Global Flora Vol 4

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The Global Flora ISSN 2398-6336 eISSN 2398-6344

A practical flora to vascular plant species of the world
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i Published online 9 February 2018

Plant Gateway’s
THE GLOBAL FLORA

GLOVAP Nomenclature Part 1
February 2018
The Global Flora
Special Edition, GLOVAP Nomenclature Part 1, Vol. 4: 1-155.
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Cover image: Hakea plurijuga (F.Muell.) Christenh. & Byng © Maarten Christenhusz
Editors
Maarten J.M. Christenhusz

Plant Gateway, Bradford & Kingston, United Kingdom and Den Haag, the Netherlands
Royal Botanic Gardens, Kew, United Kingdom.
m.christenhusz@kew.org
Michael F. Fay
Royal Botanic Gardens, Kew, United Kingdom.
m.fay@kew.org
James W. Byng
Plant Gateway, Bradford & Kingston, United Kingdom and Den Haag, the Netherlands
Naturalis Biodiversity Center, Leiden, the Netherlands.
james.byng@naturalis.nl
With contributions by
Ehoarn Bidault (Burmanniaceae)

Missouri Botanical Garden, St. Louis, United States of America.
Muséum national d’Histoire naturelle & Sorbonne Universités, Paris, France.
John E. Burrows (Myrtaceae)

Buffelskloof Nature Reserve, Lydenburg, South Africa.
Florian Jabbour (Ranunculaceae)

Muséum national d’Histoire naturelle & Sorbonne Universités, Paris, France.
Vincent S.F.T. Merckx (Burmanniaceae, Triuridaceae)

Naturalis Biodiversity Center, Leiden, the Netherlands.
Leiden University, Leiden, the Netherlands.
Greg Stull (Icacinaceae)

University of Michigan, Ann Arbor, United States of America.
Abstract
A Global List of Vascular Plants (GLOVAP) has been compiled after consulting a large number of published
phylogenetic studies and taxonomic literature. We update the classifications for a number of vascular plant families
and genera, by expanding or splitting, to provide monophyletic and morphologically well-defined groups. The first
nomenclatural changes needed to do this are presented here. We provide two new genera, four new species, 415
new names and 3286 new combinations. GLOVAP vs. 1 will adopt these new changes and hence make this one of
the largest taxonomic undertakings in botany of recent times.
Keywords
Classification – Genera – GLOVAP – Vascular plants
Background any of the standard databases cited above and hence

Generic-level classifications of many vascular plant we may have overlooked these. We apologise for these
families are still poorly known and the classification discrepancies and unintentional superfluous names,
is often not revised as new evidence comes to light. and we shall correct errors in future updates. We aim for
This is particularly true of online databases (e.g. longer term stability and it is therefore inevitable that
IPNI, Jstor, Tropicos, The Plant List, Catalogue of Life, some new names had to be coined for familiar plants
GBIF) that are usually consulted to provide a global that were nomenclaturally problematic when placed in
classification for cosmopolitan families and to identify a proper evolutionary framework. Thus, some familiar
the genus and family in which a species now should names will disappear due to these nomenclatural
be placed. Each of these online resources comes with corrections. It is like changing your surname when you
its own idiosyncrasies and errors, often resulting in have established who your parents are; people get used
incompatibility and conflicting results. Rather than to new names in the long run, particularly when they
being complementary, the information in these are unlikely to change again.
databases is often contradictory, and for the user of We cite references for each family that support these
taxonomy it can be a daunting task to know what changes and most changes are self-explanatory and
genus to choose. This is potentially leading to an error- we provide a short note for each case if appropriate.
strewn global taxonomy that is confusing and difficult to It generally results into an avoidance of naming
apply for users. A classification only has purpose if it is each clade in a grade as a separate genus, but taking
practical and useable, whilst still reflecting evolutionary historical usage into account and assuring that the
patterns. Admittedly the existence of these databases future nomenclature will remain as stable as possible
is essential in helping taxonomists and systematists based on the currently available data. Nevertheless, we
find the relevant literature, references and associated highlight a few cases below in which well-established
specimens, but as the sources and data are of varied genera have disappeared. We know that these changes
quality and the sources are often not explicitly stated, are not always positively welcomed, and therefore we
it is impossible for the general user to know which explain our motivation in the following seven cases.
database to use.
Ferns – The classification of ferns has been
We have compiled an accepted and synonymised contentious in recent times as there is a basic
linear sequence of all vascular plant genera called the discrepancy between the generalist view and the
Global List of Vascular Plants (GLOVAP), which will be specialist view. The first, supported by the classification
published soon in The Global Flora (Byng & Christenhusz, by Christenhusz & Chase (2014), opts for a broader
in prep.) After consulting hundreds of phylogenetic concept of genera, in line with historical treatments
studies and herbarium specimens, we update the of genera and avoiding taxonomic uncertainty. This
classification of many genera that have been shown to classification absorbs sister lineages into broader
be polyphyletic and hence need to be either expanded genera, avoiding intergeneric hybrids as much as
or split depending on the history of the group in order possible and focuses on the similarities rather than
to create the greatest nomenclatural stability for the differences. The latter, more finely split classification
foreseeable future. In this volume we make some of the was published by the PPG (2016), a consortium of
nomenclatural changes required and we have aimed to fern specialists. Even though this was presented as a
provide complete generic transfers whenever possible. community effort in line with APG IV (2016), the APG
Inevitably we will have omitted some combinations, method was not employed, and it resulted in the
but this is not intentional. It is also possible that new opposite, a highly divided classification with numerous
combinations already existed but were not included in genera that are difficult to identify. This results in
Published online 9 February 2018

1
genera that can only be established on molecular Proteaceae subtribe Hakeinae (Mast et al., 2015)
results after the species has been identified, and thus has shown that the large genus Grevillea R.Br. forms
the rank of genus serves little purpose. We have thus a grade leading to a monophyletic Hakea Schrad. &
followed the classification of Christenhusz & Chase J.C.Wendl., with Finschia Warb. also embedded in
(2014) and provide combinations for species to comply Grevillea. Grevillea is a well-established name, but
to the traditional broad concepts of Athyrium Roth., lacks an unambiguous synapomorphy in support of
Blechnum L., Grammitis Sw., Selliguea Bory, Thelypteris its monophyly. Nomenclatural changes have not been
Adans. and Trichomanes L. The case of Hemionitis L. is made, and even though we recognise that it will take
somewhat more contentious. Recent molecular results time before a new name for such a common and popular
on Pteridaceae subfamily Cheilanthoideae (Eiserhard horticultural genus like Grevillea will be adopted, we
et al. 2011) have shown that formerly well-established think it necessary to at least make the names available
genera like Cheilanthes Sw., Doryopteris J.Sm. and for use.
Pellaea Link, among others, are polyphyletic. There has
Rosaceae – Relationships between the genera in
been a suggestion to split all three genera, resulting in
subtribe Malinae are problematic, notably because
at least 15 poorly circumscribed genera, several only
Sorbus L. sensu lato is polyphyletic. Phylogenetic
recently described or elevated from synonymy. The
analyses have generally not provided clear resolution
result is that few botanists can recognise the genera in
between the genera in this clade and there are many
the field. Instead, we suggest that the entire subfamily
natural and synthetic intergeneric hybrids, especially if
should be merged into a single genus, as all the taxa
the lineages of Sorbus L. sensu lato are considered as
are recently evolved and hybrids across the clade are
separate genera. Sorbus intermedia (Ehrh.) Pers., for
known. As Hemionitis has nomenclatural priority, this
example, is a stabilised hybridogenic species, with S.
name is used here.
aucuparia L., S. aria (L.) Crantz (=Aria nivea Host.) and
Asparagaceae – As currently circumscribed S. torminalis (L.) Crantz (=Torminalis clusii (Roemer)
Dracaena L. (Asparagaceae) is a paraphyletic genus K.R.Robertson & J.B.Phipps) in its background.
with Sansevieria Thunb. and part of Pleomele Salisb. Thus, if the segregates of Sorbus sensu lato were to
embedded in it (Rudall et al., 2000; Lu & Morden, 2014). be recognised, this taxon would need to be called
Several authors have placed Sansevieria in synonomy ×Tormariosorbus intermedia (Ehrh.) J.M.H.Shaw,
with Dracaena based on the recent molecular with ×Tormariosorbus Mezhenskyi being the hybrid
evidence and their similar morphology (e.g. Bos, 1998; genus between Sorbus sensu stricto and ×Torminaria
Mabberley, 2017), but the new combinations required (the hybrid genus between Aria and Torminalis). An
for most of them have still not been made, leading alternative solution was proposed by Sennikov &
to increasing confusion among users. Species of both Kurtto (2017), who recognised the segregates of Sorbus
Dracaena and Sansevieria are popular ornamentals, sensu lato as genera (in the interest of recognising
outdoors in tropical areas and indoors in the temperate monophyletic units), but treated the nothogenera as
zones, which could make any potential nomenclatural genera in their own right (e.g. Borkhausenia Sennikov
changes unpopular in horticulture. In contrast, Pleomele & Kurtto = ×Sorbotorminaria). This latter solution, in
species are not of horticultural importance and new our opinion, further confuses the situation, ignoring
combinations have already been made (Jankalski, the polyphyletic relationships of their parental taxa.
2008; Mabberley, 2017). In addition a new genus was Most recently, Zhang et al. (2017), based on analyses
proposed, Chrysodracon (Jankalski) P.L.Lu & Morden of complete plastid genomes, identified four major
for a Hawaiian clade of Dracaena, but there has been clades in Malinae, two of which (Crataegus L./Mespilus
limited uptake of the latter change (e.g. WCSP, 2018). L./Amelanchier Medik./Malacomeles (Decne.) Decne./
Traditional classifications have favoured two distinct Peraphyllum Nutt. on the one hand and Pyracantha
genera based on gross morphology, primarily degree M.Roem. alone on the other) are clearly distinct from
of succulence and fruit morphology. However, great the remaining taxa. These are not discussed further
morphological variation already exists in both genera, here (although we note that Mespilus is embedded in
and there is significant morphological overlap. Notably, Crataegus and the appropriate combinations exist).
there are similarities in underground organs, stems,
For many of the remaining taxa of Malinae,
inflorescences, flowers, ecology and distribution (Bos,
combinations exist in two or more genera. For example,
1998). Recent molecular results by Lu & Morden (2014)
Sorbus torminalis has been treated as Crataegus
show that Dracaena forms a grade to an apparently
torminalis L., Pyrus torminalis (L.) Ehrh. and Torminalis
monophyletic Sansevieria. In order to retain a distinct
clusii (Roemer) K.R.Robertson & J.B.Phipps. However,
Sansevieria, Dracaena would need to be divided into
the inclusion of two major fruit crops (apples and
nearly 20 genera causing a high level of nomenclatural
pears) and the existence of a large number of apomictic
disruption and confusion.
species, notably in Cotoneaster and Sorbus, in this
Proteaceae – An extensive molecular study of group has led to people being reluctant to make the
2 Published online 9 February 2018

