red, coarsely laciniate, the laciniae finely divided at the top.
Ecology. In primary hill mixed dipterocarp forest, marshy forest, and kerangas forest,
on various types of soils, up to 1200 m altitude. Flowering in February, June and August
September; fruiting in MaySeptember.
473
vi
474
ABBREVIATIONS OF
Anderson, J.A.R. 1980. A Checklist of the Trees of Sarawak. Forestry Dept., Sarawak. CLTS
Argent, G. et al. (eds.). 1997. Manual of the Larger and More Important Non-dipterocarp Trees
of Central Kalimantan, Indonesia, Volumes 1 & 2. Forest Research Institute, Samarinda,
Indonesia.
MNDT-CK
Ashton, P.S. 1988. Manual of the Non-Dipterocarp Trees of Sarawak. Forestry Dept., Sarawak.
MNDTS
the Netherlands.
FJ
Browne, F.G. 1955. Forerst of Sarawak and Brunei. Govt. Press, Sarawak.
FTSB
TBS
Lumpur.
EPMP
TS
Coode, M.J.E. et al. (eds.). 1996. A Checklist of the Flowering Plants and Gymnosperms of
Brunei Darussalam. Forestry Department, Ministry of Industry and Primary Resources, Brunei
Darussalam and Royal Botanic Gardens, Kew.
CLBD
Corner, E.J.H. 1988. Wayside Trees of Malaya. 2 Vols. Malayan Nature Society, Kuala Lumpur.
WSTM
Flora Malesiana
FM
TNB
475
Kuala Lumpur.
OFMSP
Kochummen, K.M. 1997. Tree Flora of Pasoh Forest. Malayan Forest Records No. 44, FRIM,
Malaysia.
TFPF
PB
EPB
Merrill, E.D. 1921. A Bibliographic Enumeration of Bornean Plants. J. Str. Br. Roy. As. Soc.,
Sp. No.
EB
Merrill, E.D. 1929. Plantae Elmerianae Borneenses. University of California Publications in
Botany, Vol. 15. University of California Press, Berkeley.
PEB
Ng, F.S.P. (ed.). 1978 & 1989. Tree Flora of Malaya, Vol. 3 & 4. Longman, Malaysia.
TFM
Ng, F.S.P. 1991 & 1992. Manual of Forest Fruits, Seeds and Seedlings. Malaysian Forest Records,
No. 34. Vols. 1 & 2. FRIM, Malaysia.
MFFSS
MPESA
Plant Resources of South East Asia. Wageningen. (Various authors and volumes).
PROSEA (to be listed without editor or author names, but to include volume and page numbers).
Ridley, H.N. 19231925. The Flora of the Malay Peninsula, Vols. 15. Reeves, London.
FMP
CST
Stapf, O. 1894. On the Flora of Mount Kinabalu, in North Borneo. Transactions of the Linnean
Society, London, Ser. 2, 4, pp. 69263.
FMK
Whitmore, T.C. (ed.). 1972 & 1973. Tree Flora of Malaya, Vols. 1 & 2. Longman, Malaysia.
TFM
476
CLS
Whitmore, T.C., I.G.M. Tantra & U. Sutisna. 1990. Tree Flora of Indonesia. Check List for
Kalimantan, Part 1 & Parts 2.12.2. Forest Research & Development Centre, Bogor.
CLK
Willis, J.C. 1973. A Dictionary of the Flowering Plants & Ferns, revised by H.K. Airy Shaw.
Cambridge University Press, Cambridge.
DFPF
Wong, T.M. 1982. A Dictionary of Malaysian Timbers. Malayan Forest Records No. 30. Forest
Research Institute, Kepong, Kuala Lumpur
DMT
477
478
vi
COMMONLY USED
English
Word
Malay
Abbreviation
Word
Abbreviation
Central
Bukit
Bt.
Division
Div.
Gunung
G.
East
Kampung
Kg.
Forest Reserve
FR
Sungai
Sg.
Island
Is.
Tanjung
Tg.
Mount
Mt.
National Park
NP
North-East
NE
North-West
NW
River
R.
South
South-East
SE
South-West
SW
West
479
480
vi

Names in italics here refer to either basionyms or synonyms; family names given in upper
case are those of families revised in this volume. Pages with illustrations are listed in bolds
and italics.
A
Abauria 153
Abauria excelsa 154
Afzelia 120, 122, 123
Afzelia acuminata
124
Afzelia borneensis 124
Afzelia ?coriacea 169
Afzelia palembanica 152
Afzelia retusa 150
Afzelia rhomboidea 124, 125
Afzelia rhomboidea var. praetermissa
124
Agathis 376, 384, 389
Agathis borneensis xiv
Agathis-Casuarina 366
Alangiaceae xiii
Alseodaphne kochummenii xv
Alseodaphne lucida 22
Alstonia xiv
Anacardiaceae xiv, xv
Annonaceae xiii, 336, 463
Antcha 184
Antiaris 182, 183, 184
Antiaris toxicaria 185, 186
Apalatoa bantamensis 132
Aphananthe negrosensis 331
Apocynaceae xiv
Artocarpus ix, xii, 181, 182, 183, 184,
187, 188, 189, 199, 318,319
Artocarpus subgen. Artocarpus 187, 188
Artocarpus subgen. Pseudojaca 187, 188
Artocarpus altilis 182, 189
Artocarpus anisophyllus 187, 189, 191,
193
Artocarpus anisophyllus var. anisophyllus
192
Artocarpus anisophyllus var. sessilifolius
192
Artocarpus annulatus 190, 192
Artocarpus biformis 198
Artocarpus blumei 196
Artocarpus blumei var. kunstleri 196

Artocarpus borneensis 203
Artocarpus bracteata 320
Artocarpus brunneifolia 200
Artocarpus cerifera 323
Artocarpus champeden 199
Artocarpus clementis 201
Artocarpus communis 189
Artocarpus corneri 190, 194, 195
Artocarpus cuspidatus 208
Artocarpus dadah 191, 194, 196
Artocarpus dadah var. pubescens 196
Artocarpus dasyphylla 196
Artocarpus denisoniana 198
Artocarpus dimorphophylla 208
Artocarpus elasticus 189, 191, 196
Artocarpus elongatus 330
Artocarpus excelsus 190, 197
Artocarpus forbesii 325
Artocarpus frutescens 327
Artocarpus glaucescens 198
Artocarpus glaucescens var. tephrophylla
198
Artocarpus glaucus 190, 198
Artocarpus gomeziana var. griffithii 203
Artocarpus griffithii 203
Artocarpus heterophyllus 182, 188, 190
Artocarpus hirsutissima 199
Artocarpus humilis 203
Artocarpus incisus 189
Artocarpus inconstantissima 196
Artocarpus integer 182, 189, 190, 199
Artocarpus integer var. integer 199
Artocarpus integer var. sylvestris 199
Artocarpus jaca 199
Artocarpus jarrettiae 191, 199
Artocarpus kemando 190, 191, 200
Artocarpus kertau 208
Artocarpus klidang 191
Artocarpus kunstleri 196
Artocarpus lakoocha var. malayana 196
481
Artocarpus lamellosa 203

Artocarpus lanceifolius 190, 201
Artocarpus lanceifolius subsp. clementis
201
Artocarpus lanceolata 203
Artocarpus limpato 327
Artocarpus lingnanensis 203
Artocarpus longifolius 189, 201
Artocarpus melinoxylus 191, 202
Artocarpus melinoxylus subsp.
brevipedunculatus 202
Artocarpus melinoxylus subsp.
melinoxylus 202
Artocarpus mollis 195
Artocarpus mutabilis 205
Artocarpus nitidus 190, 191, 203, 204
Artocarpus nitidus subsp. borneensis
203, 204
Artocarpus nitidus subsp. griffithii 203,
204
Artocarpus nitidus subsp. humilis 203
Artocarpus nitidus subsp. lingnanensis
203, 204
Artocarpus nitidus subsp. nitidus 204
Artocarpus obtusus 190, 204
Artocarpus odoratissimus 182, 189, 191,
205
Artocarpus peduncularis 211
Artocarpus peltatus 191, 205, 206
Artocarpus primackiana 189, 207, 212
Artocarpus pubescens 196
Artocarpus reniformis 196
Artocarpus riedelii 323
Artocarpus rigidus 189, 191, 208, 209
Artocarpus rotunda 208
Artocarpus rufescens 195
Artocarpus sampor 203
Artocarpus sarawakensis 189, 210
Artocarpus sericicarpus 196
Artocarpus superba 191
Artocarpus tamaran 189, 210
Artocarpus tampang 195
Artocarpus tarap 205
Artocarpus tephrophylla 198
Artocarpus teysmannii 190, 211
Artocarpus tomentosulus 191, 208, 212
Artocarpus tylophylla 323
Artocarpus varians 208
Artocarpus venenosa 323

Artocarpus venenosa var. tylophylla
Artocarpus woodii 323
Artocarpus zollingeriana 198
Artocarpus sp. A 212
B
Baryxylum 158
Bauhinia 120, 122
Bignoniaceae xiv
Bleekrodea 329
Bombacaceae xiii
Bosscheria 213
Bosscheria minahassae
Brackenridgea xv
Bracteolanthus
120
Burseraceae xiii, xv
226
289
C
Caesalpinia 120, 122
Caesalpinia dasyrhachis 159
Caesalpinia inermis 160
Caesalpinia sect. Peltophorum 158
Caesalpiniaceae 119
Callaeocarpus 4
Canarium xv
Cassia 122, 123, 126
Cassia javanica 126
Cassia javanica subsp. nodosa 127, 128
Cassia nodosa 128
Castanea 3, 4
Castanea acuminatissima 7
Castanea brevicuspis 12
Castanea buruana 10
Castanea cooperta 49
Castanea costata 12
Castanea furfurella 63
Castanea javanica 20
Castanea javanica var. fuscescens 20
Castanea javanica var. montana 20
Castanea latifolia 88
Castanea ?montana 20
Castanea sessilifolia 7
Castanea spectabilis 12
Castanopsis 1, 2, 3, 4, 5
Castanopsis acuminatissima 3, 4, 5, 7
Castanopsis blumeanus 40
Castanopsis borneensis 6, 8, 9
482
Castanopsis brevicuspis 12
Castanopsis buruana 7, 8
Castanopsis buruana forma grandifolia
10
Castanopsis clemensii 6, 10, 11
Castanopsis costata 2, 6, 12, 49
Castanopsis densinervia 6, 13
Castanopsis dipersispina 19
Castanopsis elmeri 14
Castanopsis encleisocarpa 56
Castanopsis endertii 5, 14
Castanopsis evansii 5, 14
Castanopsis foxworthyi 2, 6, 15, 16
Castanopsis fulva 7, 17
Castanopsis hullettii 22
Castanopsis hypophoenicea 7, 18, 19
Castanopsis javanica 2, 6, 20, 21
Castanopsis javanica var. montana 20
Castanopsis junghuhnii 7
Castanopsis kinabaluensis 15
Castanopsis lentiginosa 20
Castanopsis lucida 6, 20
Castanopsis megacarpa 6, 22
Castanopsis microphylla 6, 23
Castanopsis motleyana 2, 6, 24, 25
Castanopsis oligoneura 6, 26
Castanopsis oviformis 2, 6, 26
Castanopsis paucispina 7, 27
Castanopsis pearsonii 24
Castanopsis pedunculata 5, 28
Castanopsis penangensis
20
Castanopsis psilophylla 5, 28, 29
Castanopsis reflexa 49
Castanopsis schlenkerae 7
Castanopsis spectabilis 12
Castanopsis trisperma 12
Castanopsis turbinata 92
Castanopsis woodii 14
Casuarina 384
Celastraceae xiii, xv
Cephalotrophis 182
Chevalierodendron glabrum 331
Chrysolepis 3
Clusiaceae xvi
Colombobalanus 3
Combretaceae xiii, 269
Connaraceae xiv
Copaifera 121, 123, 128
Copaifera palustris 129, 130

