Content uploaded by Lena Struwe
Author content
All content in this area was uploaded by Lena Struwe on Feb 14, 2016
Content may be subject to copyright.
CSIRO PUBLISHING
© CSIRO 2004 10.1071/SB03021 1030-1887/04/040399
www.publish.csiro.au/journals/asb Australian Systematic Botany 17, 399–406
Neuburgia novocaledonica, comb. nov. and the first record of domatia
in the family Loganiaceae
Jeanmaire E. Molina
A,B
and Lena Struwe
A
A
Department of Ecology, Evolution and Natural Resources, 237 Foran Hall, 59 Dudley Road, Rutgers University,
New Brunswick, New Jersey 08901, USA.
B
Corresponding author; email: jeanx@eden.rutgers.edu
Abstract. The New Caledonian species Couthovia novocaledonica Gilg & Benedict (Loganiceae) is supported as
a species distinct from Neuburgia corynocarpa (A.Gray) Leenh. Neuburgia novocaledonica is different in having
domatia on its abaxial leaf surfaces, two rings of hairs within the corolla tube, and generally obovate leaf shape.
Therefore, the new combination Neuburgia novocaledonica (Gilg & Benedict) J. Molina & Struwe is hereby made.
The presence and morphology of domatia on the lower surface of the leaves of Neuburgia novocaledonica are
discussed. This appears to be the first record of domatium occurrence in Loganiaceae.
SB03021
The first record of domatia in the f amily Logani aceaeJ. E. Mol ina a nd L. Struwe
Introduction
Neuburgia, described by Blume (1850), is a genus of woody
understory trees or shrubs often found in the everwet lowland
tropical forests of the Philippines, Celebes, Caroline Islands,
Papuasia, New Hebrides, New Caledonia and Fiji. There are
approximately 10–12 species in Neuburgia according to
Leenhouts (1963) and Conn (1995).
Couthovia was described as a new genus by the
taxonomist Gray (1858) and placed in the family
Loganiaceae. Leenhouts (1963) found Couthovia to be
identical with Blume’s Neuburgia (1850), and validly
included Couthovia in Neuburgia. The International Plant
Names Index (1999) lists 28 published species for
Couthovia, nine of which are already moved to and accepted
in Neuburgia, and 18 species reduced to synonymy
(Leenhouts 1963). In the same publication, Leenhouts
recognised six species formerly placed in Couthovia.
Couthovia novocaledonica was described by Gilg and
Benedict (1921) from New Caledonia and has often been
misspelled as C. ‘neo-caledonica’. Leenhouts (1963)
speculated that C. novocaledonica Gilg & Benedict may be
‘possibly combined with Neuburgia corynocarpa (A.Gray)
Leenhouts’ (Leenhouts 1963 p. 366). Leenhouts (1963)
reduced 13 names of Couthovia to a synonym of
N. corynocarpa. Owing to lack of material, Leenhouts was
unable to confirm the taxonomic status of
C. novocaledonica. Our recent discovery of domatia on the
lower surface of the leaves of this species strongly supports
its distinctiveness from the domatia-less N. corynocarpa.
Differences in three additional characters between the two
species are also presented (Figs 1, 2; Table 1). Couthovia
novocaledonica is therefore distinct from N. corynocarpa,
and the new combination, Neuburgia novocaledonica, is
here proposed.
Although Brouwer and Clifford (1990) published one
recorded example of domatia for the family Loganiaceae,
namely Coinochlamys angolana var. laurentii de Wild, this
species has been now reduced to the synonym of Mostuea
hirsuta (T.Anders.ex Benth. & Hook.f.) Baill ex Bak. of the
segregate family, Gelsemiaceae (Leeuwenberg 1961; Struwe
et al. 1994). Our study appears to be the first documentation
of domatia in the genus Neuburgia and also, the first
published record for true Loganiaceae.
Materials and methods
Herbarium specimens for Neuburgia novocaledonica and Neuburgia
corynocarpa (see Appendix) from Arnold Arboretum, Harvard
University (A), New York Botanical Garden (NY), United States
National Herbarium (USA), Field Museum of Natural History (F),
Chicago Academy of Sciences (CACS), and Nationaal Herbarium
Nederland, Leiden University branch (L) were examined. Type material
of N. corynocarpa at NY and C. novocaledonica at L were also
examined. For detailed studies, samples of pressed flowers, fruit and
leaves were rehydrated by boiling in water for 30–60 s in a microwave.