taxonomic changes to reflect a monophyletic group, Taxonomic list

namely Pyrus sensu lato, the generic name with
priority. In our opinion, the best solution is to recognise F5. Ophioglossaceae
Pyrus sensu lato for the remaining taxa and we make Note: The following taxon is best treated at the species
the necessary combinations here. level.
Penaeaceae – Morphologically the genera of Ophioglossum malgassicum (C.Chr.) Christenh., comb.
Penaeoideae are similar, but have traditionally been & stat. nov. Basionym: Cheiroglossa palmata var.
divided into seven genera based on the number of malgassica C.Chr., Dansk Bot. Ark. 7: 185. 1932.
pollen apertures, number and orientation of ovules,
stigma type and calyx aestivation (Dahlgren 1967a, b, c, F7. Marattiaceae. References: Murdock (2008),
1968, 1971). However, molecular data do not support Senterre (2014).
these generic delimitations as Brachysiphon A.Juss. and Note: A few species were not yet transferred to Ptisana
Stylapterus A.Juss. have both been shown to be non- Murdock (2008). African P. fraxinea appears not to
monophyletic, and several of the resulting clades are be a single species (Senterre et al., 2014) and two of
not supported by traditional morphological characters. the known species that do not yet have a name are
The lack of clear-cut boundaries between genera was described below.
noted by the authors (Rourke & McDonald, 1989;
Ptisana africana Christenh., sp. nov. (Figure 1A–B).
Rourke, 1995) of recently described species when
Similar to P. fraxinea Sm., but differs by its distal pairs
attempting to assign them to a particular genus. We
of pinnules that are similar in size and shape to the
advocate expanding the circumscription of Penaea L. to
other pinnues (vs. progressively larger distally in P.
all members of Penaeoideae.
fraxinea), the pinnules with a more or less abruptly
Cleomaceae – Until recently, Cleome L. was caudate acuminate apex (vs. progressively narrowed
considered a single large genus in keeping with floristic or acuminate), a larger rhizome, up to 40 cm tall (vs.
tradition. Since then the generic concept has narrowed up to 20 cm, hemispherical). Type: KENYA. Taita Hills,
and diverged adopting a large number of small genera P. Kamau & M.J.M. Christenhusz 638 (holotype EA;
that were long recognised as sections or series (Iltis isotype K). Etymology: This species is widespread
1952). This splitting of an otherwise monophyletic across tropical Africa, hence the name.
clade (Inda et al., 2008; Patchell et al., 2014) seems to Ptisana kingii (Copel.) Christenh., comb. nov.
be unnecessarily destabilising the classification of this Basionym: Marattia kingii Copel., Phil. J. Sci., Bot. 6,
clade and hence here we accept a broader concept pt. C: 66. 1911.
of Cleome, keeping in mind the monophyly of the
Ptisana koordersii (Alderw.) Christenh., comb. nov.
genus. Patchell et al. (2014) also suggested expanding
Basionym: Marattia koordersii Alderw., Malayan
Cleomella DC., resulting in the acceptance of four
Ferns, Suppl.: 529. 1917.
genera (Cleome, Cleomella, Peritoma DC. and Rorida
J.F.Gmel.) in Cleomaceae. Ptisana senterreana Christenh., sp. nov. (Figure 2).
Similar to P. robusta (Alston) Senterre & Rouhan,
Capparaceae – Capparis L. in its traditional concept but is smaller in habit and has shortly oblong, sessile
is polyphyletic (Tamboli et al., 2018) with the Old World pinnules (less than three times longer than wide) with
clade being separated from the New World clade abruptly caudate to acuminate apices. It is one of the
by a grade of genera. The Old World clade includes smallest species of Ptisana in Africa with petioles up
Apophyllum F.Muell. and the type of Capparis and is to to 60 cm. Type: CAMEROON. Central Province, near
be maintained as a single genus. New World Capparis Nkolbison, 7 km W. of Yaounde, 18 January 1962,
has in the past been subdivided into a several genera, F. Breteler 2500 (holotype P [P01365197]; isotype
probably to preserve the rather divergent Steriphoma WAG [WAG.1736404]). Etymology: This species is
Spreng., but this has greatly destabilised the named for Bruno Senterre, who first recognised this
nomenclature of the American clade. The oldest name species as new to science.
is Morisonia L. and we transfer the entire American Ptisana werneri (Rosenst.) Christenh., comb. nov.
clade to this genus. Basionym: Marattia werneri Rosenst., Repert. Spec.
We have done our utmost to be as precise and Nov. Regni Veg. 5: 44. 1908.
complete as possible and we hope this system will
progress to a more stable vascular plant taxonomy in
the near future. Families are numbered according to
the list that will be provided in Byng & Christenhusz (in
prep.). Ferns are marked with ‘F’, gymnosperms with
‘G’.

3
A B
Figure 1: Ferns. A-B, Ptisana africana Christenh., sp. nov.; C, Alsophila societarum (Baker) Christenh.; and D,
Pteridium pseudocaudatum (Clute) Christenh.. All images © Maarten Christenhusz.

Figure 2: Ferns. Isotype of Ptisana senterreana Christenh., sp. nov.. Image used with permission by Naturalis
Biodiversity Center.

5
F9. Hymenophyllaceae. References: Ebihara et al. Alsophila boiviniiformis (Rakotondr. & Janssen)
(2007), Christenhusz & Chase (2014). Christenh., comb. nov. Basionym: Cyathea
Note: Hymenophyllaceae is divided in two clades, one boiviniiformis Rakotondr. & Janssen, Adansonia 29:
represented by Hymenophyllum Sm., the other by 198. 2007.
Trichomanes L. Some authors have separated the Alsophila conferta (Rakotondr. & Janssen) Christenh.,
second clade into a great number of genera, which comb. nov. Basionym: Cyathea conferta Janssen &
is not in keeping with the traditional concept of Rakotondr., Adansonia 30: 272. 2008.
Trichomanes. It is therefore preferred to maintain Alsophila dilatata (Rakotondr. & Janssen) Christenh.,
Trichomanes in the broad sense. comb. nov. Basionym: Cyathea dilatata Rakotondr.
Trichomanes beaverianum (Senterre & Rouhan) & Janssen, Adansonia 30: 252. 2008.
Christenh., comb. nov. Basionym: Didymoglossum Alsophila emilei (Janssen & Rakotondr.) Christenh.,
beaverianum Senterre & Rouhan, Phytotaxa 292: comb. nov. Basionym: Cyathea emilei Janssen &
206. 2017. Rakotondr., Adansonia 30: 293. 2008.
Trichomanes franceae (Bauret & Dubuisson) Alsophila hebes (Janssen & Rakotondr.) Christenh.,
Christenh., comb. nov. Basionym: Abrodictyum comb. nov. Basionym: Cyathea hebes Janssen &
franceae Bauret & Dubuisson, Phytotaxa 284: 163. Rakotondr., Adansonia 30: 276, 2008.
2016. Alsophila impolita (Rakotondr. & Janssen) Christenh.,
Trichomanes hokurikuense (Ebihara) Christenh., comb. nov. Basionym: Cyathea impolita Rakotondr.
comb. nov. Basionym: Vandenboschia hokurikuensis & Janssen, Adansonia 29: 203. 2007.
Ebihara, Acta Phytotax. Geobot. 60: 33. 2009. Alsophila ivohibensis (C.Chr.) Christenh., comb. nov.
Trichomanes lofoushanense (Ching) Christenh., comb. Basionym: Cyathea quadrata Baker var. ivohibensis
nov. Basionym: Vandenboschia lofoushanensis C.Chr., Dansk Bot. Ark. 7: 28. 1932.
Ching in Chien & Chun, Fl. Reipubl. Pop. Sin. 2: 356. Alsophila lisyae (Janssen & Rakotondr.) Christenh.,
1959. comb. nov. Basionym: Cyathea lisyae Janssen &
Trichomanes mascarenense (Pynee & Dubuisson) Rakotondr., Adansonia 30: 258. 2008.
Christenh. comb. & stat. nov. Basionym: Alsophila longispina (Janssen & Rakotondr.) Christenh.,
Crepidomanes minutum (Blume) K.Iwats. var. comb. nov. Basionym: Cyathea longispina Janssen &
mascarenensis Pynee & Dubuisson, Phytotaxa 62: Rakotondr., Adansonia 30: 300. 2008.
28. 2012.
Alsophila meridionalis (Janssen & Rakotondr.)
Trichomanes miuraense (Ebihara) Christenh., comb. Christenh., comb. nov. Basionym: Cyathea
nov. Basionym: Vandenboschia miuraensis Ebihara, meridionalis Janssen & Rakotondr., Adansonia 30:
Acta Phytotax. Geobot. 60: 35. 2009. 260. 2008.
Trichomanes nicobaricum (R.D.Dixit & Sweta Singh) Alsophila obtecta (Rakotondr. & Janssen) Christenh.,
Christenh., comb. nov. Basionym: Microgonium comb. nov. Basionym: Cyathea obtecta Rakotondr.
nicobaricum R.D.Dixit & Shweta Singh, Indian Fern J. & Janssen, Adansonia 30: 307. 2008.
22(1-2): 182. 2005.
Alsophila pseudobellisquamara (Janssen &
Trichomanes tamonii (Y.Robert & Dubuisson) Rakotondr.) Christenh., comb. nov. Basionym:
Christenh., comb. & stat. nov. Basionym: Cyathea pseudobellisquamata Janssen & Rakotondr.,
Crepidomanes inopinatum (Pic.Serm.) J.P.Roux var. Adansonia 30: 310. 2008.
tamonii Y.Robert & Dubuisson. Phytotaxa 173: 165.
Alsophila roroka (Hovenkamp) Christenh., comb. nov.
2014.
Basionym: Cyathea roroka Hovenkamp, Blumea 33:
Trichomanes tubiflora (F.S.Wagner) Christenh., comb. 397. 1988.
nov. Basionym: Vandenboschia tubiflora F.S.Wagner,
Alsophila rouhaniana (Rakotondr. & Janssen)
Contr. Univ. Michigan Herb. 20: 243. 1995.
Christenh., comb. nov. Basionym: Cyathea
rouhaniana Rakotondr. & Janssen, Adansonia 29:
F16. Cyatheaceae. Reference: Korall et al. (2007). 207. 2007.
Note: As the genus Alsophila R.Br. appears to be Alsophila pacifica Christenh., nom. nov. Replaced
now universally accepted (e.g. Christenhusz & synonym: Cyathea samoensis Baker, J. Bot. 14: 9.
Chase, 2014; PPG, 2016), the following species are 1876, not: A. samoensis Brack. Etymology: This
transferred. species occurs in the Pacific.
Alsophila basirotundata (Rakotondr. & Janssen) Alsophila societarum (Baker) Christenh., comb. nov.
Christenh., comb. nov. Basionym: Cyathea Basionym: Cyathea societarum Baker, Syn. Fil.
basirotundata Rakotondr. & Janssen, Adansonia 30: (Hooker & Baker) ed. 2: 453. 1874. (Figure 1C)
270. 2008.