Cornaceae xiv
Cotylelobium 387
Covellia 213
Covellia albipila 257
Covellia aurata 259
Covellia didyma 283
Covellia glomerata 297
Covellia lanceolata 298
Covellia lepicarpa 283
Covellia stictocarpa 302
Covellia volkameriaefolia 283
Crudia 121, 123, 131
Crudia bantamensis 132
Crudia beccarii 132, 133
Crudia bracteolosa 134
Crudia caudata 132, 133
Crudia havilandii 139
Crudia ornata 132, 134, 135
Crudia reticulata
131, 134, 136
Crudia subsimplicifolia 131, 137
Crudia tenuipes 132, 137
Crudia velutina 132, 138
Crudia venenosa 132, 139
Crudia wrayi 132, 139
Cupuliferae 1
Cyclobalanopsis 3, 95
Cyclobalanopsis argentata 99
Cyclobalanopsis gemelliflora 102
Cyclobalanopsis horsfieldii 102
Cyclobalanopsis lineata 105
Cyclobalanopsis lowii 106
Cyclobalanopsis merkusii 102
Cyclobalanopsis merrillii 108
Cyclobalanopsis seriacea 112
Cyclobalanopsis treubiana 113
Cyclobalanopsis turbinata 102
Cyclobalanus 30
Cyclobalanus bennettii 39
Cyclobalanus blumeana 40
Cyclobalanus conocarpa 48
Cyclobalanus cyrtopoda 63
Cyclobalanus daphnoidea 51
Cyclobalanus diepenhorstii 59
Cyclobalanus encleisocarpa 56
Cyclobalanus ewyckii 57
Cyclobalanus gracilis 59
Cyclobalanus hystrix 62
483
Cyclobalanus lamponga 90
Cyclobalanus leptogyne 69
Cyclobalanus muricata 90
Cyclobalanus nitida 51
Cyclobalanus omalokos 70
Cyclobalanus rassa 82
Cynometra 120, 122, 123, 140
Cynometra bijuga 142
Cynometra elmeri 140, 141, 143
Cynometra mirabilis 141, 143
Cynometra ramiflora 141, 142, 143
Cynometra ramiflora var. bifoliolata 142
Cynometra ramiflora var. ramiflora 142
Cynometra whitfordii 142
Cynomorium sylvestre 142
Dilleniaceae xiii
Diospyros kochummenii xvi
Dipterocarpaceae xiii
Dipterocarpus sarawakensis xiii
Dodonaea 256
Dryobalanops fusca 390
E
Ebenaceae xvi
Echinocalyx
166, 170
Endocomia 335, 337, 338, 339
Endocomia macrocoma 339
Endocomia macrocoma subsp. longipes
339, 341
Endocomia rufirachis 339, 340, 341
Endocomia virella 339, 342
Endospermum 273
Entada 131
Eperua rhomboidea 124
Erythrophleum densiflorum 178
Erythrophleum unijugum 178
Erythroxylaceae xvi
Erythroxylum kochummenii xvi
Eugenia xiv, 349
Euonymus xv
Euphorbiaceae 273
D
Dacrydium beccarii 386
Dacryodes xv
Dammaropsis 213
Dansera 144, 149
Dansera procera 149
Daphne 50
Dialium 121, 123, 144, 178
Dialium subgen. Dansera 144
Dialium subgen. Uittienia 178
Dialium ambiguum 147
Dialium havilandii 147
Dialium hydnocarpoides 145
Dialium indum 145, 146
Dialium indum var. bursa 146
Dialium indum var. indum 146
Dialium kingii 147
Dialium kunstleri 144, 146
Dialium kunstleri var. kunstleri 147
Dialium kunstleri var. trifoliolatum 147
Dialium laurinum var. bursa 146
Dialium maingayi 147
Dialium marginatum 146
Dialium modestum 180
Dialium patens 146
Dialium platysepalum 145, 147
Dialium praetermissum 145
Dialium procerum 144, 148, 149
Dialium silvestre 147
Dialium trifoliolatum 147
Dialium triste 147
Dialium turbinatum 146
F
FAGACEAE ix, xvi, 1
Fagaceae subfam. Castaneoideae 3
Fagaceae subfam. Fagoideae 3
Fagaceae subfam. Quercoideae 3
Fagineae 1
Fagus 3
Fagus javanica 20
Ficus ix, xii, 181, 182, 184, 213, 214,
215, 216
Ficus subgen. Ficus 213, 214, 215
Ficus subgen. Pharmacosycea 213, 214,
215
Ficus subgen. Sycomorus 213, 214, 215
Ficus subgen. Urostigma 213, 214, 215
Ficus subsect. Congestae 290
Ficus subsect. Hispidae ser.
Tuberculifasciculatae 290
Ficus subsect. Palaeomorphe 215
Ficus acamptophylla 237, 256
Ficus acidula 298
484
Ficus acuminatissima 292

Ficus adamii 228
Ficus affinis 234
Ficus alba 280
Ficus alba var. gossypina 280
Ficus alba var. nudinervis 280
Ficus albipila 221, 257
Ficus albipila var. glabra 257
Ficus allutacea 240
Ficus allutacea var. teysmanniana 240
Ficus alnifolia 287
Ficus alutacia 240
Ficus amblyphylla 288
Ficus amboinensis 314
Ficus ampelas var. bogoriensis 290
Ficus androchaete 221, 258
Ficus angulata 309
Ficus angulidens 305
Ficus annulata 230, 233
Ficus annulata var. flavescens 230
Ficus annulata var. valida 230
Ficus antoniana 309
Ficus apiculata 276
Ficus apiocarpa 238
Ficus apiocarpa var. apiocarpa 238
Ficus apiocarpa var. villosa 238
Ficus apoensis 286
Ficus areolata 240
Ficus ashtonii 216, 258
Ficus aspera var. nota 291
Ficus aurantiaca 278
Ficus aurantiacea 214, 244, 245
Ficus aurantiacea var. angustifolia 245
Ficus aurantiacea var. aurantiacea 245
Ficus aurantiacea var. parvifolia 245
Ficus aurata 220, 259
Ficus aurata var. brevipilosa 259
Ficus aurata var. densiserra 259
Ficus aurata var. longipilosa 260
Ficus aurata var. palawanensis 260
Ficus auriculifera 261
Ficus aurita 220, 260
Ficus aurita var. auriculifera 261
Ficus aurita var. aurita 261
Ficus balabacensis 230
Ficus balansae 282
Ficus barba-jovis 251
Ficus barbata 250
Ficus basidentula 264

Ficus beccarii 220, 261, 312
Ficus beccarii var. asymmetrica 261
Ficus beccarii var. beccarii 261
Ficus beccarii var. latifolia 262
Ficus benjamina 214, 217, 226, 262
Ficus binnendykii 227
Ficus binnendykii var. binnendykii 227
Ficus binnendykii var. coriacea 227
Ficus binnendykii var. latifolia 227
Ficus borneensis 222, 223
Ficus bracteata 255
Ficus bruneiensis 219, 263, 265
Ficus brunneoaurata 219, 263
Ficus burkillii 270
Ficus callicarpa 244, 245
Ficus callicarpa var. crassinervia 248
Ficus callicarpides 244
Ficus callophylla 225
Ficus callosa 218, 264, 267
Ficus carri 252
Ficus caudatifolia 243
Ficus caulocarpa 316
Ficus celebica var. kunstleri 292
Ficus cerasiformis 248
Ficus cereicarpa 221, 264
Ficus cereicarpa var. ashtonii 266
Ficus cereicarpa var. cereicarpa 266
Ficus cerifera 314
Ficus chaii 218, 266
Ficus chartacea 218, 268
Ficus chewii 226
Ficus chloroleuca 280
Ficus chrysocarpa 259, 276
Ficus chrysocarpa var. undulata 259
Ficus chrysocoma 228
Ficus clusioides 226
Ficus compressicaulis 239
Ficus concinna 234
Ficus condensa 274
Ficus confusa 245
Ficus consociata 248
Ficus consociata var. murtoni 248
Ficus copelandii 224
Ficus cordatifolia 264
Ficus cordifolia 314
Ficus cornerii 237
Ficus crassicalyx 232
485
Ficus crassiramea 232

Ficus crassitora 297
Ficus crininervia 239
Ficus cucurbitina 229
Ficus cumingii 220, 268
Ficus cumingii var. terminalifolia 269
Ficus cuneata 309
Ficus cycloneura 223
Ficus daemonum 287
Ficus dasycaula 292
Ficus decaisanei 297
Ficus delosyce 227
Ficus delosyce var. obtusa 227
Ficus deltoidea 214, 215, 216, 269
Ficus deltoidea var. angustifolia 272
Ficus deltoidea var. arenaria 270, 272
Ficus deltoidea var. borneensis 272
Ficus deltoidea var. deltoidea 271, 272
Ficus deltoidea var. intermedia 270
Ficus deltoidea var. lutescens 270
Ficus deltoidea var. motleyana 255, 270
Ficus deltoidea var. recurvata 255
Ficus deltoidea var. subhirsuta 254
Ficus densechini 240
Ficus depressa 246
Ficus detonsa 236
Ficus diamantiphylla 219, 272
Ficus diandra 241
Ficus didymophylla 302
Ficus diepenhorstii 327
Ficus discolor 276
Ficus disticha 244
Ficus disticha var. brunneinervia 244
Ficus diversifolia 269
Ficus diversifolia var. deltoidea 269
Ficus diversifolia var. latissima 269
Ficus diversifolia var. ovoidea 272
Ficus dodonaeifolia 257
Ficus driveri 245
Ficus drupacea 228
Ficus dubia 233
Ficus dulitensis 224
Ficus elastica 214
Ficus ellipsoidea 245
Ficus elliptifolia 229
Ficus endospermifolia 219, 273, 275
Ficus eucaudata 243
Ficus eumorpha 220, 273
Ficus everettii 230

Ficus excavata 240
Ficus falcata 244
Ficus filiformis 254
Ficus fistulosa 217, 274
Ficus fistulosa var. fistulosa 274
Ficus fistulosa var. tengerensis 276
Ficus flavescens 230
Ficus flavidula 276
Ficus forstenii 251
Ficus forstenii var. villosa 251
Ficus francisci 221, 276
Ficus fulva 220, 276
Ficus gamostyla 242
Ficus gelderi 222, 230
Ficus geocarpa var. uncinata 310
Ficus geocharis 219, 277
Ficus gerontocarpa 300
Ficus gibbosa 309
Ficus gibbosa var. cuneata 309
Ficus gibbosa var. dodonaefolia 309
Ficus gibbosa var. pygmea 245
Ficus gibbosa var. rigida 309
Ficus gibbosa var. unigibba 309
Ficus gibbsiae 285
Ficus glabella 316
Ficus glabella var. concinna 234
Ficus glaberrima var. bracteata 228
Ficus glandulifera 221, 278, 279
Ficus glandulifera var. glandulifera 278
Ficus glandulifera var. villosa 278
Ficus globosa 246
Ficus glomerata 297
Ficus glomerata var. elongata 298
Ficus gossypina 280
Ficus grandiflora 236
Ficus grandifolia 248
Ficus grewiaefolia 292
Ficus grewiaefolia var. angustata 292
Ficus grossivenis 250
Ficus grossularioides 214, 218, 220, 278
Ficus gul 220, 280
Ficus haematocarpa 262
Ficus haggeri 287
Ficus hasskarlii 278
Ficus hemsleyana 219, 281
Ficus henrici 298
Ficus henschelii 278
486
Ficus heteropleura 243

Ficus heteropleura var. heteropleura 243
Ficus heteropleura var. hirta 243
Ficus hirsuta 250
Ficus hispidulosa 260
Ficus humilis 290
Ficus hunteri 280
Ficus hypogea 310
Ficus ilangoides 314
Ficus iliaspaiei 242
Ficus inaequipetiolata 260
Ficus inconstans 290
Ficus inconstatissima 196
Ficus indica var. gelderi 230
Ficus indica var. sundaica 225
Ficus intermedia 243
Ficus involucrata 231
Ficus irosinensis 287
Ficus irregularis 309
Ficus iwahigensis 246
Ficus ixoroides 214, 217, 281
Ficus jaroensis 250
Ficus johnsoni 246
Ficus kerangasensis 236
Ficus kerkhovenii 233
Ficus kinabaluensis 270
Ficus laccifera 302
Ficus laevigata 314
Ficus laevis 247
Ficus laevis var. laevis 247
Ficus laevis var. tomentosa 247
Ficus lagunensis 250
Ficus lamaoensis 233
Ficus lamponga 218, 282
Ficus lamponga var. chartacea 268
Ficus lanata 251
Ficus lanata var. foveolata 251
Ficus lanata var. lanata 251
Ficus lanceolata 298
Ficus landonii 255
Ficus latifolia 309
Ficus latiuscula 309
Ficus latsoni 314
Ficus lepicarpa 218, 282
Ficus lepicarpa var. brevibracteata 283
Ficus lepicarpa var. brevipedunculata
283
Ficus lepidosa 282
Ficus leptocalama 217, 283