Structures were observed with Zeiss Stemi 2000-C (Edmund Optics
Inc., Barrington, NJ) or Olympus SZ (Olympus America Inc., Melville,
NY) dissecting microscopes. Photographs were taken with Nikon
DXM1200 digital camera with ACT-1 version 2.11 software.
Taxonomic treatment
Neuburgia novocaledonica (Gilg & Bened.) J. Molina &
Struwe, comb. nov. Basionym: Couthovia novocaledonica
Gilg & Bened., Bot. Jahrb. 56: 547. 1921. Lectotype: NEW
CALEDONIA. Nordbezirk auf den Bergen bei Ou-Hinna,
[altitude] 500 m, 1903, Schlechter 15650 (L) (Fig. 1B)
Selected specimens examined
NEW CALEDONIA: am Kugel om Poindelen ob Kene auf Schiefer,
Däniker 1068, 1.ii.1925, L. Col d’ Amieu, Sarramea, Webster &
Hildreth 14896, 29.iv.1968, A. Col d’ Amieu, McKee 8065,
400 Australian Systematic Botany J. E. Molina and L. Struwe
A
Fig. 1. Predominant leaf shape of mature leaves (A) N. corynocarpa, broadly elliptic or oval (type, NY). (B) N. novocaledonica
(type, L), obovate.
The first record of domatia in the family Loganiaceae Australian Systematic Botany 401
B
Fig. 1. (continued)
402 Australian Systematic Botany J. E. Molina and L. Struwe
13.i.1961, USA. Col d’ Amieu, Guillaumin 8472, 21.xi.1950, A &
NY. Guillaumin 10116, 2.ii.1951, NY. Ferme Modelle, Balansa 360,
NY. L.O. de Canala, Balansa 2003, A. Route de Gomen, Hurlimann
1813, 24.viii.1951, NY & A. Col d’ Petchicara, McKee 6579,
11.ix.1958, 2 sheets, A and USA. Ourai, Lecard 20 Oct. 1879, A.
Vieillard 681, 1861–1867, A. Pancher s.n., 1870, A. Henecart s.n.,
iv.1879, A. UNITED STATES: HAWAII: (cultivated ex New
Caledonia) Kauai, Koloa Distr., Lawai Valley: National Tropical
Botanical Garden, Maidenhair Falls section, Lorence 7370,
29.iv.1993, F.
Shrub or tree up to 20 m high, glabrous throughout
except flowers; branches slightly quadrangular or terete,
solid; bark smooth or fissured. Leaves opposite, simple,
petiolate, narrow to broadly obovate, 8–19 × 3–11 cm,
chartaceous to subcoriaceous, margins entire; venation
pinnate, primary and secondary veins raised abaxially and
adaxially, tertiary veins obscure and laxly reticulate;
pit-type domatia in vein axils on abaxial leaf surface,
starting from third to sixth secondary vein from base of
midvein and regularly occurring until leaf apex, first order
[i.e. in the axils of primary nerves (Jacobs 1966)] and
second order; base cuneate; apex rounded to obtuse;
petiole 0.8–1.9 cm long; stipules interpetiolar, 0.5–1.2 ×
0.3–0.9 cm, deltoid, cupuliform, connate with petioles.
Inflorescence a dichasium, dividing three or four times,
terminal; fourth order (most distal relative to florets)
peduncle 8–11 cm, succeeding orders of peduncles
decreasing in length, with the central peduncle always
shortest; bracts broadly ovate, ciliated. Flowers white,
perfect, actinomorphic, hypogynous, terminal, 5-merous,
5.0–6.5 × 3.0–3.9 mm; pedicels 0.5–2.0 mm long, with
punctate glands. Calyx 5-lobed, synsepalous, cupuliform,
coriaceous, imbricate; calyx lobes suborbicular to
orbicular, 1.5–2.7 × 1.5–2.3 mm, apex rounded, margin
entire, ciliolate, lobes densely punctuate, light yellow
when dry. Corolla 5-lobed, sympetalous, salverform,
subcoriaceous, valvate; corolla tube 2.2–5.0 mm long,
point of insertion of stamens barbate; corolla lobes
narrowly ovate, 1.2–4.9 mm long, apex acute, margins
entire, mouth more densely barbate than basal part of
corolla tube; hairs at corolla mouth 0.9–1.8 mm long;
hairs at corolla base 0.4–0.9 mm long. Stamens 5,
alternating with corolla lobes, adnate to corolla tube,
inserted at 1/4 of tube length from corolla base, introrse;
filaments 1.0–2.6 mm long; anthers narrowly ovate,
2.0—3.5 × 1 mm, longitudinally dehiscent. Gynoecium
obclavate, c. 6.6 mm long; ovary ovoid, 2-seeded, one seed
aborted, placentation axile; style 2.0–2.6 × 0.5 mm,
gradually confluent with ovary; stigma c. 0.6–0.7 ×
0.6–0.7 mm, ovoid. Fruit a drupe, white, clavate, 2.0–2.2 ×
0.9–1.1 cm, indehiscent; exocarp with fine longitudinal
ridges; mesocarp very fibrous.