Alsophila valdesquamata (Janssen & Rakotondr.) F21. Dennstaedtiaceae

Christenh., comb. nov. Basionym: Cyathea Note: Pteridium Gled. ex Scop. has received some
valdesquamata Janssen & Rakotondr., Adansonia attention in phylogenetic studies, and a system of
30: 358. 2008. species and subspecies was proposed (Der et al.,
Cyathea flava (R.M.Tryon) Christenh., comb. nov. 2009). We think it better to elevate subspecific
Basionym: Trichipteris flava R.M.Tryon, Fieldiana, taxa as species as they are geographically isolated.
Bot. n. s. 20: 125. 1989. This will simplify the taxonomy of a genus with a
contentious nomenclatural history.
F17. Cystodiaceae Pteridium campestre (Schrad.) Christenh., comb. nov.
Basionym: Pteris campestris Schrad., Gött. gel. Anz.
Note: The following taxon is best treated at the species
Aufs. Kön. Gesellsch. Wiss. 1824: 871. 1824.
level. It is the second species known in this family.
Pteridium decompositum (Gaudich.) Christenh., comb.
Cystodium solomonense (J.R.Croft) Christenh., comb.
nov. Basionym: Pteris decomposita Gaudich., Voy.
& stat. nov. Basionym: Cystodium sorbifolium subsp.
Uranie, Bot. 4: 393. 1826.
solomonense J.R.Croft, Kew Bull. 41: 902. 1986.
Pteridium pseudocaudatum (Clute) Christenh., comb.
& stat. nov. Basionym: Pteridium aquilinum (L.)
F19. Saccolomataceae. Reference: Luong et al. (2015). Kuhn var. pseudocaudatum Clute, Fern Bull. 8: 39.
Note: The genus Orthiopteris Copel. is similar 1900. (Figure 1D)
morphologically to Saccoloma Kaulf. (Luong et al. Pteridium pubescens (Underw.) Christenh., comb. &
2015), and molecular studies do not discriminate stat. nov. Basionym: Pteridium aquilinum (L.) Kuhn
between the two genera. As it is clear that the two var. pubescens Underw., Our Native Ferns, ed. 6: 91.
are closely related and to prevent recognition of a 1900.
paraphyletic genus, Saccoloma is here expanded
to include Orthiopteris. Most of the species of
Orthiopteris have already been combined into F22. Pteridaceae. Reference: Eiserhardt et al. (2011).
Saccoloma in the past. Note: Subfamily Cheilanthoideae consisted of several
Saccoloma acuminatum (Rosenst.) Christenh., comb. genera that were found not to be monophyletic.
nov. Basionym: Dennstaedtia acuminata Rosenst., As a result a number of genera were coined that
Hedwigia 56: 350. 1915. were morphologically indiscriminate. As numerous
hybrids are known, a number of intergeneric hybrids
Saccoloma cicutarioides (Baker) Christenh., comb.
would additionally need to be coined. As this has
nov. Basionym: Davallia cicutarioides Baker, J. Bot.
been destabalising the nomenclature of genera like
(Britten) 28: 106. 1890.
Cheilanthes Sw., Doryopteris J.Sm. and Pellaea Link,
Saccoloma samoense (Hovenkamp & T.T.Luong) it seems reasonable to treat the entire subfamily
Christenh., comb. nov. Basionym: Orthiopteris as a single genus, stabalising this genus in future
samoensis Hovenkamp & T.T.Luong, PhytoKeys 53: taxonomic study, which is in line with the treatment
60. 2015. of other large genera in ferns like Asplenium and
Pteris L.. Hemionitis L. has nomenclatural priority
F20. Lindsaeaceae. Reference: Lehtonen et al. (2010). and thus all taxa are transferred into this genus.
Note: A few species were omitted when the family Hemionitis achariorum Christenh., nom. nov. Replaced
was revised. They are hereby transfered into their synonym: Pteris palmata Willd. sp. 5. 357. 1810,
correct genus. not: Hemionitis palmata L., Sp. Pl. 2: 1077. 1753.
Lindsaea stortii (Alderw.) Christenh., comb. nov. Etymology: Named for the Acharis, a race of sad
Basionym: Schizoloma stortii Alderw., Bull. Jard. Bot. wormlike creatures from The Neverending Story
Buitenzorg Ser. 2, XVI: 36. 1914. (Ende, 1979).
Odontosoria goudotiana (Kunze) Christenh., comb. Hemionitis aconitifolia Christenh., nom. nov. Replaced
nov. Basionym: Davallia goudotiana Kunze, Analecta synonym: Cassebeera paradoxa Fée, Mém. Fam.
Pterid.: 35. 1837. Foug. 7: 30. 1857, not: Hemionitis paradoxa (R.Br.)
Christenh. Etymology: This species have leaves that
Odontosoria humbertii (Tardieu) Christenh., comb.
resemble Aconitum (Ranunculaceae).
nov. Basionym: Sphenomeris humbertii Tardieu,
Notul. Syst. (Paris) 15: 182. 1956. Hemionitis acta (Sehnem) Christenh., comb. nov.
Basionym: Adiantopsis acta Sehnem, Pesquisas 3:
Odontosoria madagascariensis (Baker) Christenh.,
508. 1959.
comb. nov. Basionym: Lindsaea madagascariensis
Baker, J. Linn. Soc., Bot. 16: 198. 1877, as ‘Lindsaya Hemionitis acutiloba (Prantl) Christenh., comb. nov.
madagascariensis’. Basionym: Pellaea acutiloba Prantl, Bot. Jahrb. Syst.
3: 425. 1882.

7
A B
C D
E F
Figure 3: Ferns. A, Hemionitis amarganthica Christenh.; B, Hemionitis concolor (Langsd. & Fisch.) Christenh.; C,
Hemionitis himalaica (Fraser-Jenk.) Christenh.; D, Hemionitis hispida (Mett. ex Kuhn) Christenh.; E, Hemionitis
querquobadii Christenh.; and F, Hemionitis sinuata (Lag. ex Sw.) Christenh.. All images © Maarten Christenhusz.