Ficus leptocarpa 238
Ficus leptocarpa var. borneensis 239
Ficus leptocarpa var. crassa 239
Ficus leptocarpa var. subglabra 239
Ficus leptocarpa var. timorensis 239
Ficus leptogramma 218, 284
Ficus leucopleura 302
Ficus littoralis 288
Ficus lobata 280
Ficus lobbii 292
Ficus longispathulata 264
Ficus longistipulata 232
Ficus lucescens 298
Ficus macilenta 218, 284
Ficus macilenta var. gibbsiae 285
Ficus macilenta var. ilicifolia 285
Ficus macilenta var. macilenta 285
Ficus macrocalyx 231
Ficus macrocarpa 244
Ficus macrostyla 217, 221, 285
Ficus magnoliifolia 216, 286
Ficus malaica 283
Ficus mallotoides 257
Ficus malunuensis 264
Ficus manilensis 280
Ficus manok 246
Ficus mappan 280
Ficus megaleia 219, 286
Ficus megaleia var. megaleia 287
Ficus megaleia var. multinervia 287
Ficus megaleia var. subuncinata 287
Ficus melinocarpa 220, 287
Ficus microcarpa 214, 217, 228, 288
Ficus microstoma 223
Ficus microsyce 292
Ficus microtricherinos 257
Ficus microtus 292
Ficus microtus var. borneensis 292
Ficus midotis 217, 218, 289
Ficus minahassae 221, 289
Ficus moderata 221, 290
Ficus mollis var. albipila 257
Ficus montana 220, 290
Ficus motleyana 255
Ficus neglecta 262
Ficus nervosa 286
Ficus nitida 262
487
Ficus nivea 280

Ficus nota 221, 291
Ficus obliqua 287
Ficus obpyramidata 219, 221, 291
Ficus obscura 220, 292
Ficus obscura var. angustata 292
Ficus obscura var. borneensis 292
Ficus obscura var. kunstleri 292
Ficus obscura var. lanata 293
Ficus obscura var. obscura 293
Ficus obscura var. scaberrima 292
Ficus obtusidens 305
Ficus oleifolia 214, 215, 216 217, 293
Ficus oleifolia var. calcicola 295
Ficus oleifolia var. dodonaeifolia 256
Ficus oleifolia var. impressicostata 256
Ficus oleifolia var. linearifolia 294
Ficus oleifolia var. major 293
Ficus oleifolia var. memecylifolia 295
Ficus oleifolia var. myrsinoides 295
Ficus oleifolia var. oleifolia 296
Ficus oleifolia var. riparia 294
Ficus oleifolia var. valida 296
Ficus ovoidea 272
Ficus pachyphylla 256
Ficus palaquiifolia 234
Ficus palawanensis 251
Ficus palmata 280
Ficus paloensis 300
Ficus paracamptophylla 237
Ficus paradoa 309
Ficus paramorpha 220, 296
Ficus parietalis 248
Ficus parvifolia 234
Ficus patens 276
Ficus payapa 228
Ficus pellucido-punctata 222, 230
Ficus pendens 253
Ficus peracuta 227
Ficus philippinensis 302
Ficus phlebophylla 249
Ficus pilosa 228
Ficus piperifolia 254
Ficus piperifolia var. borneensis 254
Ficus pisifera 292
Ficus pisocarpa 223
Ficus polysyce 314
Ficus procera 231
Ficus procera var. crassiramea 232

Ficus producta 269
Ficus propinqua 250
Ficus pruniformis 246
Ficus pseudocamptophylla 223
Ficus pseudorubra 223
Ficus pseudotarennifolia 217, 296
Ficus pubinervis 221, 297
Ficus pumila 214
Ficus punctata 214, 244
Ficus pyrifolia 225
Ficus quercifolia 290
Ficus racemifera 314
Ficus racemosa 214, 217, 297
Ficus racemosa var. elongata 298
Ficus radicans 243
Ficus radicans var. angulosa 305
Ficus ramentacea 239
Ficus ramentacea var. urnigera 253
Ficus raridens 305
Ficus recurva 238, 241, 254
Ficus recurva var. bridelioides 239
Ficus recurva var. elegantior 239
Ficus recurva var. lasiocarpa 239
Ficus recurva var. recurva 239
Ficus recurva var. ribesoides 239
Ficus recurva var. urnigera 253
Ficus reinwardtii 276
Ficus remblas 292
Ficus retusa 216, 298
Ficus retusa var. borneensis 299
Ficus retusa var. ovoidea 272
Ficus retusa var. pisifera 288
Ficus retusa var. retusa 299
Ficus riedelii 289
Ficus riparia 298
Ficus rubescens 225
Ficus rubrocuspidata 217, 299
Ficus rubromidotis 217, 300
Ficus rudis 280
Ficus ruficaulis 222, 300
Ficus ruficaulis var. paloensis 300
Ficus rufipila 249
Ficus ruginervia 248
Ficus sabahana 241
Ficus sagittata 239
Ficus sagittata var. adhaerens 239
Ficus sagittata var. oligosperma 239
488
Ficus sagittata var. sagittata 239

Ficus samarensis 238
Ficus sarawakensis 247
Ficus saxatalis 245
Ficus scaberrima 292
Ficus scaberrima var. angustata 292
Ficus schwarzii 217, 301
Ficus sclerocoma 290
Ficus scleroptera 264
Ficus scortechinii 218, 302
Ficus scortechinii var. lanceata 302
Ficus septica 216, 302, 303
Ficus setiflora 220, 304
Ficus setiflora var. adelpha 305
Ficus setiflora var. puberula 305
Ficus setiflora var. setiflora 304
Ficus sikkimensis 245
Ficus similis 297
Ficus sinuata 218, 305
Ficus soepadmoi 237
Ficus spanogheana 238
Ficus spathulata 272
Ficus spathulifolia 216, 305
Ficus spathulifolia var. stipitata 306
Ficus spiralis 249
Ficus stictocarpa 302
Ficus stolonifera 219, 306
Ficus stricta 226
Ficus strigosa 238
Ficus stupenda 229
Ficus stupenda var. minor 229
Ficus subcordata 242
Ficus subcordata var. malayana 242
Ficus subdenticulata 292
Ficus subfulva 221, 307
Ficus subfulva var. subfulva 307
Ficus subfulva var. villosula 307
Ficus subgelderi 222
Ficus subobliqua 309
Ficus subopaca 314
Ficus suborbicularis 276
Ficus subpedunculata 247
Ficus subracemosa 314
Ficus subrigida 239
Ficus subsidens 235
Ficus subsubulata 224
Ficus subtecta 231
Ficus subterranea 221, 307
Ficus subulata 245

Ficus sumatrana 223, 226, 228
Ficus sumatrana var. microcyse 223
Ficus sumatrana var. subsumatrana 223
Ficus sumatrana var. sumatrana 223
Ficus sundaica 225
Ficus sundaica var. beccarii 225
Ficus sundaica var. impressicostata 225
Ficus sundaica var. sundaica 225
Ficus supperforata 249, 251
Ficus tabing 249
Ficus tajam 292
Ficus tampang 195
Ficus tarennifolia 218, 296, 308
Ficus tengerensis 276
Ficus terminalifolia 269
Ficus tetangis 238
Ficus teysmanniana 240
Ficus tinctoria 217, 227, 308
Ficus tinctoria ssp. gibbosa 309
Ficus tinctoria var. gibbosa 309
Ficus tinctoria var. rigida 309
Ficus travancorica 228
Ficus treubii 219, 220, 310
Ficus trichocarpa 254
Ficus trichocarpa var. piperifolia 254
Ficus trichocarpa var. trichocarpa 254
Ficus tristaniifolia 224
Ficus tulipifera 235, 246
Ficus tylophylla 226
Ficus uncinata 219, 310
Ficus uncinata var. A 311
Ficus uncinata var. B 313
Ficus uncinata var. C 311
Ficus uncinata var. gracilis 312
Ficus uncinata var. pilosior 312
Ficus uncinata var. strigosa 312
Ficus uncinata var. subbeccarii 312
Ficus uncinata var. truncata 313
Ficus uncinata var. uncinata 311, 312,
313
Ficus uncinulata 252
Ficus uniglandulosa 224
Ficus uniglandulosa var. latior 224
Ficus uniglandulosa var. parvifolia 224
Ficus urnigera 253
Ficus urophylla 243
Ficus valida 230
489
Ficus variegata 214, 218, 314, 315

Ficus variegata var. ilangoides 314
Ficus variegata var. pilosior 314
Ficus variegata var. variegata 314
Ficus vesca 297
Ficus vidaliana 228
Ficus villamilii 228
Ficus villipes 238
Ficus villosa 250, 254
Ficus villosa var. adpressa 250
Ficus villosa var. subglobosa 250
Ficus villosa var. villosa 250
Ficus virens 218, 233, 316
Ficus virens var. glabella 316
Ficus virescens 219, 316
Ficus viridifolia 292
Ficus xylophylla 216, 222, 317
Ficus zambalensis 300
Ficus zollingeriana 223
Formanodendron 3
Gymnacranthera forbesii 345, 350

Gymnacranthera forbesii var. crassinervis
350
Gymnacranthera forbesii var. forbesii
351
Gymnacranthera murtonii 345
Gymnacranthera murtonii var. borneensis
345
Gymnacranthera ocellata 344, 346, 351
Gymnacranthera paniculata var.
zippeliana 349
Gymnacranthera stenophylla 442
Gymnacranthera suluensis 349
Gymnacranthera zippeliana 349
Gymnartocarpus 318
Gymnartocarpus triandra 325
Gymnartocarpus venenosus 323
Gymnostoma 387
H
Haplolobus xv
Hopea xv
Horsfieldia 337, 338, 352, 353, 357
Horsfieldia acuminata 373
Horsfieldia affinis 385
Horsfieldia amklaal 373
Horsfieldia amplomontana 357, 360,
361
Horsfieldia androphora 357, 360, 362
Horsfieldia borneensis 356, 359, 363
Horsfieldia brachiata 355, 358, 363
Horsfieldia brachiata var. brachiata 364
Horsfieldia brachiata var. laticostata
374
Horsfieldia brachiata var. sumatrana
384
Horsfieldia bracteosa 393
Horsfieldia bracteosa var. bracteosa
393
Horsfieldia bracteosa var. microcarya
381
Horsfieldia carnosa 355, 360, 364, 365
Horsfieldia congestiflora 373
Horsfieldia crassifolia 353, 357, 366
Horsfieldia discolor 355, 360, 367
Horsfieldia disticha 356, 361, 368
Horsfieldia endertii 356, 360, 368
Horsfieldia fragillima 354, 356, 358,
360, 369, 370
G
Gironneira glabra 331
Gomphia xv
Guatteria 463
Gymnacranthera 336, 337, 343, 344
Gymnacranthera apiculata 349
Gymnacranthera bancana 343, 344, 345,
346
Gymnacranthera bancana var. borneensis
345
Gymnacranthera contracta 345, 347
Gymnacranthera crassinervis 351
Gymnacranthera eugeniifolia 349
Gymnacranthera eugeniifolia var.
eugeniifolia 349
Gymnacranthera eugeniifolia var. griffithii
349
Gymnacranthera farquhariana 345, 348
Gymnacranthera farquhariana var.
eugeniifolia 349
farquhariana 349
griffithii 349
Gymnacranthera farquhariana var. major
349
zippeliana 349
490
Horsfieldia fulva var. paludicola 366