Results
Neuburgia corynocarpa and N. novocaledonica primarily
differ in three morphological characters, namely leaf shape,
position of hairs in corolla and domatium occurrence, all of
which were studied and analysed in this paper.
Neuburgia corynocarpa leaves are ovate or broadly
elliptic (Fig. 1A), whereas N. novocaledonica leaves are
obovate, with the widest part conspicuously displaced
towards the apex (Fig. 1B). Furthermore, the leaves have
subacute apices in the type of N. corynocarpa (Fig. 1A),
whereas leaves of N. novocaledonica are obtuse (Fig. 1B).
Fig. 2. Position of hairs on inside of corolla (indicated by arrow)
(A) N. corynocarpa (AC Smith 7133, NY), hairs only in mouth of
corolla. (B) N. novocaledonica (Guillaumin 10116, NY), hairs both in
corolla mouth (M) and base (T) of tube (1, stamen). Bars = 1 mm
(A, B).
The first record of domatia in the family Loganiaceae Australian Systematic Botany 403
In Neuburgia corynocarpa, hairs are inserted only at the
entrance to the corolla mouth (Fig. 2A), unlike
N. novocaledonica, where hairs are inserted in the corolla
both at the base of the stamens and at the mouth (Fig. 2B).
Domatia are absent in the vein axils of the abaxial surface
of the leaves of Neuburgia corynocarpa (Fig. 3A), but are
persistently present in the first (Fig. 3B) and second (Fig. 3C)
order vein axils of N. novocaledonica. N. novocaledonica
has pit-type domatia (Fig. 3B, C).
Discussion
Domatia (Lundstroem 1887) refer to plant structures that
may provide shelter to animals. However, Jacobs (1965,
1966) restricted the usage of the term to spatial structures
occurring in or near vein axils of the first, second or third
order, and were restricted to the abaxial surface of leaves. He
further confined the term to features that accommodate only
mite symbionts, hence excluding ants and bacteria (Jacobs
1966). Domatia are plant-produced formations and are not
induced by their resident symbionts (O’Dowd and Willson
1991). They appear early in plant development (Brouwer and
Clifford 1990), but may not be inhabited until maturity
(Willson 1991). Interestingly, domatia are also restricted to
woody dicotyledons growing in humid habitats (Jacobs
1965).
The ecological significance of domatia has long been the
subject of debate, but recent studies demonstrate the
beneficial role of domatia in promoting plant–arthropod
mutualism (Pemberton and Turner 1989; O’Dowd and
Pemberton 1994; Agrawal and Karban 1997). Domatia
provide protective domiciles and nurseries for mites, and
additionally, these mites keep the leaf lamina healthy by
feeding on pathogenic fungi and herbivorous mites (O’Dowd
and Pemberton 1994).
Domatia have been recorded in 277 families and in almost
2000 species (Agrawal and Karban 1997) in many
geographical regions (O’Dowd and Pemberton 1994). Since
domatia are heritable structures, Brouwer and Clifford
(1990) considered domatium occurrence as a reliable
taxonomic character. Robbrecht (1988) suggested that
domatia may only be taxonomically useful in species
discrimination, but also commented that higher taxa
(subgenera, genera) may sometimes exhibit a particular
domatium type. In Jacobs’ comprehensive discussion of
domatia, he noted that there are studies supporting the
usefulness of domatia in species delineation, in much the
same way as hairs and glands do (Jacobs 1966). van Steenis
(1967, p. 1568) also affirmed its ‘systematical value in
specified cases.’ Hence, domatium occurrence on the abaxial
leaf surfaces of Neuburgia novocaledonica is a satisfactory
discriminatory character at the species level. Domatia have
never been observed by us in N. corynocarpa, N. tubiflora,
N. rumphiana, N. macrocarpa, N. kochii, N. macroloba, and
N. moluccana (see Appendix).
Without careful inspection, it is difficult to distinguish
between Neuburgia corynocarpa and Neuburgia
novocaledonica, and the latter was treated as a synonym of
the former by Leenhouts (1963). However, the presence of
domatia shows the distinctiveness of N. novocaledonica
(Table 1). Two additional traits are presented here, which may
be supplementary features that can aid in species delimitation
and determination of N. novocaledonica (Figs 1–3).