Hemionitis adiantoides (T.C.Chambers & P.A.Farrant) Hemionitis amarganthica Christenh., nom. nov.
Christenh., comb. nov. Basionym: Cheilanthes Replaced synonym: Pteris argentea S.G.Gmel., Novi
adiantoides T.C.Chambers & P.A.Farrant, Telopea 4: Comment. Acad. Sci. Imp. Petrop.12: 519. 1769, not:
532. 1991. Hemionitis argentea Willd. Etymology: Named for
Hemionitis adiantopsis Christenh., nom. nov. Replaced Amarganth, a city in the magical land of Fantastica
synonym: Hypolepis pedata Hook., Sp. Fil. 2: 73. from The Neverending Story (Ende, 1979). (Figure
1852, not: Hemionitis pedata Sw. Etymology: Named 3A)
for the former genus Adiantopsis, now included Hemionitis ambigua (Fée) Christenh., comb. nov.
in Hemionitis. It means ‘resembling Adiantum’ Basionym: Synochlamys ambigua Fée, Mém. Soc.
(Pteridaceae) Sci. Nat. Strasbourg 5: 35. 1857.
Hemionitis adornata (Yesilyurt) Christenh., comb. nov. Hemionitis anceps (Blanf.) Christenh., comb. nov.
Basionym: Doryopteris adornata Yesilyurt, Bot. J. Basionym: Cheilanthes anceps Blanf., J. Simla
Linn. Soc. 158: 73. 2008. Natural. Soc. 1: 21. 1886.
Hemionitis aemula (Maxon) Christenh., comb. nov. Hemionitis andromedifolia (Kaulf.) Christenh., comb.
Basionym: Cheilanthes aemula Maxon, Contr. U.S. nov. Basionym: Pteris andromedifolia Kaulf., Enum.
Natl. Herb. 10: 495. 1908. Filic.: 188. 1824.
Hemionitis aethiopica (Saiki) Christenh., comb. Hemionitis angelica (K.Wood & W.H.Wagner)
nov. Basionym: Aleuritopteris aethiopica Saiki, J. Christenhusz, comb. nov. Basionym: Doryopteris
Phytogeogr. Taxon. 32: 81. 1984. angelica K.Wood & W.H.Wagner, Contr. Univ.
Hemionitis affinis (Mett.) Christenh., comb. nov. Michigan Herb. 22: 147. 1999.
Basionym: Cheilanthes affinis Mett., Abh. Hemionitis angolensis (Schelpe) Christenh., comb.
Senckenberg. Naturf. Ges. 3: 63. 1859. nov. Basionym: Pellaea angolensis Schelpe, Garcia
Hemionitis agetae (Saiki) Christenh., comb. nov. de Orta, Ser. Bot. 3: 53. 1976.
Basionym: Aleuritopteris agetae Saiki, J. Phytogeogr. Hemionitis angulosa (Bory ex Willd.) Christenh., comb.
Taxon. 32: 1. 1984. nov. Basionym: Pteris angulosa Bory ex Willd., Sp. Pl.
Hemionitis aiuolae Christenh., nom. nov. Replaced ed. quarta: 377. 1810.
synonym: Cheilanthes boivini Mett. ex Kuhn, Fil. Afr. Hemionitis angusta (R.M.Tryon) Christenh., comb.
69. 1868, not: Hemionitis boivinii (Hook.) Christenh. nov. Basionym: Notholaena angusta R.M.Tryon,
Etymology: Named for Dame Eyola (or Aiuóla), a Rhodora 63: 80. 1961.
plant taking the form of a motherly woman who Hemionitis angustata (Sehnem) Christenh., comb.
lives in the House of Change; from The Neverending nov. Basionym: Doryopteris angustata Sehnem, Fl.
Story (Ende, 1979). Ilustr. Catarin. 1: 140. 1972.
Hemionitis alabamensis (Buckley) Christenh., comb. Hemionitis angustifolia (Kunth) Christenh., comb. nov.
nov. Basionym: Pteris alabamensis Buckley, Amer. J. Basionym: Cheilanthes angustifolia Kunth, Nov. Gen.
Sci. Arts 45: 177. 1843. Sp., quarto ed. 1: 21. 1815.
Hemionitis alata (Prantl) Christenh., comb. nov. Hemionitis angustifrondosa (Alston) Christenh., comb.
Basionym: Adiantopsis alata Prantl, Gartenflora 32: nov. Basionym: Cheilanthes angustifrondosa Alston,
99. 1883. Bull. Brit. Mus. (Nat. Hist.), Bot. 1: 48. 1952.
Hemionitis albofusca (Baker) Christenh., comb. nov. Hemionitis apiacea (Mickel) Christenh., comb. nov.
Basionym: Cheilanthes albofusca Baker, Bull. Misc. Basionym: Cheilanthes apiacea Mickel, Mem. New
Inf. Kew 1895(99): 54. 1895. York Bot. Gard. 88: 182. 2004.
Hemionitis albomarginata (Clarke) Christenh., comb. Hemionitis apparicioi (Brade) Christenh., comb. nov.
nov. Basionym: Cheilanthes albomarginata Clarke, Basionym: Doryopteris apparicioi Brade, Arq. Jard.
Trans. Linn. Soc. Lond. II. Bot. 1: 456. 1880. Bot. Rio de Janeiro 18: 27. 1965.
Hemionitis aliena (Maxon) Christenh., comb. nov. Hemionitis arequipensis (Maxon) Christenh., comb.
Basionym: Notholaena aliena Maxon, Contr. U.S. nov. Basionym: Notholaena arequipensis Maxon,
Natl. Herb. 17: 605. 1916. Smithsonian Misc. Collect. 65: 9. 1915.
Hemionitis alleniae (R.M.Tryon) Christenh., comb. nov. Hemionitis argyrophylla (Sw.) Christenh., comb. nov.
Basionym: Doryopteris alleniae R.M.Tryon, Contrib. Basionym: Pteris argyrophylla Sw., Syn. Fil.: 105.
Gray Herb. 191: 91. 1962. 1806.
Hemionitis allosuroides (Mett.) Christenh., comb. Hemionitis arizonica (Maxon) Christenh., comb. &
nov. Basionym: Cheilanthes allosuroides Mett., Abh. stat. nov. Basionym: Cheilanthes pyramidalis Fée
Senckenberg. Naturf. Ges. 3: 76. 1859. subsp. arizonica Maxon, Amer. Fern J. 8: 116. 1918.

9
Hemionitis artax Christenh., nom. nov. Replaced Hemionitis bhutanica (Fraser-Jenk. & Wangdi)
synonym: Cheilanthes dealbata Pursh, Fl. Amer. Christenh., comb. nov. Basionym: Cheilanthes
Sept. 2: 671. 1814, not: Hemionitis dealbata bhutanica Fraser-Jenk. & Wangdi, Fern Gaz. 18: 220.
Willd. Etymology: Named for Atreju’s horse in The 2009.
Neverending Story (Ende, 1979). Hemionitis bicolor (Roxb.) Christenh., comb. nov.
Hemionitis aschenborniana (Klotzsch) Christenh., Basionym: Pteris bicolor Roxb., Calcutta J. Nat. Hist.
comb. nov. Basionym: Notholaena aschenborniana 4: 507. 1844.
Klotzsch, Linnaea 20: 417. 1847. Hemionitis boivinii (Hook.) Christenh., comb. nov.
Hemionitis atreyu Christenh., nom. nov. Replaced Basionym: Pellaea boivinii Hook., Sp. Fil. 2: 147.
synonym: Cheilanthes acrostica Tod., Giorn. Sci. 1858.
Nat. Econ. Palermo 1: 215. 1866. not: Hemionitis Hemionitis bolborrhiza (Mickel & Beitel) Christenh.,
acrosticha Noronha (1827). Etymology: Named for comb. nov. Basionym: Cheilanthes bolborrhiza
Atreyu, the boy warrior in The Neverending Story Mickel & Beitel, Mem. New York Bot. Gard. 46: 107.
(Ende, 1979). 1988.
Hemionitis atropurpurea (L.) Christenh., comb. nov. Hemionitis bonapartei (J.P.Roux) Christenh., comb.
Basionym: Pteris atropurpurea L., Sp. Pl. 2: 1076. nov. Basionym: Cheilanthes bonapartei J.P.Roux,
1753. Strelitzia 23: 180. 2009.
Hemionitis aurantiaca (Cav.) Christenh., comb. nov. Hemionitis bonariensis (Willd.) Christenh., comb. nov.
Basionym: Pteris aurantiaca Cav., Descr. Pl.: 266. Basionym: Acrostichum bonariense Willd., Sp. Pl. ed.
1801. quarta 5: 114. 1810.
Hemionitis aurea (Link-Pérez, Seabolt & Ledford) Hemionitis botswanae (Schelpe & N.C.Anthony)
Christenh., comb. nov. Basionym: Adiantopsis aurea Christenh., comb. nov. Basionym: Cheilanthes
Link-Pérez, Seabolt & Ledford, Syst. Bot. 41: 913. botswanae Schelpe & N.C.Anthony, Contr. Bolus
2016. Herb. 10: 151. 1982.
Hemionitis aureolina (Yatsk. & Arbeláez). Christenh., Hemionitis brachyptera (T.Moore) Christenh., comb.
comb. nov. Basionym: Notholaena aureolina Yatsk. & nov. Basionym: Platyloma brachyptera T.Moore,
Arbeláez, Novon 18: 120. 2008. Replaced synonym: Gard. Chron. 1873: 141. 1873.
Cheilanthes aurea Baker, Syn. Fil.: 452. 1868, not: Hemionitis brachypus (Kunze) Christenh., comb. nov.
Hemionitis aurea Willd. Basionym: Cheilanthes squamosa Gillies & Hook. ex
Hemionitis auryn Christenh., nom. nov. Replaced Grev. var. brachypus Kunze, Linnaea 18: 340. 1844.
synonym: Notholaena aurantiaca D.C.Eaton, Proc. Hemionitis bradburii (Hook.) Christenh., comb. nov.
Amer. Acad. Arts Sci. 22: 462. 1887, not: Hemionitis Basionym: Cheilanthes bradburii Hook., Sp. Filic. 2:
aurantiaca (Cav.) Christenh. Eymology: Named for 97. 1852. Replaced synonym: Cheilanthes tomentosa
AURYN a protective amulet in The Neverending Story Link, Hort. Berol. 2: 42. 1833, not: Hemionitis
(Ende, 1979). tomentosa Raddi
Hemionitis austrotenuifolia (H.M.Quirk & Hemionitis brandegeei (D.C.Eaton) Christenh., comb.
T.C.Chambers) Christenh., comb. nov. Basionym: nov. Basionym: Cheilanthes brandegeei D.C.Eaton,
Cheilanthes austrotenuifolia H.M.Quirk & Bull. Torrey Bot. Club 17: 215. 1890.
T.C.Chambers, Austral. J. Bot. 31: 510. 1983.
Hemionitis brandtii (Franch & Sav.) Christenh., comb.
Hemionitis baturitensis (Brade) Christenh., comb. nov. Basionym: Cheilanthes brandtii Franch. & Sav.,
nov. Basionym: Doryopteris baturitensis Brade, Enum. Pl. Jap. 2: 212. 1879.
Rodriguesia 4: 297. 1940.
Hemionitis branneri (Copel.) Christenh., comb. nov.
Hemionitis beitelii (Mickel) Christenh., comb. nov. Basionym: Doryopteris branneri Copel., Phil. J. Sci.
Basionym: Cheilanthes beitelii Mickel, Mem. New 38: 148. 1929.
York Bot. Gard. 46: 107. 1988.
Hemionitis brasiliensis (Baker) Christenh., comb. nov.
Hemionitis belangeri (Bory) Christenh., comb. nov. Basionym: Pellaea brasiliensis Baker, Bot. Jahrb. Syst.
Basionym: Pteris belangeri Bory in Belang., Voy. 17: 522. 1893, not: Hemionitis brasiliana Desv.
Indes Or. 2: 44. 1833.
Hemionitis breweri (D.C.Eaton) Christenh., comb. nov.
Hemionitis belensis (Weath. ex Copel.) Christenh., Basionym: Pellaea breweri D.C.Eaton, Proc. Amer.
comb. nov. Basionym: Cheilanthes belensis Weath. Acad. Arts Sci. 6: 555. 1865.
ex Copel., Univ. Calif. Publ. Bot. 18: 218. 1942.
Hemionitis bridgesii (Hook.) Christenh., comb. nov.
Hemionitis bergiana (Schltdl.) Christenh., comb. nov. Basionym: Pellaea bridgesii Hook., Sp. Fil. 2: 238.
Basionym: Cheilanthes bergiana Schltdl., Adumbr. Pl. 1858.
1: 51. 1825.