Horsfieldia gracilis 354, 355, 359, 371
Horsfieldia grandis 354, 358, 372
Horsfieldia irya 339, 353, 357, 373
Horsfieldia labillardieri 373
Horsfieldia laticostata 356, 360, 374
Horsfieldia lemanniana 373
Horsfieldia macilenta 356, 361, 375
Horsfieldia macrobotrys 376
Horsfieldia macrocoma 339
Horsfieldia macrocoma var. rufirachis
340
Horsfieldia montana 357, 360, 361, 375
Horsfieldia motleyi 354, 358, 376
Horsfieldia nervosa 356, 360, 377
Horsfieldia nunu 373
Horsfieldia obscura 357, 360, 378
Horsfieldia obtusa 355, 361, 379
Horsfieldia oligocarpa 355, 359, 379
Horsfieldia pallidicaula 354, 355, 358,
359, 380
Horsfieldia pallidicaula var. macrocarya
381
Horsfieldia pallidicaula var. microcarpa
381
Horsfieldia pallidicaula var. pallidicaula
381
Horsfieldia paucinervis 354, 359, 381
Horsfieldia penangiana 357, 359, 382
Horsfieldia penangiana subsp. obtusifolia
382
Horsfieldia polyspherula 356, 361, 383
Horsfieldia polyspherula var. maxima
383
Horsfieldia polyspherula var. oligocarpa
379
Horsfieldia polyspherula var. polyspherula
384
Horsfieldia polyspherula var. sumatrana
384
Horsfieldia polyspherula var. tenuifolia
394
Horsfieldia punctatifolia 357, 359, 384
Horsfieldia reticulata 354, 359, 385
Horsfieldia ridleyana 355, 361, 386
Horsfieldia rufo-lanata
354, 355, 358,
387
Horsfieldia sabulosa 352, 354, 358, 387,
388
Horsfieldia sessilifolia 354, 358, 389

Horsfieldia sparsa 393
Horsfieldia splendida 355, 358, 390, 391
Horsfieldia sterilis 353, 358, 390
Horsfieldia subalpina 357, 360, 392
Horsfieldia subalpina subsp. kinabaluensis
392
Horsfieldia subglobosa 373
Horsfieldia subglobosa var. brachiata
364
Horsfieldia sucosa 353, 358, 393
Horsfieldia sucosa subsp. bifissa 393
Horsfieldia tenuifolia 356, 361, 394, 395
Horsfieldia tristis 354, 360, 396
Horsfieldia wallichii 354, 355, 358, 359,
396
Horsfieldia xanthina 357, 360, 397
Horsfieldia xanthina subsp. macrophylla
397, 398
Horsfieldia xanthina subsp. xanthina 398
Hypericaceae xiii
I
Icacinaceae xiv
Ilex 285, 333
Inga pterocarpa 160
Intsia 120, 122, 123, 149, 150
Intsia acuminata
124
Intsia bijuga 150, 151
Intsia palembanica 150, 151, 152
Intsia plurijuga 152
Intsia retusa 150
Ipo 184
Iripa 142
Ixora 281
J
Jonesia 162
Jonesia declinata 163
Jonesia palembanica 163
K
Knema 335, 336, 337, 338, 352, 399, 400
Knema acuminata 412
Knema ashtonii 405
Knema ashtonii var. ashtonii 403, 405
Knema ashtonii var. cinnamomea 400,
405
491
Knema cenabrei 416

Knema cinerea var. alpina 418
Knema cinerea var. cordata 439
Knema cinerea var. patentinervia forma
longipedicellata 442
Knema cinerea var. sumatrana 410, 412
Knema conferta 401, 406
Knema conferta var. borneensis 420
Knema coriacea 419
Knema curtisii 402, 406
Knema curtisii var. amoena 407
Knema curtisii var. arenosa 408
Knema curtisii var. curtisii 407
Knema curtisii var. linguiformis 407,
423
Knema curtisii var. paludosa 408
Knema elmeri 400, 408
Knema emmae 403, 409
Knema galeata 400, 401, 409
Knema geminata 412
Knema gitingensis 412
Knema glauca 404, 410
Knema glauca var. glauca 411
Knema glauca var. riparia 411
Knema glauca var. typica 410
Knema glaucescens 404, 411
Knema glaucescens var. cordata 439
Knema glaucescens var. glaucescens 410
Knema glomerata 402, 403, 412
Knema heterophylla 412
Knema heterophylla var. pubescens 412
Knema hirtella 404, 413
Knema hirtella var. hirtella 413, 414
Knema hirtella var. pilocarpa 414
Knema hirtella var. stylosa 443
Knema insularis 416
Knema intermedia 402, 414
Knema kinabaluensis 403, 415
Knema korthalsii 416
Knema korthalsii subsp. korthalsii 402,
417
Knema korthalsii subsp. rimosa 400,
417
Knema kostermansiana 403, 417
Knema kunstleri 401, 418
Knema kunstleri subsp. alpina 418
Knema kunstleri subsp. coriacea 419
Knema latericia
400, 419
Knema latericia subsp. albifolia 420

Knema latericia subsp. ridleyi 420
Knema latericia subsp. ridleyi f. nana
420
Knema latericia var. albifolia 420
Knema latericia var. lunduensis 419,
426
Knema latifolia 403, 421, 422
Knema laurina 402, 421
Knema linguiformis 402, 423, 424
Knema longepilosa 400, 425
Knema lunduensis 401, 426
Knema luteola 404, 426
Knema membranifolia 400, 401, 427
Knema minima 420
Knema mogeana 404, 428
Knema muscosa 401, 428
Knema nitida 421
Knema oblongata 402, 429
Knema oblongata subsp. oblonga 429
Knema oblongata subsp. oblongata 430,
431
Knema oblongata subsp. parviflora 430
Knema oblongata subsp. pedunculata
430
Knema obovoidea 430
Knema pallens 400, 432, 433
Knema pectinata 403, 432
Knema pectinata subsp. pectinata 434
Knema pectinata subsp. vestita 434
Knema pedicellata 401, 434, 435
Knema percoriacea 401, 436
Knema percoriacea forma fusca 437
Knema percoriacea forma longepilosa
425
Knema percoriacea forma sarawakensis
437
Knema piriformis 404, 437
Knema psilantha 400, 438
Knema pubiflora 401, 438
Knema pulchra 403, 439
Knema riangensis 404, 440
Knema rufa 402, 441
Knema sericea 400, 441
Knema stenocarpa 412
Knema stenophylla 442
Knema stenophylla subsp. longipedicellata
404, 442
492
Knema stenophylla subsp. stenophylla

442
Knema stylosa 404, 443
Knema subhirtella 404, 442, 443
Knema tridactyia 444
Knema tridactyla 402, 403, 444
Knema tridactyla subsp. pachydactyla
446
Knema tridactyla subsp. salicifolia 446
Knema tridactyla subsp. sublaevis 446
Knema tridactyla subsp. tridactyla 445,
446
Knema uliginosa 402, 447
Knema umbellata 421
Knema vidalii 412
Knema viridis 401, 447, 448
Knema winkleri 421
Knema woodii 403, 449
Kochummenia xvi
Kokoona xv
Koompassia 121, 122, 153
Koompassia beccariana 156
Koompassia excelsa 153, 155,156, 257
Koompassia malaccensis 153, 154, 155
Koompassia parvifolia 154
Koompassia parviflora 154
Lithocarpus brochidodromus 33, 41, 42

Lithocarpus bullatus 2, 34, 43
Lithocarpus bulusanensis 45
Lithocarpus cantleyanus 36, 44
Lithocarpus caudatifolius 34, 44
Lithocarpus clathrata 85
Lithocarpus clementianus 36, 45
Lithocarpus clementis 76
Lithocarpus confertus 36, 46
Lithocarpus confragosus 33, 47
Lithocarpus conocarpus 2, 34, 48
Lithocarpus coopertus 33, 49
Lithocarpus corneri 32, 50
Lithocarpus crateriformis 76
Lithocarpus craterophora 93
Lithocarpus cyathiformis 59
Lithocarpus cyrtopoda 63
Lithocarpus cyrtorhyncha 59
Lithocarpus daphnoideus 36, 50
Lithocarpus dasystachyus 34, 51
Lithocarpus densiflorus 31
Lithocarpus diepenhorstii 59
Lithocarpus echinifer 32, 52, 53
Lithocarpus echinulatus 34, 54
Lithocarpus elegans 35, 55
Lithocarpus encleisacarpus 33, 56
Lithocarpus ewyckii 36, 57
Lithocarpus ferrugineus 35, 58
Lithocarpus gracilis 36, 59
Lithocarpus grandifolius 55
Lithocarpus grandifrons 90
Lithocarpus hallieri 32, 60
Lithocarpus hatusimae 35, 61
Lithocarpus havilandii 34, 62
Lithocarpus hypophoenicea 19
Lithocarpus hystrix 33, 62
Lithocarpus jacobsii 35, 63
Lithocarpus kalkmanii 32, 64
Lithocarpus keningauensis 32, 65, 66
Lithocarpus kochummenii xvi, 33, 66,
67
Lithocarpus kunstleri 80
Lithocarpus lampadarius 35, 68
Lithocarpus leptogyne 35, 69
Lithocarpus lucidus 36, 70, 71
Lithocarpus luteus 36, 72
Lithocarpus mariae 33, 73
Lithocarpus meijeri 37, 73
L
Lauraceae xiv, xv
Laurus lucida 22
Leguminosae 119, 257
LEGUMINOSAE subfam.
CAESALPINIOIDEAE 119
Leguminosae suborder Caesalpinieae
119
Lepurandra 184
Leucostegane 122, 123, 156
Leucostegane grandis 157, 158
Lithocarpus ix, 1, 2, 3, 4, 30, 31
Lithocarpus abendanonii 62
Lithocarpus andersonii 2, 35, 37
Lithocarpus argentata 99
Lithocarpus bancanus 35, 38
Lithocarpus beccarianus 32, 38
Lithocarpus bennettii 2, 35, 39
Lithocarpus blumeanus 33, 40
Lithocarpus boholensis 49
Lithocarpus borneensis 76
493
Lithocarpus melataiensis 34, 74

Lithocarpus microcalyx 55
Lithocarpus minahassae 45
Lithocarpus muluensis 34, 75
Lithocarpus nieuwenhuisii 34, 75
Lithocarpus nitida 51
Lithocarpus nodosus 35, 76
Lithocarpus oblancifolius 36, 77
Lithocarpus ochracea 76
Lithocarpus omalokos 70
Lithocarpus papillifer 2, 34, 78
Lithocarpus poculiformis 51
Lithocarpus porcatus 32, 79
Lithocarpus pruinosa 88
Lithocarpus pseudokunstleri 35, 80
Lithocarpus pseudolamponga 57
Lithocarpus pulcher 32, 80
Lithocarpus pusillus 35, 81
Lithocarpus pyriformis 85
Lithocarpus rajah 38
Lithocarpus rangeriana 82
Lithocarpus rassa 2, 35, 82
Lithocarpus reflexa
49
Lithocarpus revolutus 34, 83
Lithocarpus rhioensis 55
Lithocarpus ridleyana 82
Lithocarpus rigidus 35, 84
Lithocarpus rotundatus 32, 85
Lithocarpus ruminatus 32, 85
Lithocarpus sandakanensis 36, 86
Lithocarpus sarawakensis 51
Lithocarpus scyphigera var. riedelii 38
Lithocarpus sericobalanus 37, 87
Lithocarpus spicata var. elegans 55
Lithocarpus spicata var. glaberrima 55
Lithocarpus spicata var. gracilipes 55
Lithocarpus spicata var. placentaria 55
Lithocarpus stonei 36, 88, 89
Lithocarpus sundaicus 2, 33, 88
Lithocarpus symingtonia 82
Lithocarpus tawaiensis 2, 31, 90
Lithocarpus turbinatus 32, 91, 92
Lithocarpus urceolaris 33, 93
Lithocarpus wallichianus 34, 94
Lithocarpus wenzigiana 82
Lithocarpus winkleriana 51
Loganiaceae xiii, 187
Lysiphyllum 120
M
Maclura 182, 183, 184
Maclura cochinchinensis 183
Macrolobium bijugum 150
Magnolia 286
Magnoliaceae 286
Malaisia 182
Malaisia scandens 183
Malvaceae xiii
Melanochylla xv
Melastomataceae 295
Memecylon 295
Mesua kochummenii xvi
MORACEAE ix, xiv, xv, 181, 182, 207,
318
Moraceae tribe Artocarpeae 182
Moraceae tribe Castilleae 182
Moraceae tribe Dorstenieae 182
Moraceae tribe Ficeae 182
Moraceae tribe Moreae 182
Morus alba 182
Morus leucophylla 257
Myristica 335, 337, 344, 449, 451
Myristica sect. Fatua 451
Myristica sect. Gymnacranthera 343
Myristica sect. Myristica 450
Myristica amplifolia 345
Myristica apiculata 349
Myristica bancana 345
Myristica beccarii 453
Myristica borneensis 452, 454, 455
Myristica brachiata 364
Myristica cantleyi 423
Myristica carnosa 366
Myristica cinnamomea 451, 456
Myristica collettiana 384
Myristica conferta 406
Myristica conferta var. borneensis 420
Myristica contracta 347
Myristica cookii 463
Myristica coriacea 419
Myristica corticata 453, 457
Myristica corticosa var. decipiens 414
Myristica crassa 452, 458, 465
Myristica crassifolia 366
Myristica crassinervis 351
Myristica cumingii var. floribunda 464
Myristica curtisii 407
494
Myristica depressa 451, 459