Neuburgia corynocarpa has broadly elliptic or oval leaves and
occurs at an altitudinal range of 30–200 m. Smith (1942)
claimed that occurrence above 500 m is unusual. Neuburgia
novocaledonica has obovate leaves and occurs at a higher
elevation of 300–650 m. Leaf morphology and altitude of
occurrence are possibly highly variable characters in
N. corynocarpa. Further investigations of more herbarium
material, as well as field studies, are required to see if these
are consistent throughout the broad distributional range of
N. corynocarpa. In N. novocaledonica, the development of
hairs in both corolla base and mouth (Fig. 2B) also
differentiates it from most individuals of N. corynocarpa,
which has only hairs at the corolla mouth. Documented
exceptions (Conn 1979) in N. corynocarpa warrant further
examination. In addition, Conn (1981) had described the
lectotype of N. corynocarpa var. sarcantha as being ‘laxly
wooly in tube, at and above point of insertion of stamens’. We
have only seen three specimens of N. corynocarpa (White
10299, Streismann 44393, Streismann & Lelean 18332) that
are sparsely pubescent at the base of the inner corolla tube
Table 1. Character differences between N. corynocarpa and N. novocaledonica based on protologues, personal observations and
herbarium sheet notes
Character Protologue Personal observations
N. corynocarpa N. novocaldeonica N. corynocarpa N. novocaldeonica
Leaf shape Suborbicular or broadly
elliptic to obovate
(Gillespie 1931)
Obovate or broadly obovate
(Engler 1921)
Oval to slightly ovate, very
rarely obovate;
sometimes orbicular
Obovate; sometimes
oval
Position of hairs on corolla Corolla throat (Gillespie
1931)
Corolla mouth (Engler
1921)
Inner surface of corolla
lobes only
Inner surface of corolla
lobes and base of
corolla tube
Domatia Not mentioned Not mentioned Absent Present
Altitude of aoccurrence Not mentioned 500 m (Engler 1921) 30–200 m 300–650 m
404 Australian Systematic Botany J. E. Molina and L. Struwe
relative to the corolla mouth. These three were all collected
in Papua New Guinea. Hairs at the corolla base are also
distinctly shorter (0.15–0.21 mm) than those in the corolla
mouth (0.33–0.76 mm). This observation may suggest that
position of pubescence within the corolla tube is not a good
species-delimiting feature. However, we also suspect that
N. corynocarpa, in its current circumscription, is a composite
of several species. For example, Conn (1979) observed
variants of N. corynocarpa that possessed only four calyx
lobes, which we have not seen so far in N. novocaledonica.
The taxonomic significance of the occurrence of domatia in
Neuburgia novocaledonica should not be ignored (Fig. 3).
The pit is the predominant domatium type in all examined
specimens of N. novocaledonica (Fig. 3B, C). Intermediates
between the types are likely, hence it is not the type, but it is
the presence or lack of domatia, which must be considered
(Jacobs 1965). These axillary structures or domatia in
N. novocaledonica could not be mistaken for galls, as they are
very regular in their position (Jacobs 1966), being regularly
present in the first and second order vein axils of the leaf’s
abaxial surface. Moreover, of 15 herbarium sheets of
N. novocaledonica examined, all exhibited domatia, except
for one specimen (Balansa 360, NY), which was difficult to
ascertain since a significant portion of the abaxial surface of
Fig. 3. Presence of domatia in vein axils (indicated by arrow) (A) N. corynocarpa (AC Smith 6646, HUH), none in first (1) and second (2) order
axils. (B, C) N. novocaledonica (Guillaumin 8472, NY), pit-type domatia present in both first (1) and second (2) order axils. Bars = 7 mm
(A), 2 mm (B), 3 mm (C).
The first record of domatia in the family Loganiaceae Australian Systematic Botany 405
the leaves were glued to the sheet. The occurrence of domatia
in N. novocaledonica is indisputably a taxonomic character
that delineates it from N. corynocarpa. Therefore,
N. novocaledonica is here recognised a distinct species, and
the transfer from Couthovia to Neuburgia is proposed here.