Hemionitis brownii (Desv.) Christenh., comb. nov. Hemionitis cheilanthoides (R.M.Senna) Christenh.,
Basionym: Notholaena brownii Desv., Mém. Soc. comb. nov. Basionym: Adiantopsis cheilanthoides
Linn. Paris 6: 220. 1827. R.M.Senna, Iheringia, Bot. 59: 108. 2004.
Hemionitis bryopoda (Maxon) Christenh., comb. nov. Hemionitis cheiloglyphis (Fée) Christenh., comb. nov.
Basionym: Notholaena bryopoda Maxon, Proc. Biol. Basionym: Myriopteris cheiloglyphis Fée, Mém.
Soc. Washington 18: 205. 1905. Foug. 8: 77. 1857.
Hemionitis buchtienii (Rosenst.) Christenh., comb. Hemionitis cheiloplecton Christenh., nom. nov.
nov. Basionym: Notholaena buchtienii Rosenst., Replaced synonym: Pteris rigida Sw., Syn. Filic.: 104,
Repert. Spec. Nov. Regni Veg. 5: 238. 1908. 299. 1806, not: Hemionitis rigida Cav. Etymology:
Hemionitis bullosa (Kunze) Christenh., comb. nov. Named for former genus Cheiloplecton, now
Basionym: Cheilanthes bullosa Kunze, Linnaea 24: included in Hemionitis.
274. 1851. Hemionitis chihuahuaensis (Saiki) Christenh., comb.
Hemionitis caesia (Christ) Christenh., comb. nov. nov. Basionym: Aleuritopteris chihuahuaensis Saiki,
Basionym: Cheilanthes caesia Christ, Bull. Acad. Int. J. Phytogreogr. Taxon. 32: 85. 1984.
Géogr. Bot. Mans 16: 133. 1906. Hemionitis chilensis (Fée & J.Remy) Christenh., comb.
Hemionitis cairon Christenh., nom. nov. Replaced nov. Basionym: Cincinalis chilensis Fée & J.Rémy, Fl.
synonym: Notholaena peninsularis Maxon & Weath., Chil. 6: 497. 1853.
Contr. Gray Herb. 127: 15. 1939. Etymology: Named Hemionitis chinensis (Baker) Christenh., comb. nov.
for centaur Cairon in The Neverending Story (Ende, Basionym: Notholaena chinensis Baker, Gard. Chr. n.
1979). s. 14: 494. 1880.
Hemionitis calidirupium (Brownsey & Lovis) Christenh., Hemionitis chipinquensis (Knobloch & Lellinger)
comb. nov. Basionym: Pellaea calidirupium Brownsey Christenh., comb. nov. Basionym: Cheilanthes
& Lovis, New Zealand J. Bot. 28: 197. 1990. chipinquensis Knobloch & Lellinger, Amer. Fern J. 59:
Hemionitis californica (Hook.) Christenh., comb. nov. 8. 1969.
Basionym: Hypolepis californica Hook., Sp. Fil. 2: 71. Hemionitis chlorophylla (Sw.) Christenh., comb. nov.
1851. Basionym: Cheilanthes chlorophylla Sw., Kongl.
Hemionitis calomelanos (Sw.) Christenh., comb. nov. Vetensk. Acad. Handl. 1817: 76. 1817.
Basionym: Pteris calomelanos Sw., J. Bot. (Schrader) Hemionitis christensenii Christenh., nom. nov.
1800(2): 70. 1801. Replaced synonym: Pellaea smithii C.Chr., Acta Horti
Hemionitis camposportoi (Brade) Christenh., comb. Gothob. 1: 84. 1924, not: Hemionitis smithii (Trev.)
nov. Basionym: Doryopteris camposportoi Brade, C.Chr. Etymology: Named for Carl Christensen, who
Arq. Inst. Biol. Veg. Rio de Janeiro 3: 3. 1936. first described this species.
Hemionitis candida (M.Martens & Galeotti) Christenh., Hemionitis christii (Fraser-Jenk. & Yatsk.) Christenh.,
comb. nov. Basionym: Cheilanthes candida comb. nov. Basionym: Cheilanthes christii Fraser-
M.Martens & Galeotti, Nouv. Mém. Acad. Roy. Sci. Jenk. & Yatsk., Indian Fern J. 27: 213. 2011. Replaced
Bruxelles 15: 73. 1842. synonym: Pteris paupercula Christ, Bull. Acad.
Int. Géogr. Bot. 16: 131. 1906, not: Hemionitis
Hemionitis cantangensis (R.M.Tryon) Christenh.,
paupercula (Kunze) Christenh.
comb. nov. Basionym: Notholaena cantangensis
R.M.Tryon, Rhodora 63: 81. 1961. Hemionitis chrysophylla (Hook.) Christenh., comb.
nov. Basionym: Cheilanthes chrysophylla Hook., Sp.
Hemionitis capensis (Thunb.) Christenh., comb. nov.
Fil. 2: 113. 1852.
Basionym: Adiantum capense Thunb., Prodr. Pl.
Cap.: 173. 1800. Hemionitis chusana (Hook.) Christenh., comb. nov.
Basionym: Cheilanthes chusana Hook., Sp. Fil. 2: 95.
Hemionitis carlottahalliae (W.H.Wagner & E.F.Gilbert)
1852.
Christenh., comb. nov. Basionym: Cheilanthes
carlottahalliae W.H.Wagner & E.F.Gilbert, Amer. J. Hemionitis cinnamomea (Baker) Christenh., comb.
Bot. 44: 738. 1957. nov. Basionym: Notholaena cinnamomea Baker, Syn.
Filic. (ed. 2): 515. 1874.
Hemionitis caudata (R.Br.) Christenh., comb. nov.
Basionym: Cheilanthes caudata R.Br., Prodr. Fl. Nov. Hemionitis clevelandii (D.C.Eaton) Christenh., comb.
Holland.: 156. 1810. nov. Basionym: Cheilanthes clevelandii D.C.Eaton,
Bull. Torrey Bot. Club 6: 33. 1875.
Hemionitis ceterachoides (A.W.Klopper & Klopper)
Christenh., comb. nov. Basionym: Cheilanthes Hemionitis cochisensis (Goodd.) Christenh., comb.
ceterachoides A.W.Klopper & Klopper, Ferns S. Afr. nov. Basionym: Notholaena cochisensis Goodd.,
Compreh. Guide: 747. 2011. Muhlenbergia 8: 93. 1912.