Myristica elliptica 451, 460
Myristica elliptica var. elliptica 460
Myristica elliptica var. simiarum 469
Myristica eugeniifolia 349
Myristica extensa 452, 461
Myristica fallax 452, 461
Myristica farquhariana 348
Myristica farquhariana var. major 349
Myristica ferruginea 345
Myristica forbesii 350
Myristica fragrans 336, 451
Myristica geminata 412
Myristica gigantea 452, 453, 462
Myristica glabra var. sumatrana 384
Myristica glauca 410
Myristica glaucescens 412
Myristica grandis 372
Myristica guatteriaefolia 463
Myristica guatteriifolia
453, 463
Myristica hackenbergii 465
Myristica heterophylla 412
Myristica iners 452, 453, 464
Myristica intermedia 414
Myristica irya 373
Myristica irya var. longifolia 373
Myristica irya var. wallichii 373
Myristica javanica 373
Myristica korthalsii 416
Myristica kunstleri 418
Myristica laurina 423
Myristica laurina var. borneensis 423
Myristica littoralis 463
Myristica lowiana 453, 465
Myristica macrocoma 339
Myristica malaccensis 452, 466
Myristica malaccensis subsp. malaccensis
467
Myristica malaccensis subsp. papillosa
466, 467
Myristica maxima 451, 452, 467
Myristica motleyi 376
Myristica murtonii 345
Myristica murtonii var. borneensis 345
Myristica oligocarpa 379
Myristica palawanensis 463
Myristica paludicola 366
Myristica pandurifolia 466
Myristica papyracea 452, 468

Myristica paucinervis 381
Myristica polyspherula 383
Myristica pulchra 439
Myristica reticulata 385
Myristica ridleyana 386
Myristica ridleyi 420
Myristica riedelii 463
Myristica rubiginosa 372
Myristica simiarum 453, 469
Myristica simiarum subsp. calcarea
470
Myristica smythiesii 453, 470
Myristica spherocarpa 373
Myristica stenophylla 442
Myristica suavis 458
Myristica sublanceolata 464
Myristica sucosa 393
Myristica umbellata 421
Myristica villosa 452, 471, 472
Myristica vordermannii 464
Myristica vrieseana 373
Myristica wallichii 396
Myristica zippeliana 349
MYRISTICACEAE 335, 336
Myrsinaceae 295
Myrsine 295
Myrtaceae xiv, 224, 349
469,
N
Nothofagaceae 3
Nothofagus 3
Nyssaceae xiii
O
Ochnaceae xv
Olea 256, 293
Oleaceae 256, 293
Olmedia 182
Orthosiphon aristatus
xiii
P
Pahudia 123
Pahudia acuminata
124
Pahudia borneensis
124
Pahudia rhomboidea 124
Pahudia rhomboidea var. praetermissa
124
495
Palaquium 234
Parartocarpus 182, 183, 184, 318, 319
Parartocarpus beccarianus 323
Parartocarpus borneensis 325
Parartocarpus bracteatus 319
Parartocarpus excelsa 325
Parartocarpus microcarpus 319, 320
Parartocarpus spinulosus 319, 321, 322
Parartocarpus triandra 325
Parartocarpus venenosus 319, 323, 324
Parartocarpus venenosus subsp. borneensis
321, 323, 325
Parartocarpus venenosus subsp. forbesii
325
Parartocarpus venenosus subsp. venenosus
325
Paratrophis 329
Paratrophis caudata 333
Paratrophis glabra 331
Parishia maingayi xv
Pasania 30
Pasania acuminatissima 7
Pasania bancana 38
Pasania beccariana 39
Pasania bennettii 39
Pasania cantleyana 44
Pasania clementiana 45
Pasania clementis 76
Pasania confragosa 47
Pasania conocarpa 48
Pasania craterophora 93
Pasania cyrtorhyncha 59
Pasania daphnoidea 51
Pasania dasystachya 51
Pasania encleisocarpa 56
Pasania ewyckii 57
Pasania glomerata 55
Pasania grandifrons 90
Pasania hystrix 62
Pasania korthalsii 90
Pasania lampadaria 68
Pasania lamponga 90
Pasania lamponga var. ewyckioides 57
Pasania lucida 70
Pasania ochracea 76
Pasania oligoneura 93
Pasania omalokos 70
Pasania pruinosa 88
Pasania rajah 38
Pasania rassa 82
Pasania rotundata 85
Pasania spicata var. gracilipes 55
Pasania spicata var. microcalyx 55
Pasania spicata var. placentaria 55
Pasania sundaica 88
Pasania urceolaris 93
Pasania wallichiana 94
Pasania wenzigiana 82
Pasania winkleriana 51
Peltophorum 121, 122, 158, 159
Peltophorum dasyrhachis 159, 161
Peltophorum inerme 160
Peltophorum pterocarpum 159, 160
Peltophorum racemosum 159, 160, 161
Perrottetia xv
Phanera 120
Pharmacosycea 213
Pimelodendron xv
Pogonotrophe 213
Pogonotrophe alnifolia 287
Pogonotrophe aurantiaca 278
Pogonotrophe borneensis 254
Pogonotrophe emodi 247
Pogonotrophe emodi var. cordata 247
Pogonotrophe emodi var. oblongata 247
Pogonotrophe glandulifera 278
Pogonotrophe piperifolia 254
Pogonotrophe sumatrana 278
Poinciana 120
Polyphema champeden 199
Prainea 182, 183, 184, 326
Prainea cuspidata 327
Prainea frutescens 326, 327
Prainea limpato 326, 327, 328
Prainea multinervia 327
Proteaceae xiii
Pryona 131
Pryona bantamensis 132
Pseudosindora 129
Pseudosindora palustris 129
Pseudostreblus 329
Pseudostreblus caudatus 331
Pseudotrophis 329
Pseudotrophis laxifolia 333
Pseudotrophis mindanaensis 333
Pterolobium 121, 122
496
Pterolobium borneense
Quercus glaberrima 55
Quercus glomerata 55
Quercus gracilipes 55
Quercus gracilis 59
Quercus grandifolius 55
Quercus grandifrons 90
Quercus hallieri 60
Quercus havilandii 62
Quercus hendersoniana 105
Quercus horsfieldii 102
Quercus hypophoenicea 19
Quercus hystrix 62
Quercus hystrix var. mappacea 90
Quercus hystrix. var. longispica 94
Quercus javanica 20
Quercus junghuhnii 7
Quercus kerangensis 104
Quercus kerangasensis 97, 98, 103
Quercus kinabaluensis 2, 97, 98, 104
Quercus korthalsii 90
Quercus korthalsii var. hystrix 62
Quercus korthalsii var. mappacea 90
Quercus lampadaria 68
Quercus lamponga 69, 90
Quercus leptogyne 69
Quercus lineata 96, 99, 105
Quercus lineata var. heterochroa 105
Quercus lineata var. merkusii 102
Quercus lineata var. oxyrhyncha 105
Quercus lineata var. typica 105
Quercus lowii 2, 97, 98, 106, 107
Quercus lucida 70
Quercus mappacea 90
Quercus mappacea var. hystrix 62
Quercus merkusii 102
Quercus merrillii 96, 98, 108
Quercus mespilifolia var. borneensis 114
Quercus microcalyx 55
Quercus minahassae 45
Quercus muricata 88
Quercus nieuwenhuisii 76
Quercus nitida 51
Quercus nivea 96, 98, 108
Quercus ochracea 76
Quercus oligoneura 93
Quercus omalokos 70
Quercus oxyrhyncha 105
Quercus percoriacea 96, 98, 109
121
Q
Quercus 1, 2, 3, 94, 95, 97
Quercus subdiv. Castanopsis 4
Quercus subgen. Cyclobalanopsis 95
Quercus abendanonii 62
Quercus anceps 55
Quercus arbutifolia 100
Quercus arcaula var. microcalyx 55
Quercus arcaula var. racemosa 54
Quercus argentata 96, 98, 99
Quercus argentata var. concolor 99
Quercus bancana 38
Quercus beccariana 39
Quercus bennettii 39
Quercus blumeana 40
Quercus boholensis 49
Quercus borneensis 76
Quercus brevipetiolata 63
Quercus bulusanensis 45
Quercus cantleyana 44
Quercus caudatifolius 45
Quercus chrysotricha 96, 98, 100
Quercus clathrata 85
Quercus clementiana 45
Quercus clementis 76
Quercus confragosa 47
Quercus conocarpa 48, 69
Quercus cooperta 49
Quercus crassilamellata 102
Quercus cuneata 70
Quercus cyrtopoda 63
Quercus cyrtorhyncha 59
Quercus daphnoidea 50
Quercus dasystachya 51
Quercus depressa 55
Quercus diepenhorstii 59
Quercus discocarpa 20
Quercus echinifera 52
Quercus elegans 55
Quercus elmeri 96, 99, 101
Quercus encleisacarpa 56
Quercus ewyckii 57
Quercus fagiformis
7
Quercus fernandezii
49
Quercus gaharuensis 96, 97, 101
Quercus gemelliflora 96, 98, 102
497
R
Radermachia integra 199
Radermachia rotunda 208
Rhizophoraceae xiv
Ribes grossularia 278
Rubiaceae xvi, 281, 308
Quercus pinanga 99
Quercus placentaria 55
Quercus poculiformis 51
Quercus polyneura 105
Quercus pruinosa 49, 88
Quercus pseudomolucca var. korthalsii
90
Quercus pseudomolucca var. sundaica
88
Quercus pseudoverticillata 96, 97, 110,
111
Quercus pulchra 80
Quercus pyrifolia 55
Quercus pyriformis 85
Quercus racemosa 55
Quercus rajah 38
Quercus rassa 82
Quercus rassa var. lanuginosa 82
Quercus rassa var. montana 82
Quercus reflexa 49
Quercus rhioensis 55
Quercus rotundata 85
Quercus scyphigera var. riedelii 38
Quercus seriacea 112
Quercus sphacelata 55
Quercus spicata 55
Quercus spicata var. depressa 55
Quercus spicata var. glaberrima 55
Quercus spicata var. gracilipes 55
Quercus spicata var. latifolia 55
Quercus spicata var. microcalyx 55
Quercus spicata var. placentaria 55
Quercus spicata var. racemosa 55
Quercus subsericea 97, 98, 112
Quercus sumatrana 97, 98, 112
Quercus sundaica 88
Quercus teysmannii 45
Quercus treubiana 97, 99, 113
Quercus turbinata 102
Quercus turbinata var. crassilamellata
102
Quercus urceolaris 93
Quercus valdinervosa 96, 98, 114
Quercus varingaefolia 7
Quercus wallichiana 94
Quercus wenzigiana 82
Quercus wilhelminae 99
Quercus winkleriana 51
S
Salix 446
Santiria xv
Sapindaceae 256
Sapotaceae 234
Saraca 121, 123, 162, 163, 468
Saraca crassifolia 163
Saraca declinata 163, 164
Saraca elegans 163
Saraca elmeri 163
Saraca hullettii 163, 164, 165
Saraca lanceolata 163
Saraca longistyla 163
Saraca macroptera 163
Saraca palembanica 163
Saraca triandra 163
Scrophulariaceae 123
Serianthes gigalobium 178
Seymeria 123
Shorea xv
Shorea laevifolia 394
Simaroubaceae xiii
Sindora 122, 123, 129, 166, 167
Sindora affinis 167, 168
Sindora beccariana 167, 168, 169
Sindora coriacea 167, 168, 169
Sindora echinocalyx 167, 170
Sindora intermedia 175
Sindora irpicina 167, 168, 171
Sindora leiocarpa 167, 168, 172
Sindora mucronata 173
Sindora parvifoliola 173
Sindora velutina 167, 168, 172
Sindora wallichii 167, 168, 170, 173,
174
Sindora wallichii var. intermedia 175
Sloetia 329
Sloetia elongata 330
Sloetia pinangeana 330
Sloetia sideroxylon 330
Sloetia wallichii 330
498
Sterculia decandra 412