This may be the first record of the occurrence of domatia
for the genus Neuburgia, and perhaps for the family
Loganiaceae. How this species uniquely evolved domatia
within an angiosperm family that is not known to produce
domatia has interesting implications. Loganiaceae is closely
related with Rubiaceae (Backlund et al. 2000), and both
belong to the order Gentianales, along with three other
families, Gelsemiaceae, Gentianaceae, and Apocynaceae
(APG 1998). Robbrecht (1988) suggested that Rubiaceae has
the highest ratio of occurrence of domatia among all plant
families. The close relationship between Loganiaceae and
Rubiaceae may explain domatium occurrence in the former.
Further studies must be undertaken with respect to the
evolution and ecology of these enigmatic leaf formations.
Acknowledgments
We thank the staff of the New York Botanical Garden, Field
Museum of Natural History, United States National
Herbarium, Nationaal Herbarium Nederland, Leiden
University (Gerard Thijsse), and the Arnold Arboretum of
Harvard University (Gustavo Romero, Emily Wood and Judy
Warnement) for providing us important references and vital
herbarium material. Barbara Zilinskas of Rutgers University
had let us borrow a high-performance dissecting microscope
with a digital camera that was necessary for this project. We
also thank Barry Conn of the Royal Botanic Garden, P. Morat
of the Muséum National d'Histoire Naturelle of Paris,
France, Tran Cong Khanh of the Hanoi College of Pharmacy,
and Leonard Co of the University the Philippines for
responding to our emails regarding domatium occurrence in
Loganiaceae. We finally thank our colleagues and friends,
Cindy Frasier, Juanita Choo, as well as Sandra Yap and Maya
Medalla who were helpful and supportive throughout the
duration of this research. This research and the resulting
paper were developed during a graduate course and we thank
all students of the course for their constructive criticism as
peer-reviewers.
References
Agrawal A, Karban R (1997) Domatia mediate plant–arthropod
mutualism. Nature 387, 562–563. doi:10.1038/42384
Angiosperm Phylogeny Group (1998) An ordinal classification for the
families of flowering plants. Annals of the Missouri Botanical
Garden 85, 531–553.
Backlund M, Oxelman B, Bremer B (2000) Phylogenetic relationships
within the Gentianales based on ndhF and rbcL sequences, with
particular reference to theLoganiaceae. American Journal of Botany
87, 1029–1043.
Blume CL (1850) Neuburgia. Museum Botanicum Lugduno Batavum 1,
156–157.
Brouwer YM, Clifford HT (1990) An annotated list of domatia-bearing
species. Notes of the Jodrell Laboratory 12, 1–33.
Conn BJ (1979) Notes on Neuburgia Blume (Loganiaceae) in Papuasia.
Brunonia 2, 99–105.
Conn BJ (1981) Lectotypification of Neuburgia corynocarpa var.
sarcantha (Loganiaceae). Brunonia 4, 209–211.
Conn BJ (1995) Neuburgia. In ‘Handbooks of the flora of Papua New
Guinea. Vol. 3’. (Ed. BJ Conn) pp. 174–182. (Melbourne University
Press: Australia)
Engler A (1921) Couthovia novocaledonica Gilg & Bened. Botanische
Jahrbücher fürSystematik 56, 547–548.
Gillespie JW (1931) Couthovia corynocarpa Gray. Bernice P. Bishop
Museum Bulletin 83, 29.
Gray A (1858) Couthovia nov. gen. Proceedings of the American
Academy of Arts and Sciences 4, 324.
International Plant Names Index (1999) http://www.ipni.org [Online
verified 25 June 2004]
Jacobs M (1965) Domatia. Flora Malesiana Bulletin 20, 1272–1273.
Jacobs M (1966) On domatia—the viewpoints and some facts.
Proceedings of theKoninklijke Nederlandse Akademie van
Wetenschappen (Section C) 69, 275–316.
Leenhouts PW (1963) Neuburgia. Flora Malesiana (Series 1) 6,
363–369.
Leeuwenberg AJM (1961) The Loganiaceae of Africa: a revision of
Mostuea Didr. Mededeelingen van de Landbouwhogeschool 61,
1–31.
Lundstroem AN (1887) Pflanznbiolische Studien II. Die Anpassungen
der Pflanzen an Thiere. Nova Acta Regiae Societatis Scientiarum
Upsaliensis (Series 3) 2, 1–74.
O’Dowd DJ, Pemberton RW (1994) Leaf domatia in Korean plants:
floristics, frequencyand biogeography. Vegetatio 114, 137–148.