11
Hemionitis collina (Raddi) Christenh., comb. nov. Hemionitis cubensis (Weath ex R.M.Tryon) Christenh.,
Basionym: Pteris collina Raddi, Opusc. Sci. 3: 292. comb. nov. Basionym: Notholaena cubensis Weath.
1819. ex R.M.Tryon, Contr Gray Herb. 179: 49. 1956.
Hemionitis columbina (Hook.) Christenh., comb. & Hemionitis cucullans (Fée) Christenh., comb. nov.
stat. nov. Basionym: Pellaea lomariacea Hook. var. Basionym: Cheilanthes cucullans Fée, Mém. Foug. 7:
columbina Hook., Sp. Filic. 2: 133. 1858. 39. 1857.
Hemionitis complanata (A.R.Sm.) Christenh., comb. Hemionitis cuneata (Kaulf. ex Link) Christenh. comb.
nov. Basionym: Cheilanthes complanata A.R.Sm., nov. Basionym: Cheilanthes cuneata Kaulf. ex Link,
Amer. Fern J. 70: 19. 1980. Hort. Berol. 2: 40. 1833.
Hemionitis concolor (Langsd. & Fisch.) Christenh., Hemionitis cuspidata (Copel.) Christenh., comb. nov.
comb. nov. Basionym: Pteris concolor Langsd. & Basionym: Doryopteris cuspidata Copel., Phil. J. Sci.
Fisch., Icon. Filic. 19: pl. 21. 1810. (Figure 3B) 38: 148. 1929.
Hemionitis conformis (K.U.Kramer & R.M.Tryon) Hemionitis cyclophylla (A.R.Sm.) Christenh., comb.
Christenh., comb. nov. Basionym: Doryopteris nov. Basionym: Doryopteris cyclophylla A.R.Sm.,
conformis K.U.Kramer & R.M.Tryon, Ann. Missouri Ann. Missouri Bot. Gard. 77: 262. 1990.
Bot. Gard. 42: 213. 1955. Hemionitis cymbiformis (Prado) Christenh., comb. nov.
Hemionitis connectens (C.Chr.) Christenh., comb. nov. Basionym: Pellaea cymbiformis Prado, Amer. Fern J.
Basionym: Pellaea connectens C.Chr., Acta Horti 83: 129. 1993.
Gothob. 1: 84. 1924. Hemionitis dactylifera (Link-Pérez & Hickey) Christenh.,
Hemionitis contigua (Baker) Christenh., comb. nov. comb. nov. Basionym: Adiantopsis dactylifera Link-
Basionym: Cheilanthes contigua Baker, Syn. Fil. 476. Pérez & Hickey, Syst. Bot. 36: 576. 2011.
1874. Hemionitis dalhousiae (Hook.) Christenh., comb. nov.
Hemionitis contracta (Kunze) Christenh., comb. nov. Basionym: Cheilanthes dalhousiae Hook., Sp. Fil. 2:
Basionym: Cheilanthes contracta Kunze, Linnaea 28: 80. 1852.
243. 1850. Hemionitis davidsei (A.R.Sm.) Christenh., comb.
Hemionitis cooperae (D.C.Eaton) Christenh., comb. nov. Basionym: Doryopteris davidsei A.R.Sm., Ann.
nov. Basionym: Cheilanthes cooperae D.C.Eaton, Missouri Bot. Gard. 77: 262. 1990.
Bull. Torrey Bot. Club 6: 33. 1875. Hemionitis deboeri (Verdc.) Christenh., comb. nov.
Hemionitis cordifolia (Sessé & Moc.) Christenh., comb. Basionym: Cheilanthes deboeri Verdc., Fl. Trop. E.
nov. Basionym: Adiantum cordifolium Sessé & Moc., Africa, Adiantac.: 40. 2002.
Naturaleza (Mexico City) ser. 2, 1(App.): 182. 1890, Hemionitis decipiens (Hook.) Christenh., comb. nov.
not: Hemionitis cordifolia Roxb. ex Wall., nom. inval., Basionym: Pteris decipiens Hook., Sp. Fil. 2: 209.
nom. nud. 1858.
Hemionitis covillei (Maxon) Christenh., comb. nov. Hemionitis decomposita (M.Martens & Galeotti)
Basionym: Cheilanthes covillei Maxon, Proc. Biol. Christenh., comb. nov. Basionym: Allosorus
Soc. Washington 31: 147. 1918. decompositus M.Martens & Galeotti, Mém. Foug.
Hemionitis crassifolia (Houlston & T.Moore) Christenh., Mex.: 48. 1842.
comb. nov. Basionym: Notholaena crassifolia Hemionitis decora (Brackenr.) Christenh., comb. nov.
Houlston & T.Moore, Gard. Mag. Bot. 3: 20. 1851. Basionym: Doryopteris decora Brackenr., U.S. Expl.
Hemionitis cremea (Ching) Christenh., comb. nov. Exped., Filic. 16: 103. 1854.
Basionym: Aleuritopteris cremea Ching, Acta Hemionitis decurrens (Mickel) Christenh., comb. nov.
Phytotax. Sin. 19: 65. 1981. Basionym: Cheilanthes decurrens Mickel, Mem. New
Hemionitis crenata (R.M.Tryon) Christenh., comb. nov. York Bot. Gard. 88: 191. 2004.
Basionym: Pellaea crenata R.M.Tryon, Contr. Gray Hemionitis decursiva (Forssk.) Christenh., comb. nov.
Herb. 143: 68. 1942. Basionym: Pteris decursiva Forssk., Fl. Aegypt.-Arab.:
Hemionitis crenulans (Fée) Christenh., comb. nov. 186. 1775.
Basionym: Pellaea crenulans Fée, Crypt. Vasc. Brésil Hemionitis delavayi (Baker) Christenh., comb. nov.
2: 27. 1872. Basionym: Gymnogramme delavayi Baker, Ann. Bot.
Hemionitis crinoidea (Link-Pérez & Hickey) Christenh., (Oxford) 5: 484. 1891.
comb. nov. Basionym: Adiantopsis crinoidea Link- Hemionitis delicatessa Christenh., nom. nov. Replaced
Pérez & Hickey, Syst. Bot. 36: 571. 2011. synonym: Cheilanthes delicatula Tagawa & K.lwats.,
Hemionitis crispatula (Baker) Christenh., comb. nov. Acta Phytotax. Geobot. 25: 19. 1971, not: Hemionitis
Basionym: Pellaea crispatula Baker, Ann. Bot. delicatula (Maxon & Weath.) Christenh. Etymology:
(Oxford) 5: 215. 1891. From Latin delicatus, delightful.

Hemionitis delicatula (Maxon & Weath.) Christenh., Hemionitis ebenipes (X.C.Zhang) Christenh., comb.
comb. nov. Basionym: Notholaena delicatula Maxon nov. Basionym: Aleuritopteris ebenipes X.C.Zhang,
& Weath., Contr. Gray Herb. 127: 7. 1939. Acta Phytotax. Sin. 32: 94. 1994.
Hemionitis deltoidea (Kunze) Christenh., comb. nov. Hemionitis eckloniana (Kunze) Christenh., comb. nov.
Basionym: Cheilanthes deltoidea Kunze, Linnaea 10: Basionym: Notholaena eckloniana Kunze, Linnaea
536. 1836. 10: 501. 1836.
Hemionitis densa (Brackenr.) Christenh., comb. nov. Hemionitis ehrenbergiana (Klotzsch) Christenh., comb.
Basionym: Onychium densa Brackenr., U.S. Expl. nov. Basionym: Gymnogramma ehrenbergiana
Exped., Filic. 16: 120. 1854. Klotzsch, Linnaea 20: 411. 1847.
Hemionitis depauperata (Baker) Christenh., comb. Hemionitis ekmanii (Maxon) Christenh., comb. nov.
nov. Basionym: Cheilanthes depauperata Baker, Basionym: Notholaena ekmanii Maxon, Amer. Fern
Ann. Bot. (Oxford) 6: 210. 1891. J. 16: 9. 1926.
Hemionitis dichotoma (Sw.) Christenh., comb. nov. Hemionitis emperatricella Christenh., nom. nov.
Basionym: Cheilanthes dichotoma Sw., Syn. Filic. Replaced synonym: Pteris sagittata Cav., Descr. Pl.:
129: 335. 1806. 267. 1802, not: Hemionitis sagittata Fée. Etymology:
Hemionitis dilimanensis (Zamora) Christenh., comb. Named for the Child-like Empress (Moonchild), ruler
nov. Basionym: Cheilanthes dilimanensis Zamora, of Fantástica in The Neverending Story (Ende, 1979).
Kalikasan 3: 193. 1974. Hemionitis engywookii Christenh., nom. nov. Replaced
Hemionitis dinteri (Brause) Christenh., comb. nov. synonym: Notholaena fendleri Kunze, Farrnkr. 2: 87.
Basionym: Cheilanthes dinteri Brause, Bot. Jahrb. 1851. Etymology: Named for Engywook, a gnome-
Syst. 53: 385. 1915. scientist specialised in the Southern Oracle in The
Neverending Story (Ende, 1979).
Hemionitis distans (R.Br.) Christenh., comb. nov.
Basionym: Notholaena distans R.Br., Prod. Fl. Nov. Hemionitis eriophora (Fée) Christenh., comb. nov.
Holland.: 146. 1810. Basionym: Notholaena eriophora Fée, Mém. Foug.
5: 159. 1850.
Hemionitis dolomiticola (Schelpe) Christenh., comb.
nov. Basionym: Pellaea dolomiticola Schelpe, J. S. Hemionitis erythraea (Pichi-Serm.) Christenh., comb.
African Bot. 34: 239. 1968. nov. Basionym: Cheilanthes erythraea Pichi-Serm.,
Webbia 27: 420. 1972.
Hemionitis doniana (J.Sm. ex Hook.) Christenh., comb.
nov. Basionym: Pellaea doniana J.Sm. ex Hook., Sp. Hemionitis falkor Christenh., nom. nov. Replaced
Fil. 2: 137. 1858. synonym: Doryopteris feei Brade, Arq. Inst. Biol.
Veg. Rio de Janeiro 1: 226. 1935, not: Hemionitis
Hemionitis doryopteris Christenh., nom. nov. Replaced
feei (T.Moore) Christenh. Etymology: Named for
synonym: Pteris pedata L., Sp. Pl. 2: 1075. 1753, not:
Falkor (or Fuchur), the Luck Dragon, one of the main
Hemionitis pedata Sw. Etymology: Named for former
characters in The Neverending Story (Ende, 1979).
genus Doryopteris, now included in Hemionitis.
Hemionitis fantastica Christenh., nom. nov. Replaced
Hemionitis drakeana (Jeanp. ex C.Chr.) Christenh.,
synonym: Notholaena californica D.C.Eaton, Bull.
comb. nov. Basionym: Trachypteris drakeana Jeanp.
Torrey Bot. Club 10: 27. 1883. Etymology: Named
ex C.Chr. in Bonap., Notes Ptérid. 16: 194. 1925.
for Fantástica, the mythical land in The Neverending
Hemionitis dubia (Hope) Christenh., comb. nov. Story (Ende, 1979).
Basionym: Cheilanthes dubia Hope, J. Bombay Nat.
Hemionitis farinosa (Forssk.) Christenh., comb. nov.
Hist. Soc. 12: 528. 1899.
Basionym: Pteris farinosa Forssk., Fl. Aegypt.-Arab.:
Hemionitis duclouxii (Christ) Christenh., comb. nov. 187. 1775.
Basionym: Doryopteris duclouxii Christ, Bull. Acad.
Hemionitis feei (T.Moore) Christenh., comb. nov.
Int. Géogr. Bot. 11: 231. 1902.
Basionym: Cheilanthes feei T.Moore, Index Fil.: 38.
Hemionitis dura (Willd.) Christenh., comb. nov. 1857.
Basionym: Pteris dura Willd., Sp. Pl. ed. quarta: 376.
Hemionitis fendleri (Hook.) Christenh., comb. nov.
1810.
Basionym: Cheilanthes fendleri Hook., Sp. Fil. 2: 103.
Hemionitis duthiei (Baker) Christenh., comb. nov. 1852.
Basionym: Cheilanthes duthiei Baker, Ann. Bot.
Hemionitis flavescens (Fée) Christenh., comb. nov.
(Oxford) 5: 210. 1891.
Basionym: Pellaea flavescens Fée, Crypt. Vasc. Brésil
Hemionitis eatonii (Baker) Christenh., comb. nov. 1: 44. 1869.
Basionym: Cheilanthes eatonii Baker, Syn. Fil. 4: 140.
Hemionitis flexuosa (Kunze) Christenh., comb. nov.
1867.
Basionym: Cheilanthes flexuosa Kunze, Linnaea 22:
578. 1849.