Sterculia glomerata 412
Sterculiaceae xiii
Streblus 181, 182, 184, 329, 330
Streblus elongatus 330
Streblus glaber 330, 331, 332
Streblus ilicifolius 330, 333
Streblus laevifolius 331
Streblus macrophyllus 330, 333
Strychnos 187
Sycomorphe 213
Sycomorus riparia 298
Sympetalandra 121, 122, 123, 175, 176
Sympetalandra borneensis 176, 178
Sympetalandra unijuga 176, 177, 178
Symplocaceae xiv
Synaedrys 30
Synaedrys bancana 38
Synaedrys beccariana 39
Synaedrys bennettii 39
Synaedrys blumeanus 40
Synaedrys cantleyana 44
Synaedrys caudatifolia 45
Synaedrys clathrata 85
Synaedrys clementi 76
Synaedrys clementiana 45
Synaedrys conocarpa 48
Synaedrys cooperta 49
Synaedrys cyrtorhyncha 59
Synaedrys daphnoidea 51
Synaedrys dasystachya 51
Synaedrys diepenhorstii 59
Synaedrys encleisocarpa 56
Synaedrys ewyckii 57
Synaedrys fagiformis 7
Synaedrys grandifrons 90
Synaedrys hallieri 60
Synaedrys hystrix 62
Synaedrys lampadaria 68
Synaedrys lamponga 90
Synaedrys lucida 70
Synaedrys nieuwenhuisii 76
Synaedrys omalokos 70
Synaedrys poculiformis 51
Synaedrys pruinosa 88
Synaedrys pulchra 80
Synaedrys pyriformis 85
Synaedrys rajah 38
Synaedrys rassa 82
Synaedrys reflexa 49
Synaedrys rotundata 85
Synaedrys seriacea 112
Synaedrys sundaica 88
Synaedrys wallichiana 94
Synaedrys wenzigiana 82
Synoecia 213
Synoecia apiocarpa 238
Synoecia diversifolia var. angustifolia 272
Synoecia sumatrana 245
T
Tarenna 308
Taxotrophis 329
Taxotrophis balansae 333
Taxotrophis ilicifolia 333
Taxotrophis macrophylla 333
Taxotropis obtusa 333
Taxotropis triapiculata 333
Terminalia 269
Thymelaeaceae 50
Tiliaceae xiii
Touchiroa bantamensis 132
Toxicaria 184
Trigonobalanus 1, 2, 3, 115
Trigonobalanus doichangensis 115
Trigonobalanus excelsa 115
Trigonobalanus verticillata 115, 116,
117
Tristania 224
Tristaniopsis 387
Trophis 182, 184
Trophis scandens 183
U
Uittienia 121, 123, 178
Uittienia modesta 179, 180
Ulmaceae 182
Urostigma acamptophyllum 256
Urostigma amblyphyllum 288
Urostigma annulatum 230
Urostigma apiocarpa 238
Urostigma benjaminum 262
Urostigma binnendykii 227
Urostigma biverrucellum 230
Urostigma bracteatum 255
Urostigma callophyllum 226
499
Urostigma caulocarpum 316

Urostigma clusioides 226
Urostigma concinnum 234
Urostigma consociatum 248
Urostigma crassirameum 232
Urostigma cycloneuron 223
Urostigma diepenhorstii 327
Urostigma euneuron 286
Urostigma giganteum 229
Urostigma globosum 246
Urostigma haematocarpum 262
Urostigma involucratum 231
Urostigma javanicum 314
Urostigma lucescens 298
Urostigma manok 246
Urostigma microcarpum 288
Urostigma modestum 286
Urostigma monadenum 223
Urostigma neglectum 262

Urostigma nitidum 262
Urostigma onustum 246
Urostigma peracutum 227
Urostigma pisiferum 288
Urostigma pisocarpum 223
Urostigma procerum 231
Urostigma pseudorubrum 225
Urostigma pyrifolium 225
Urostigma rubescens 225
Urostigma strictum 226
Urostigma sumatranum 223
Urostigma sundaicum 225
Urostigma zollingeriana 223
Urticaceae 181, 182
V
Verbenaceae
500
xiv

Scientific names in brackets are the correct names corresponding to vernacular names listed
below.
A
alanagni (Myristica guatteriifolia A. DC.) 464
andai (Ficus midotis Corner) 289
angar-angar (Crudia reticulata Merr.) 136
antjar (Antiaris Lesch.) 184
apas (Koompassia malaccensis Maingay ex Benth.)
ara (Ficus L.) 213, 214
ara (Ficus deltoidea Jack) 270
ara (Ficus lamponga Miq.) 282
ara (Ficus oleifolia King) 294
ara (Ficus retusa L.) 299
ara (Ficus schwarzii Koord.) 301
ara (Ficus spathulifolia Corner) 306
asoka (Saraca L.) 162
156
B
babai (Saraca L.) 162
babai (Saraca declinata (Jack) Miq.) 165
babayi (Castanopsis motleyana King) 24
bakis (Saraca declinata (Jack) Miq.) 165
balanut (Saraca declinata (Jack) Miq.) 165
bani (Ficus benjamina L.) 262
bangas (Lithocarpus dasystachyus (Miq.) Rehder) 52
bangkil (Parartocarpus venenosus (Zoll. & Moritzi) Becc.) 323
barung-barung (Cassia javanica L. subsp. nodosa (Buch.-Ham. ex Roxb.) K. Larsen & S.S.
Larsen) 128
bedayang (Castanopsis hypophoenicea (Seemen) Soepadmo) 19
beluing (Castanopsis foxworthyi Schottky) 17
beluno (Artocarpus integer (Thunb.) Merr.) 199
beranang (Castanopsis hypophoenicea (Seemen) Soepadmo) 19
berangan (Castanopsis (D. Don) Spach) 2, 4
berangan (Castanopsis borneensis King) 8
berangan (Castanopsis pedunculata Soepadmo) 28
berangan beluyan (Trigonobalanus Forman) 115
berangan beluyan (Trigonobalanus verticillata Forman) 116
berangan bo (Castanopsis hypophoenicea (Seemen) Soepadmo) 19
berangan bris (Lithocarpus pusillus Soepadmo) 82
berangan bu (Quercus percoriacea Soepadmo) 110
berangan daun besar (Castanopsis motleyana King) 24
berangan entadu (Castanopsis psilophylla Soepadmo) 30
berangan lingkau (Castanopsis foxworthyi Schottky) 17
501
berangan padi (Castanopsis borneensis King) 8

berangan padi (Castanopsis fulva Gamble) 17
berangan paya (Castanopsis foxworthyi Schottky) 17
berangan pipit (Catanopsis oviformis Soepadmo) 27
beringin (Ficus benjamina L.) 262
berungulad (Lithocarpus elegans (Blume) Hatus. ex Soepadmo) 56
beruni (Artocarpus J.R. & G. Forster) 187, 188
beruni (Artocarpus dadah Miq.) 196
beruni (Artocarpus nitidus Trc.) 203
beruni (Artocarpus primackiana Kochummen) 207
beruni (Artocarpus rigidus Blume) 210
beruni (Prainea limpato (Miq.) Buemee ex Heyne) 329
bi-n (Saraca hullettii Prain) 165
binarah (Myristicaceae) 335
binarak (Horsfieldia endertii W.J. de Wilde) 369
bindara (Myristicaceae) 335
bintau (Artocarpus anisophyllus Miq.) 192
bintawak (Artocarpus anisophyllus Miq.) 192
biyau (Saraca L.) 162
bokis (Saraca declinata (Jack) Miq.) 165
borak (Sindora Miq.) 166
buah abung (Ficus beccarii King) 261
buah abung (Ficus geocharis Corner) 277
buah klidang (Parartocarpus venenosus (Zoll. & Moritzi) Becc.) 323
buah tepur (Ficus tarennifolia Corner) 308
bukoh (Artocarpus annulatus Jarrett) 194
bukoh (Artocarpus integer (Thunb.) Merr.) 199
buling (Castanopsis evansii Elmer) 15
busuk-busuk (Cassia javanica L. subsp. nodosa (Buch.-Ham. ex Roxb.) K. Larsen & S.S. Larsen)
128
butoh terampayoh (Castanopsis oviformis Soepadmo) 27
C
chempedak (Artocarpus integer (Thunb.) Merr.) 189, 199
chempedak ayer (Artocarpus teysmannii Miq.) 211
copaiba (Copaifera L.) 128
D
dadah (Artocarpus dadah Miq.) 194, 196
dadah (Artocarpus primackiana Kochummen) 207
dadak (Artocarpus dadah Miq.) 196
dadak (Artocarpus nitidus Trc.) 203
darah-darah (Gymnacranthera (A. DC.) Warb.) 343
darah-darah (Gymnacranthera forbesii (King) Warb. var. forbesii)
darah-darah (Horsfieldia Willd.) 352
darah-darah (Horsfieldia grandis (Hook. f.) Warb.) 372
darah-darah (Knema Lour.) 399
darah-darah (Myristica Gronov.) 449
502
351
darah-darah (Myristica iners Blume) 465

darah-darah (Myristicaceae) 335
darah-darah kerantu (Knema laurina (Blume) Warb.) 423
denging (Artocarpus anisophyllus Miq.) 192
dialion (Dialium L.) 144
diparak (Lithocarpus gracilis (Korth.) Soepadmo) 60
dulanut (Saraca declinata (Jack) Miq.) 165
dunduk (Ficus obscura Blume) 293
dunuk (Ficus benjamina L.) 262
E
empaka (Artocarpus nitidus Trc.) 203
empatak (Prainea limpato (Miq.) Beumee ex Heyne) 329
empeli (Quercus kinabaluensis Soepadmo) 105
empenit (Lithocarpus Blume) 30
empenit (Quercus L.) 94
empenit (Quercus kerangasensis Soepadmo) 104
empenit batu (Lithocarpus andersonii Soepadmo) 37
empenit daun halus (Lithocarpus pusillus Soepadmo) 82
empenit gading (Lithocarpus pusillus Soepadmo) 82
empenit jangkar (Lithocarpus andersonii Soepadmo) 37
empenit johari (Lithocarpus pseudokunstleri A. Camus) 80
empenit padang (Lithocarpus andersonii Soepadmo) 37
empenit padang (Lithocarpus dasystachyus (Miq.) Rehder) 52
empili (Lithocarpus Blume) 30
empili (Quercus L.) 94
empili (Quercus argentata Korth.) 100
empili (Quercus merrillii Seemen) 108
empili (Quercus percoriacea Soepadmo) 110
empili (Quercus treubiana Seemen) 114
empili batu (Lithocarpus nieuwenhuisii (Seemen) A. Camus) 76
empili pipit (Lithocarpus andersonii Soepadmo) 37
empili pipit (Lithocarpus bennettii (Miq.) Rehder) 40
engkabuloh (Ficus schwarzii Koord.) 301
engkeruroh (Ficus schwarzii Koord.) 301
engkunoh (Ficus fistulosa Reinw. ex Blume) 274
engkunoh (Ficus obpyramidata King) 292
engkunoh (Ficus schwarzii Koord.) 301
engkunoh (Ficus scortechinii King var. lanceata Corner) 302
engkuroh (Ficus fistulosa Reinw. ex Blume) 274
engkusuroh (Ficus lepicarpa Blume) 283
ensurut (Sindora leiocarpa Backer ex de Wit) 172
entawa (Artocarpus anisophyllus Miq.) 192
entimau (Ficus beccarii King) 261
entimau (Ficus francisci H. Winkl.) 276
entimau (Ficus geocharis Corner) 277
entimau (Ficus stolonifera King) 306
epil (Intsia bijuga (Colebr.) Kuntze) 150
503
G
gaparang (Saraca L.) 162
gapis (Saraca L.) 162
gapis (Saraca declinata (Jack) Miq.)
gili-gili (Streblus Lour.) 329
gul (Ficus gul K. Schum. & Lauterb.)
165
280
I
impas (Koompassia Maingay ex Benth.) 153
impas (Koompassia malaccensis Maingay ex Benth.)
ipil (Intsia bijuga (Colebr.) Kuntze) 150
ipil (Intsia palembanica Miq.) 152
ipil darat (Afzelia rhomboidea (Blanco) Vidal) 126
ipil laut (Intsia bijuga (Colebr.) Kuntze) 150
ipoh (Antiaris Lesch.) 184
ipoh (Antiaris toxicaria Lesch.) 187
irya (Horsfieldia irya (Gaertn.) Warb.) 373
J
jambakia (Saraca declinata (Jack) Miq.)
jela bala (Myristicaceae) 335
156
165
K
kachang kachang (Saraca declinata (Jack) Miq.) 165
kaher (Parartocarpus bracteatus (King ex Hook. f.) Becc.) 320
kakang (Artocarpus tamaran Becc.) 211
kalan (Lithocarpus leptogyne (Korth.) Soepadmo) 70
kaliput (Artocarpus lanceifolius Roxb.) 201
kalit (Lithocarpus jacobsii Soepadmo) 64
kalong (Artocarpus tamaran Becc.) 211
kamuning (Streblus macrophyllus Blume) 334
kara (Ficus deltoidea Jack) 270
kara (Ficus fistulosa Reinw. ex Blume) 274
kara (Ficus hemsleyana King) 281
kara (Ficus lepicarpa Blume) 283
kara (Ficus midotis Corner) 289
kara (Ficus obscura Blume) 293
kara (Ficus rubrocuspidata Corner) 300
kara (Ficus rubromidotis Corner) 300
karen (Prainea limpato (Miq.) Beumee ex Heyne) 329
karon (Artocarpus nitidus Trc.) 203
karon (Prainea limpato (Miq.) Beumee ex Heyne) 329
kassia (Cassia L.) 126
kasusur aboi (Prainea limpato (Miq.) Beumee ex Heyne) 329
kateh (Parartocarpus bracteatus (King ex Hook. f.) Becc.) 320
kateh (Parartocarpus microcarpus Corner) 321
kateh (Parartocarpus venenosus (Zoll. & Moritzi) Becc.) 323
kati (Parartocarpus microcarpus Corner) 321
504
kati (Parartocarpus venenosus (Zoll. & Moritzi) Becc.) 323