O’Dowd D, Willson M (1991) Associations between mites and leaf
domatia. Trends in Ecology and Evolution 6, 179–182.
Pemberton R, Turner C (1989) Occurrence of predatory and
fungivorous mites in leaf domatia. American Journal of Botany 76,
105–112.
Robbrecht E (1988) Tropical woody Rubiaceae. Opera Botanica
Belgica 1, 49–50.
Smith AC (1942) Couthovia. Sargentia 1, 99–107.
Struwe L, Albert VA, Bremer B (1994) Cladistics and family level
classification of the Gentianales. Cladistics 10, 175–206.
doi:10.1006/CLAD.1994.1011
van Steenis C (1967) Occurrence of domatia as a systematic character.
Flora Malesiana Bulletin 22, 1568–1570.
Willson M (1991) Foliar shelters for mites in the Eastern deciduous
forest. American Midland Naturalist 126, 111–117.
Manuscript received 11 August 2003, accepted 22 June 2004
406 Australian Systematic Botany J. E. Molina and L. Struwe
http://www.publish.csiro.au/journals/asb
Appendix
Specimens examined for Neuburgia corynocarpa (A.Gray) Leenh.
ISOTYPE: FIJI. Wilkes Expedition (NY).
FIJI: (1) Namosi: Veivatuloa, Rova & Gutavsson 2433, 11.ii.1999, NY. (2) Vanua Levu: Mathuata, Smith 6646, 25.xi.–8.xii.1947, NY. (3) Vanua
Levu: Mathuata: Southern slopes of Mt Numbuiloa, east of Lambasa, Smith 6330, 27.x.–10.ii.1947, A (4) Viti Levu: Namosi: Hills between Navua
R. and Suva, Greenwood 1056, v.1943, A. (5) Viti Levu: Namosi, 28.ix.1953, Smith 8750, NY. (6) Viti Levu: Namosi: Hills bordering Wainavindrau
Creek, in vicinity of Wainimakutu, Smith 8520, 17.ix.–8 Oct. 1953, USA. (7) Wailoku, 8 km NW of Suva, Regalado and Vodonaivalu 535,
12.iv.1993, CACS. (8) Serua, Smith 8991, 20.x.1953, NY. (9) Serua, Smith 9308, 23.xi.–7.xii.1953, NY. (10) Tailevu, Smith 7133, 15–27.iv.1953,
NY.
HAWAII: (11) Oahu Isalnd, HSPA Forest Nursery, Manoa Valley, Yuncker 3598, 10 Apr. 1933, CACS.
INDONESIA: (12) Bogor: Island Buru, West Buru, Bara, Wae Duna, base camp 9, Mogea 5431, 30 Nov. 1984, A.
PAPUA NEW GUINEA: (13) East New Britain District: Kokopo Subdistrict: Trans Vudal, Streimann 44393, 18.viii.1969, A. (14) Madang District:
Josephstaal, White 10299, ix.1958, A. (15) Morobe Province: logging road near Markham Village, 6°43' S, 146°53' E, Takeuchi 7144, 10.vii.1991,
NY. (16) Western District: Kiunga Subdistrict: Tuidemasuk Road, Kiunga, Streimann & Lelean 18332, 17.ix.1972, A.
Specimens examined for other species of Neuburgia
(1) Neuburgia macroloba (A.C. Sm.) A.C. Sm.
ISOTYPE: FIJI: Borders of lake east of Somosomo, Smith 917, 29.xii.1933, L.
(2) Neuburgia moluccana (Boerl.) Leenh.
ISOLECTOTYPE: Unknown. Cult. In Hort. Bot. Bog. Java, Rusdy 3, 3.iv.1960, L.
ISOLECTOTYPE: INDONESIA: Moluccas, Unknown IV.E.92, 4.vi.1912, L.
SYNTYPE: INDONESIA: Moluccas, Unknown s.n., N.A., L.
(3) Neuburgia rumphiana Leenh.
HOLOTYPE: INDONESIA: New Guinea: Holtekang, E. of Hollandia, Lam 7787, 12.x.1954, L.
(4) Neuburgia sarcantha (Gilg & Benedek) Leenh.
HOLOTYPE: PAPUA NEW GUINEA: New Guinea Western Highlands District Terr. of New Guinea W. of Togoba, Hoogland & Pullen 6207,
9.xii.1956, L.
(5) Neuburgia tubiflora Blume
ISOLECTOTYPE: INDONESIA: New Guinea, Markgraf s.n., 1926, L.