13
Hemionitis formosa (Liebm.) Christenh., comb. nov. Hemionitis glanduloviscida (Schuettp. & Windham)
Basionym: Allosorus formosus Liebm., Kongel. Christenh., comb. nov. Basionym: Pentagramma
Dankse Vidensk. Selsk. Skr., Naturvidensk. Math. glanduloviscida Schuettp. & Windham, Syst. Bot. 40:
Afd., ser. 5, 1: 220. 1849. Replaced synonym: 639. 2015.
Allosorus pulchellus M.Martens & Galeotti, Nouv. Hemionitis glauca (Cav.) Christenh., comb. nov.
Mém. Acad. Roy. Sci. Bruxelles 15: 47. 1842, not: Basionym: Pteris glauca Cav., Descr. Pl.: 269. 1802.
Hemionitis pulchella (Bory ex Willd.) Christenh. Hemionitis gleichenioides (Gardner) Christenh., comb.
Hemionitis formosana (Hayata) Christenh. comb. nov. nov. Basionym: Cassebeera gleichenioides Gardner,
Basionym: Cheilanthes formosana Hayata, Enum. Pl. Icon. Pl. 6: t. 507. 1843.
Form.: 612. 1906. Hemionitis glutinosa (M.Kessler & A.R.Sm.) Christenh.,
Hemionitis fractifera (R.M.Tryon) Christenh., comb. comb. nov. Basionym: Cheilanthes glutinosa
nov. Basionym: Cheilanthes fractifera R.M.Tryon, M.Kessler & A.R.Sm., Brittonia 59: 188. 2007.
Rhodora 62: 7. 1960. Hemionitis gmorgana Christenh., nom. nov. Replaced
Hemionitis fragilis (Hook.) Christenh., comb. nov. synonym: Cheilanthes rufa D.Don, Prod. Fl. Nepal.:
Basionym: Cheilanthes fragilis Hook., Filic. Exot.: t. 16. 1825, not: Hemionitis rufa Sw. Etymology:
96. 1859. Named for Gmorg a wolf-like servant of The Nothing
Hemionitis fragillima (F.Muell) Christenh, comb. nov. in The Neverending Story (Ende, 1979).
Basionym: Cheilanthes fragillima F.Muell., Fragm. Hemionitis goabica Christenh., nom. nov. Replaced
Phyt. Austr. 5: 126. 1866. Replaced synonym: synonym: Pteris cordifolia Baker, J. Bot. 29: 4. 1891,
Notholaena fragilis Hook., Sp. Fil. 5: 114. 1864, not: not Hemionitis cordifolia (Sessé & Moc.) Christenh.,
Hemionitis fragilis (Hook.) Christenh. nor H. cordifolia Roxb. ex Wall, nom. inval., nom.
Hemionitis fraseri (Mett. ex Kuhn) Christenh., comb. nud. Etymology: Named for Goab, the Desert of
nov. Basionym: Cheilanthes fraseri Mett. ex Kuhn, Colours in The Neverending Story (Ende, 1979).
Linnaea 36: 83. 1869. Hemionitis goyazensis (Taub.) Christenh., comb. nov.
Hemionitis gaga Christenh., nom. nov. Replaced Basionym: Notholaena goyazensis Taub., Bot. Jahrb.
synonym: Allosorus chaerophyllus M.Martens & Syst. 21: 421. 1896.
Galeotti, Nouv. Mém. Acad. Roy. Sci. Bruxelles 15: 47. Hemionitis gracillima (D.C.Eaton) Christenh., comb.
1842, not: Hemionitis chaerophylla Poir. Etymology: nov. Basionym: Cheilanthes gracillima D.C.Eaton in
Named for the former genus Gaga, now a synonym Emory, Rep. U.S. Mex. Bound. 2: 234. 1859.
of Hemionitis. Hemionitis grandis Christenh., nom. nov. Replaced
Hemionitis galapagensis (Weath. & Svenson) synonym: Aleuritopteris gigantea Saiki, J. Phytogeogr.
Christenh., comb. nov. Basionym: Notholaena Taxon. 32: 86. 1984, not: Hemionitis gigantea Desv.
galapagensis Weath. & Svenson, Bull. Torrey Bot. Etymology: Named for its large size.
Club 65: 319. 1938. Hemionitis graograman Christenh., nom. nov. Replaced
Hemionitis galeottii (Fée) Christenh., comb. nov. synonym: Cheilanthes pilosa Goldm., Nov. Act. Acad.
Basionym: Notholaena galeottii Fée, Mém. Foug. 5: Caes. Leop.-Carol. Nat. Cur. 19 (suppl. 1): 455. 1843,
159. 1852. not: Hemionitis pilosa (Poir.) Christenh. Etymology:
Hemionitis gastonyi (Windham) Christenh., comb. Named for Graograman, the Many-Coloured Death
nov. Basionym: Pellaea gastonyi Windham, Contr. in The Neverending Story (Ende, 1979).
Univ. Michigan Herb. 19: 36. 1993. Hemionitis grayii (Davenp.) Christenh., comb. nov.
Hemionitis geraniifolia (Weath.) Christenh., comb. Basionym: Notholaena grayi Davenp. in D.C.Eaton,
nov. Basionym: Cheilanthes geraniifolia Weath., J. Ferns N. Amer. 2: 31. 1879.
Arnold Arbor. 27: 367. 1946. Hemionitis greggii (Mett. ex Kuhn) Christenh., comb.
Hemionitis germanotta (Fay W. Li & Windham) nov. Basionym: Pellaea greggii Mett. ex Kuhn,
Christenh., comb. nov. Basionym: Gaga germanotta Linnaea 36: 86. 1869.
Fay W.Li & Windham, Syst. Bot. 37: 857. 2012. Hemionitis grevilleoides (Christ) Christenh., comb.
Hemionitis glabella (Mett. ex Kuhn) Christenh., comb. nov. Basionym: Cheilanthes grevilleoides Christ.
nov. Basionym: Pellaea glabella Mett. ex Kuhn, Notul. Syst. (Paris) 1: 51. 1909.
Linnaea 36: 87. 1869. Hemionitis grisea (Blanf.) Christenh., comb. nov.
Hemionitis glaberrima (Fée) Christenh., comb. nov. Basionym: Cheilanthes grisea Blanf., J. Simla Natural.
Basionym: Cheilanthes glaberrima Fée, Crypt. Casc. Soc. 1: 21. 1886.
Brésil 1: 54. 1869. Hemionitis ×gryphus (Mickel) Christenh., comb. nov.
Basionym: Cheilanthes ×gryphus Mickel, Amer. Fern
J. 77: 112. 1987.