katong-katong (Cynometra L.) 140
katong-katong (Cynometra mirabilis Meeuwen) 141
katong-katong (Cynometra ramiflora L. var. ramiflora) 142
kayo bakek (Artocarpus anisophyllus Miq.) 192
kayo bela (Myristicaceae) 335
kayo bibungan (Artocarpus anisophyllus Miq.) 192
kayo raha (Myristicaceae) 335
kayu ara (Ficus L.) 213, 214
kayu ara (Ficus callosa Willd.) 264
kayu kikai (Lithocarpus andersonii Soepadmo) 37
kayu susu (Prainea limpato (Miq.) Beumee ex Heyne) 329
kayu tanyit (Ficus oleifolia King) 294
keiran (Artocarpus odoratissimus Blanco) 205
kekatong (Cynometra L.) 140
kelangan (Saraca L.) 162
kelat (Crudia ornata de Wit) 136
keledang (Artocarpus J.R. & G. Forster) 188
keledang (Artocarpus lanceifolius Roxb.) 201
kelidang (Artocarpus anisophyllus Miq.) 192
kemando (Artocarpus kemando Miq.) 200
kempan (Ficus bruneiensis Corner) 263
kempas (Koompassia Maingay ex Benth.) 153
kempas (Koompassia malaccensis Maingay ex Benth.) 156
keraki (Lithocarpus cantleyanus (King ex Hook. f.) Rehder) 44
keraki (Lithocarpus hallieri (Seemen) A. Camus) 60
keraki (Lithocarpus pulcher (King) Markgr.) 81
keraki boras (Lithocarpus andersonii Soepadmo) 37
keraki padi (Lithocarpus rassa (Miq.) Rehder) 83
keranji (Dialium L.) 144
keranji (Dialium indum L.) 146
kesusu (Prainea limpato (Miq.) Beumee ex Heyne) 329
kian (Artocarpus elasticus Reinw. ex Blume) 197
kihan (Artocarpus tamaran Becc.) 211
kogopon (Castanopsis buruana Miq.) 10
kogopon (Castanopsis clemensii Soepadmo) 12
kogopon (Castanopsis endertii Hatus. ex Soepadmo) 14
kogopon (Castanopsis microphylla Soepadmo) 24
kogopon (Castanopsis psilophylla Soepadmo) 30
kukan (Ficus grossularioides Burm. f.) 280
kukor (Artocarpus altilis (Parkinson) Fosberg) 189
kumpan (Ficus androchaete Corner) 258
kumpan (Ficus aurata (Miq.) Miq.) 259
kumpan (Ficus bruneiensis Corner) 263
kumpan (Ficus grossularioides Burm. f.) 280
kumpang (Gymnacranthera (A. DC.) Warb.) 343
kumpang (Gymnacranthera forbesii (King) Warb. var. forbesii) 351
kumpang (Knema Lour.) 399
505
kumpang (Myristica Gronov.) 449

kumpang (Myristicaceae) 335
kumpang daun panjang (Knema intermedia (Blume) Warb.) 415
kumpang ensluai (Knema glauca (Blume) Petermann var. riparia W.J. de Wilde) 411
kumpang kabang (Knema uliginosa J. Sinclair) 447
kumpang limau (Endocomia macrocoma (Miq.) W.J. de Wilde subsp. longipes W.J. de Wilde)
340
kumpang paya (Horsfieldia crassifolia (Hook. f. & Thomson) Warb.) 367
kumpang pendarahan (Myristica lowiana King) 466
kumpang putih (Gymnacranthera farquhariana (Hook. f. & Thomson) Warb.) 348
kumpang tembaga (Horsfieldia sessilifolia W.J. de Wilde) 390
kumpas (Koompassia malaccensis Maingay ex Benth.) 156
kumpat (Castanopsis evansii Elmer)
15
kupan (Ficus brunneoaurata Corner) 264
kupang (Peltophorum pterocarpum (DC.) Backer ex. K. Heyne)
160
kupat (Castanopsis javanica (Blume) A. DC.) 20
L
lahu (Gymnacranthera forbesii (King) Warb. var. forbesii) 351
lampasih (Streblus macrophyllus Blume) 334
lanau (Gymnacranthera contracta Warb.) 347
langgong (Cassia javanica L. subsp. nodosa (Buch.-Ham. ex Roxb.) K. Larsen & S.S. Larsen)
128
langsat monyet (Saraca declinata (Jack) Miq.) 165
languwai (Lithocarpus pusillus Soepadmo) 82
lelingkan (Ficus grossularioides Burm. f.) 280
lemelai (Intsia palembanica Miq.) 152
lempoyang paya (Horsfieldia irya (Gaertn.) Warb.) 373
lenakan (Ficus grossularioides Burm. f.) 280
lengan (Ficus fulva Reinw. ex Blume) 277
lengkan (Ficus grossularioides Burm. f.) 280
liang (Horsfieldia androphora W.J. de Wilde) 362
limpatu (Prainea limpato (Miq.) Beumee ex Heyne) 327
lumau (Ficus magnoliifolia Blume) 286
lunok (Ficus androchaete Corner) 258
lunok (Ficus deltoidea Jack) 270
lunok (Ficus hemsleyana King) 281
lunok (Ficus microcarpa L. f.) 288
lunok (Ficus oleifolia King) 294
M
maguwau (Crudia reticulata Merr.) 136
maid maid (Crudia subsimplicifolia Merr.) 137
mangaris (Koompassia excelsa (Becc.) Taub.) 154
marang parang (Saraca declinata (Jack) Miq.) 165
maruwit (Castanopsis foxworthyi Schottky) 17
mempening (Lithocarpus Blume) 2, 30
mempening (Quercus L.) 2, 94
mempening (Quercus argentata Korth.) 100
506
mempening (Quercus chrysotricha A. Camus) 100

mempening (Quercus elmeri Merr.) 101
mempening (Quercus gaharuensis Soepadmo) 102
mempening (Quercus gemelliflora Blume) 103
mempening (Quercus kerangasensis Soepadmo) 104
mempening (Quercus kinabaluensis Soepadmo) 105
mempening (Quercus lineata Blume) 106
mempening (Quercus lowii King) 106
mempening (Quercus merrillii Seemen) 108
mempening (Quercus nivea King) 109
mempening (Quercus percoriacea Soepadmo) 110
mempening (Quercus pseudoverticillata Soepadmo) 110
mempening (Quercus subsericea A. Camus) 112
mempening (Quercus sumatrana Soepadmo) 113
mempening (Quercus treubiana Seemen) 114
mempening (Quercus valdinervosa Soepadmo) 115
mempening babi (Trigonobalanus Forman) 115
mempening babi (Trigonobalanus verticillata Forman) 116
mengarung (Castanopsis psilophylla Soepadmo) 30
menggris (Koompassia Maingay ex Benth.) 153
menggris (Koompassia malaccensis Maingay ex Benth.) 156
mengkusub (Trigonobalanus verticillata Forman) 116
mentawa (Artocarpus anisophyllus Miq.) 192
merbau (Intsia Thouars) 149
merbau ayer (Intsia palembanica Miq.) 152
merbau bukit (Intsia palembanica Miq.) 152
merbau lalat (Sympetalandra borneensis Stapf) 176
merbau lalat (Sympetalandra unijuga (Airy Shaw) Steenis) 178
merbau paya (Intsia bijuga (Colebr.) Kuntze) 150
merbau tanduk (Afzelia rhomboidea (Blanco) Vidal) 126
merubi (Artocarpus dadah Miq.) 196
merubi (Artocarpus glaucus Blume) 198
minggi (Parartocarpus Baill.) 318
minggi (Parartocarpus bracteatus (King ex Hook f.) Becc.) 320
minggi (Parartocarpus microcarpus Corner) 321
minggi (Parartocarpus venenosus (Zoll. & Moritzi) Becc.) 323
N
nakan (Artocarpus integer (Thunb.) Merr.) 199
nangka (Artocarpus heterophyllus Lamarck) 190
ngidunuk (Artocarpus nitidus Trc.) 203
nunuk daun berduri (Streblus ilicifolius (Vidal) Corner)
333
P
padeh siah (Lithocarpus conocarpus (Oudem.) Rehder) 49
pala (Myristica fragrans Houtt.) 336
pala munsoh (Artocarpus melinoxylus Gagnep. subsp. brevipedunculatus Jarrett)
pala munsoh (Artocarpus rigidus Blume) 208
pala tupai (Artocarpus lanceifolius Roxb.) 201
507
203
pala tupai (Artocarpus melinoxylus Gagnep. subsp. brevipedunculatus Jarrett)

palan (Lithocarpus conocarpus (Oudem.) Rehder) 49
palan baboi (Quercus percoriacea Soepadmo) 110
pang (Myristicaceae) 335
pasoi (Saraca declinata (Jack) Miq.) 165
patat (Artocarpus annulatus Jarrett) 194
pedalai (Artocarpus elasticus Reinw. ex Blume) 197
pedalai (Artocarpus sarawakensis Jarrett) 210
pedayang (Castanopsis hypophoenicea (Seemen) Soepadmo) 19
penarahan (Gymnacranthera (A. DC.) Warb.) 343
penarahan (Knema Lour.) 399
penarahan (Myristica Gronov.) 449
penarahan (Myristicaceae) 335, 336
pendarahan (Myristicaceae) 335
penarahan pianggu (Horsfieldia irya (Gaertn.) Warb.) 373
pening-peningan (Lithocarpus leptogyne (Korth.) Soepadmo) 70
penyibong (Lithocarpus andersonii Soepadmo) 37
peranak (Ficus schwarzii Koord.) 301
perupak (Gymnacranthera ocellata R.T.A. Schouten) 352
petir (Sindora Miq.) 166
pien (Artocarpus elasticus Reinw. ex Blume) 197
pili (Lithocarpus pusillus Soepadmo) 82
pingan (Artocarpus odoratissimus Blanco) 205
pingan (Artocarpus sarawakensis Jarrett) 210
pinggi (Parartocarpus Baill.) 318
pinggi (Parartocarpus bracteatus (King ex Hook f.) Becc.) 320
pinggi (Parartocarpus venenosus (Zoll. & Moritzi) Becc.) 323
pipin (Ficus schwarzii Koord.) 301
piraus (Lithocarpus conocarpus (Oudem.) Rehder) 49
pisau-pisau (Horsfieldia laticostata (J. Sinclair) W.J. de Wilde) 374
pisiah (Horsfieldia Willd.) 352
pudau (Artocarpus J.R. & G. Forster) 187, 188
pudau (Artocarpus kemando Miq.) 200
pudau (Artocarpus rigidus Blume) 208
pudau putih (Artocarpus glaucus Blume) 198
pudu (Artocarpus kemando Miq.) 200
puduh (Artocarpus kemando Miq.) 200
pulon (Artocarpus altilis (Parkinson) Fosberg) 189
pulutan (Artocarpus integer (Thunb.) Merr.) 199
pumpu (Myristicaceae) 335
punoh (Ficus aurita Reinw. ex Blume) 260
pupud (Artocarpus kemando Miq.) 200
puro (Ficus magnoliifolia Blume) 286
puroh (Artocarpus kemando Miq.) 200
R
raha meban (Myristicaceae) 335
rasa (Lithocarpus rassa (Miq.) Rehder)
82
508
203
S
salad (Lithocarpus Blume) 30
salad (Lithocarpus hallieri (Seemen) A. Camus) 60
salad (Lithocarpus hatusimae Soepadmo) 61
salad (Lithocarpus kochummenii S. Julia & Soepadmo) 68
salad (Quercus percoriacea Soepadmo) 110
salad kup (Lithocarpus lampadarius (Gamble) A. Camus) 69
salad padeh (Lithocarpus papillifer Soepadmo) 78
salad repak (Lithocarpus ewyckii (Korth.) Rehder) 58
salad rettan (Trigonobalanus verticillata Forman) 116
salad urong (Lithocarpus pulcher (King) Markgr.) 81
saled (Castanopsis paucispina Soepadmo) 28
saled (Lithocarpus conocarpus (Oudem.) Rehder) 49
saled (Lithocarpus gracilis (Korth.) Soepadmo) 60
saled (Lithocarpus pusillus Soepadmo) 82
saled arong (Lithocarpus jacobsii Soepadmo) 64
saled arong (Lithocarpus lucidus (Roxb.) Rehder) 72
saled birar (Lithocarpus confertus Soepadmo) 47
saled kup (Lithocarpus hallieri (Seemen) A. Camus) 60
saled urong (Lithocarpus revolutus Hatus. ex Soepadmo) 84
saled urong (Lithocarpus stonei S. Julia & Soepadmo) 88
salud kup (Lithocarpus ruminatus Soepadmo) 86
sampa rantu (Sindora beccariana Backer ex de Wit) 169
sampandai (Ficus cereicarpa Corner) 266
sansag (Crudia ornata de Wit) 136
sansag (Crudia venenosa de Wit) 139
sapetir (Sindora coriacea (Baker) Prain) 170
sarason (Crudia subsimplicifolia Merr.) 137
sarusup tembaung (Lithocarpus lucidus (Roxb.) Rehder) 72
selangapid (Crudia reticulata Merr.) 136
selangkin (Artocarpus nitidus Trc.) 203
selangking (Artocarpus J.R. & G. Forster) 187, 188
selangking (Artocarpus dadah Miq.) 196
selangking (Artocarpus glaucus Blume) 198
selangking (Artocarpus rigidus Blume) 210
selangking (Parartocarpus bracteatus (King ex Hook f.) Becc.)
selangking (Prainea King) 326
selangking (Prainea frutescens Becc.) 327
selangking (Streblus glaber (Merr.) Corner) 331
selangking batu (Prainea frutescens Becc.) 327
selibut (Artocarpus kemando Miq.) 200
selungapid (Saraca declinata (Jack) Miq.) 165
sepetir (Sindora Miq.) 166
sepetir (Sindora beccariana Backer ex de Wit) 169
sepetir (Sindora coriacea (Baker) Prain) 170
sepetir (Sindora irpicina de Wit) 171
sepetir (Sindora leiocarpa Backer ex de Wit) 172
sepetir (Sindora velutina Baker) 173
509
320
sepetir paya (Copaifera palustris (Symington) de Wit) 129

sepetir paya (Sindora leiocarpa Backer ex de Wit) 172
sepitir (Sindora affinis de Wit) 168
sikut nabalu (Endocomia rufirachis (J. Sinclair) W.J. de Wilde)
sinampar (Sindora beccariana Backer ex de Wit) 169
sindur (Sindora Miq.) 166
sindur (Sindora beccariana Backer ex de Wit) 169
sinojoh (Artocarpus nitidus Trc.) 203
soka (Saraca L.) 162
sukun (Artocarpus altilis (Parkinson) Fosberg) 189
sulok kondat (Ficus oleifolia King) 294
342
T
taalan (Lithocarpus ewyckii (Korth.) Rehder) 58
taburakin (Artocarpus nitidus Trc.) 203
tagandop (Saraca declinata (Jack) Miq.) 165
tahid (Koompassia excelsa (Becc.) Taub.) 154
takalot (Lithocarpus jacobsii Soepadmo) 64
talatad (Ficus macilenta King) 284
talolonandim (Intsia bijuga (Colebr.) Kuntze) 150
tamaran (Artocarpus tamaran Becc.) 210, 211
tambakis (Saraca declinata (Jack) Miq.) 165
tampang (Prainea limpato (Miq.) Beumee ex Heyne) 329
tampar hantu (Sindora leiocarpa Backer ex de Wit) 172
tampar hantu paya (Sindora leiocarpa Backer ex de Wit) 172
tanggajun (Parartocarpus venenosus (Zoll. & Moritzi) Becc.) 323
tanggar-angor (Crudia reticulata Merr.) 136
tanid (Koompassia excelsa (Becc.) Taub.) 154
tanyit (Koompassia excelsa (Becc.) Taub.) 154
tapang (Koompassia Maingay ex Benth.) 153
tapang (Koompassia excelsa (Becc.) Taub.) 154
tapian diwit (Ficus minahassae (Teijsm. & de Vries) Miq.) 290
tawak (Artocarpus anisophyllus Miq.) 192
tegong (Ficus brunneoaurata Corner) 264
tekalat (Lithocarpus brochidodromus S. Julia & Soepadmo) 43
tekalat (Lithocarpus gracilis (Korth.) Soepadmo) 60
tekalat uak (Lithocarpus gracilis (Korth.) Soepadmo) 60
tekaled (Lithocarpus andersonii Soepadmo) 37
tekalet kapuong (Lithocarpus porcatus Soepadmo) 79
tekalong (Artocarpus elasticus Reinw. ex Blume) 197
tekalong (Artocarpus odoratissimus Blanco) 205
tekulud (Lithocarpus Blume) 30
tembaran (Artocarpus tamaran Becc.) 211
temedak (Artocarpus integer (Thunb.) Merr.) 199
tempan (Ficus aurata (Miq.) Miq.) 259
tempan (Ficus bruneiensis Corner) 263
tempan (Ficus brunneoaurata Corner) 264
tempan (Ficus eumorpha Corner) 274
tempan (Ficus fulva Reinw. ex Blume) 277
510
tempan (Ficus lamponga Miq.) 282

tempan (Ficus macilenta King) 284
tempan (Ficus ruficaulis Merr.) 301
tempan (Ficus subfulva Corner) 307
tempinis (Streblus Lour.) 329
tempinis (Streblus elongatus (Miq.) Corner) 331
temponek (Artocarpus melinoxylus Gagnep. subsp. brevipedunculatus Jarrett)
temponek (Artocarpus rigidus Blume) 208
tepih (Copaifera palustris (Symington) de Wit) 129
terampayoh (Castanopsis oviformis Soepadmo) 27
terap (Artocarpus J.R. & G. Forster) 187, 188
terap (Artocarpus elasticus Reinw. ex Blume) 197
terap (Artocarpus glaucus Blume) 198
terap (Artocarpus kemando Miq.) 201
terap (Artocarpus nitidus Trc.) 204
terap (Artocarpus odoratissimus Blanco) 205
terap (Artocarpus tamaran Becc.) 211
terap (Artocarpus teysmannii Miq.) 212
terap (Parartocarpus Baill.) 318
terap (Parartocarpus bracteatus (King ex Hook f.) Becc.) 320
terap ikal (Artocarpus anisophyllus Miq.) 192
terap togop (Artocarpus elasticus Reinw. ex Blume) 197
terbudak (Ficus sinuata Thunb.) 305
tian (Artocarpus elasticus Reinw. ex Blume) 197
tibadak (Artocarpus integer (Thunb.) Merr.) 199
tibudak (Artocarpus integer (Thunb.) Merr.) 199
tihaig (Lithocarpus elegans (Blume) Hatus. ex. Soepadmo) 56
tikalod (Lithocarpus brochidodromus S. Julia & Soepadmo) 43
tikalod (Lithocarpus cantleyanus (King ex Hook. f.) Rehder) 44
tikalod (Lithocarpus conocarpus (Oudem.) Rehder) 49
tikalod (Lithocarpus gracilis (Korth.) Soepadmo) 60
tikalod (Lithocarpus leptogyne (Korth.) Soepadmo) 70
timadang (Artocarpus odoratissimus Blanco) 205
timbangan (Artocarpus tamaran Becc.) 211
timbarayong (Peltophorum racemosum Merr.) 162
timopada (Ficus geocharis Corner) 277
tukalod (Castanopsis psilophylla Soepadmo) 30
tululs hobui (Saraca declinata (Jack) Miq.) 165
turug (Lithocarpus leptogyne (Korth.) Soepadmo) 70
tusuk pending memaluh (Lithocarpus jacobsii Soepadmo) 64
U
umpo (Horsfieldia Willd.) 352
uok (Ficus septica Burm. f.) 304
upang (Horsfieldia Willd.) 352
usu (Prainea limpato (Miq.) Beumee ex Heyne)
329
W
waringin (Ficus benjamina L.) 262
wau (Castanopsis hypophoenicea (Seemen) Soepadmo)
511
19
203
vi

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TREE FLORA

Produced with the financial support of

INTERNATIONAL TROPICAL TIMBER ORGANIZATION

OVERSEAS DEVELOPMENT ADMINISTRATION, U.K.

Tree Flora of Sabah and Sarawak Editorial Committee Chairman: E. Soepadmo

E. Soepadmo and K.M. Wong

Copyright Forest Research Institute Malaysia (FRIM),

First published 1995

Perpustakaan Negara Malaysia Cataloguing-in-Publication Data

Electronic version by L. G. Saw & B. Aslina, Forest Research Institute

THE TREE FLORA PROPER

Abbreviations of Frequently Cited References

significance to the continuing and future undertaking of conservation and forestry

Dato Dr. Salleh Mohd. Nor

A Brief History of Botanical Collecting and Documentation in Borneo

Biogeography and Ecology

TREE FLORA OF SABAH AND SARAWAK

BACKGROUND TO THE TREE FLORA

Why a Tree Flora of Sabah and Sarawak?

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)

The need for a Flora of Borneo

he flora of a given region provides an inventory of plant species occurring in that

The Tree Flora of Sabah and Sarawak Project

To publish 8 volumes (each containing 300400 species) of a concise Tree Flora of

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)

NomenclatureNames of families and genera will normally follow Brummitt (1992),

ReferencesOnly selected references most relevant to the taxonomy and distribution

Derivation of genus and species namesLanguage and meaning of root words to be

Basionym and synonymBasionym to be given if it indicates a new taxonomic

TypificationWhenever possible, citation of type specimen(s) of accepted species

Vernacular namesOnly names commonly used in Sabah and Sarawak will be

EcologyFor a family or a genus to be given only if general trends can be

UsesFor a family or a genus to be provided only if there are general or interesting

TaxonomyTo be provided only if there are taxonomic controversies regarding the

Key(s) to lower rank taxaBracket key structure is to be used. Couplets to be

Other project activities

Organisation of collecting expeditions to a number of botanically little-known localities

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)

TREE FLORA OF SABAH AND SARAWAK

A BRIEF HISTORY OF BOTANICAL

uch of the information on early botanical collections in Borneo is summarised, by

Early collectors and documentation

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)

BRIEF HISTORY (WONG)

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)

Early 20th Century collectors

BRIEF HISTORY (WONG)

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)

BRIEF HISTORY (WONG)

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)

BRIEF HISTORY (WONG)

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)

BRIEF HISTORY (WONG)

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)

BRIEF HISTORY (WONG)

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)

Of check-lists and floras

BRIEF HISTORY (WONG)

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)

BRIEF HISTORY (WONG)

TREE FLORA OF SABAH AND SARAWAK VOL. 1 (1995)

BRIEF HISTORY (WONG)