Hemionitis guanchica (Bolle) Christenh., comb. nov. Hemionitis hispida (Mett. ex Kuhn) Christenh., comb.
Basionym: Cheilanthes guanchica Bolle, Bonplandia nov. Basionym: Gymnogramma hispida Mett. ex
7: 107. 1859. Kuhn, Linnaea 36: 72. 1869. (Figure 3D)
Hemionitis gluckuk Christenh., nom. nov. Replaced Hemionitis hookeri (E.J.Lowe) Christenh., comb. nov.
synonym: Notholaena subcordata Desv., J. Bot. Agric. Basionym: Notholaena hookeri E.J.Lowe, Ferns 1:
1: 92. 1813, not: Hemionitis subcordata (D.C.Eaton & 59. 1856. Replaced synonym: Notholaena standleyi
Davenp.) Mickel. Etymology: Named for Gluckuk, a Maxon, Amer. Fern J. 5: 1. 1915.
winzling riding a fast snail in The Neverending Story Hemionitis hopeana (C.Chr.) Christenh., comb. nov.
(Ende, 1979). Basionym: Cheilanthes hopeana C.Chr., Ind. Filic.,
Hemionitis gymnocarpa (Ching) Christenh., comb. Suppl. 1: 18. 1913. Replaced synonym: Pellaea
nov. Basionym: Aleuritopteris gymnocarpa Ching squamosa C.Hope & C.H.Wright, not: Hemionitis
in Y.L.Chang et al., Sporae Pterid. Sin.: 158. 1976. squamosa (Gillies ex Hook. & Grev.) Christenh.
Replaced synonym: Aleuritopteris doniana S.K.Wu, Hemionitis horokana Christenh., nom. nov. Replaced
nom. illeg. superfl., Cheilanthes dealbata D.Don, synonym: Pellaea intermedia Mett. ex Kuhn, Linnaea
nom. illeg., not: Hemionitis dealbata Willd., nor 38: 84. 1869, not: Hemionitis intermedia Fée.
Hemionitis doniana (J.Sm.) Christenh. Etymology: Named for Horok Castle, the Allseeing
Hemionitis hancockii (Baker) Christenh., comb. nov. Hand in The Neverending Story (Ende, 1979).
Basionym: Cheilanthes hancockii Baker, Bull. Misc. Hemionitis horridula (Maxon) Christenh., comb. nov.
Inf. Kew 1895(99): 54. 1895. Basionym: Cheilanthes horridula Maxon, Amer. Fern
Hemionitis harrisii (Maxon) Christenh., comb. nov. J. 8: 94. 1918.
Basionym: Cheilanthes harrisii Maxon, Contr. U.S. Hemionitis humatoides (Saiki) Christenh., comb.
Natl. Herb. 24: 51. 1922. nov. Basionym: Aleuritopteris humatoides Saiki, J.
Hemionitis harrisonae (Jenm.) Christenh., comb. nov. Phytogeogr. Taxon. 32: 5. 1984.
Basionym: Pteris harrisonae Jenm., Gard. Chron. ser. Hemionitis humbertii (Tardieu) Christenh., comb.
3, 24: 414. 1898. nov. Basionym: Doryopteris humbertii Tardieu, Fl.
Hemionitis hassleri (Weath.) Christenh., comb. nov. Madagasc. 5(1): 144. 1958.
Basionym: Notholaena hassleri Weath., Lilloa 6: 274. Hemionitis ×hybrida (Brade & Rosenst.) Christenh.,
1941. comb. nov. Basionym: Doryopteris ×hybrida Brade &
Hemionitis hickeyi (Link-Pérez, Seabolt & Ledford) Rosenst., Repert. Sp. Nov. Regni Veg. 21: 346. 1925.
Christenh., comb. nov. Basionym: Adiantopsis Hemionitis hydornii Christenh., nom. nov. Replaced
hickeyi Link-Pérez, Seabolt & Ledford, Syst. Bot. 41: synonym: Pellaea tomentosa Bonap., Notes Ptértid.
915. 2016. 5: 64. 1917, not: Hemionitis tomentosa Raddi.
Hemionitis hieronymi (Herter) Christenh., comb. nov. Etymology: Named for one of the four heroes in The
Basionym: Cheilanthes hieronymi Herter, Anal. Mus. Neverending Story (Ende, 1979).
Hist. Nat. Montevideo 1: 360. 1925. Hemionitis hykrionii Christenh., nom. nov. Replaced
Hemionitis himalaica (Fraser-Jenk.) Christenh., comb. synonym: Pellaea prolifera Schelpe, Bull. Jard. Bot.
nov. Basionym: Notholaena himalaica Fraser-Jenk., Natl. Belg. 38: 193. 1968, not: Hemionitis prolifera
New Sp. Syndr. Indian Pteridol. & Ferns Nepal: 183. Retz. Etymology: Named for one of the four heroes
1997. Replaced synonym: Gymnogramma vestita in The Neverending Story (Ende, 1979).
Hook., not: Hemionitis vestita (Desv.) Christenh. Hemionitis hynreckii Christenh., nom. nov. Replaced
(Figure 3C) synonym: Gymnogramma muelleri Hook., Sp.
Hemionitis hintoniorum (Mendenh. & G.L.Nesom) Filic. 5: 143. 1864, not: Hemionitis muelleri J.Sm.
Christenh., comb. nov. Basionym: Cheilanthes Etymology: Named for one of the four heroes in The
hintoniorum Mendenh. & G.L.Nesom, Sida 14: 551. Neverending Story (Ende, 1979).
1991. Hemionitis hypoleuca (Kunze) Christenh., comb. nov.
Hemionitis hirsuta (Link) Christenh., comb. nov. Basionym: Notholaena hypoleuca Kunze, Linnaea 9:
Basionym: Cheilanthes hirsuta Link, Hort. Berol. 2: 54. 1834.
40. 1833. Hemionitis hysbaldii Christenh., nom. nov. Replaced
Hemionitis hirta (Sw.) Christenh., comb. nov. Basionym: synonym: Pellaea pringlei Davenp., Gard. and Forest
Cheilanthes hirta Sw., Syn. Filic.: 128, 329. 1806. 4: 555. 1891, not: Hemionitis pringlei (Davenp.)
Hemionitis hispanica (Mett.) Christenh., comb. nov. Christenh. Etymology: Named for one of the four
Basionym: Cheilanthes hispanica Mett., Abh. heroes in The Neverending Story (Ende, 1979).
Senckenberg. Naturf. Ges. 3: 74. 1859. Hemionitis ilwan Christenh., nom. nov. Replaced

15
synonym: Saffordia induta Maxon, Rhodora 83: 137. Hemionitis keralensis (N.C.Nair & S.R.Ghosh)
1981, not: Hemionitis induta (Kunze) Christenh. Christenh., comb. nov. Basionym: Cheilanthes
Etymology: Named for Ilwan, a blue genie in The keralensis N.C.Nair & S.R.Ghosh, J. Indian Bot. Soc.
Neverending Story (Ende, 1979). 55: 52. 1976.
Hemionitis inaequalis (Kunze) Christenh., comb. nov. Hemionitis kitchingii (Baker) Christenh., comb. nov.
Basionym: Notholaena inaequalis Kunze, Farrnkr. 1: Basionym: Pellaea kitchingii Baker, J. Bot. 18: 327.
146. 1844. 1880.
Hemionitis incana (C.Presl) Christenh., comb. nov. Hemionitis krameri (Franch. & Sav.) Christenh., comb.
Basionym: Notholaena incana C.Presl, Reliq. Haenk. nov. Basionym: Cheilanthes krameri Franch. & Sav.,
1: 19. 1825. Enum. Pl. Jap. 2: 212, 619. 1879.
Hemionitis incarum (Maxon) Christenh., comb. nov. Hemionitis kuhnii (Milde) Christenh., comb. nov.
Basionym: Cheilanthes incarum Maxon, Smithsonian Basionym: Cheilanthes kuhnii Milde, Bot. Zeitung
Misc. Coll. 65(8): 5. 1915. (Berlin) 25: 149. 1867.
Hemionitis induta (Kunze) Christenh., comb. nov. Hemionitis kunzei (Mett.) Christenh., comb. nov.
Basionym: Cheilanthes induta Kunze., Linnaea 10: Basionym: Cheilanthes kunzei Mett., Abh.
538. 1836. Senckenberg. Naturf. Gesell. 3: 71. 1859.
Hemionitis insignis (Ching) Christenh., comb. nov. Hemionitis laciniata (Sodiro) Christenh., comb. nov.
Basionym: Cheilanthes insignis Ching, Fl. Tsinling. 2: Basionym: Cheilanthes laciniata Sodiro, Anal. Univ.
208. 1974. Centr. Ecuador 7: 189. 1892.
Hemionitis integerrima (Hook.) Christenh., comb. & Hemionitis lanosa (Michx.) Christenh., comb. nov.
stat. nov. Basionym: Notholaena sinuata (Lag. ex Basionym: Nephrodium lanosum Michx., Fl. Bor.-
Sw.) Kaulf. var. integerrima Hook., Sp. Filic. 5: 108. Amer. 2: 270. 1803.
1864. Hemionitis lanuginosa (Desv. ex Poir.) Christenh.,
Hemionitis intertexta (Maxon) Christenh., comb. comb. nov. Basionym: Notholaena lanuginosa Desf.
nov. Basionym: Cheilanthes colvillei Maxon subsp. ex Poir. in Lam., Encycl. Meth. Bot. Suppl. 4: 110.
intertexta Maxon, Proc. Biol. Soc. Washington 31: 1816.
149. 1918. Hemionitis latiloba (C.Chr.) Christenh., comb. nov.
Hemionitis intramarginalis (Kaulf. ex Link) Christenh., Basionym: Doryopteris latiloba C.Chr., Arch. Bot.
comb. nov. Basionym: Pteris intramarginalis Kaulf. (Caen) 2, Bull. Mens.: 213. 1928.
ex Link, Hort. Berol. 2: 34. 1833. Hemionitis leachii (Schelpe) Christenh., comb. nov.
Hemionitis itatiaiensis (Fée) Christenh., comb. nov. Basionym: Notholaena leachii Schelpe, J. S. Afr. Bot.
Basionym: Pellaea itatiaiensis Fée, Crypt. Vasc. Brésil 30: 185, f. la (1964)
2: 26. 1872. Hemionitis lemmonii (D.C.Eaton) Christenh., comb.
Hemionitis jacalensis (Pray) Christenh., comb. nov. nov. Basionym: Notholaena lemmonii D.C.Eaton,
Basionym: Notholaena jacalensis Pray, Amer. Fern J. Bull. Torrey Bot. Club 7: 63. 1880.
57: 101. 1967. Hemionitis lendigera (Cav.) Christenh., comb. nov.
Hemionitis jaliscana (Yatsk. & Arbeláez) Christenh., Basionym: Pteris lendigera Cav., Descr. Pl.: 268. 1802.
comb. nov. Basionym: Notholaena jaliscana Yatsk. & Hemionitis leonardii (Maxon) Christenh., comb.
Arbeláez, Novon 18: 121. 2008. Replaced synonym: nov. Basionym: Cheilanthes leonardii Maxon, J.
Cheilanthes palmeri D.C.Eaton, Proc. Amer. Acad Washington Acad. Sci. 14: 87. 1924.
Arts & Sci. 22: 464. 1887, not Hemionitis palmeri Hemionitis leonina (Maxon) Christenh., comb. nov.
(Baker) Christenh. Basionym: Notholaena leonina Maxon, Contr. U.S.
Hemionitis jamaicensis (Maxon) Christenh., comb. Natl. Herb. 16: 58. 1912.
nov. Basionym: Cheilanthes jamaicensis Maxon, Hemionitis leptolepis (Fraser-Jenk.) Christenh., comb.
Contr. U.S. Nat. Herb. 24: 51. 1922. nov. Basionym: Cheilanthes leptolepis Fraser-Jenk.,
Hemionitis jequitinhonhensis (Salino) Christenh., Bot. Helv. 102: 144. 1992.
comb. nov. Basionym: Doryopteris jequitinhonhensis Hemionitis lerstenii (Mickel & Beitel) Christenh., comb.
Salino, J. Bot. Res. Inst. Texas 2: 843. 2008. nov. Basionym: Cheilanthes lerstenii Mickel & Beitel,
Hemionitis jonesii (Maxon) Christenh., comb. nov. Mem. New York Bot. Gard. 46: 117. 1988.
Basionym: Notholaena jonesii Maxon, Amer. Fern J. Hemionitis leucopoda (Link) Christenh., comb. nov.
7: 108. 1917. Basionym: Cheilanthes leucopoda Link, Filic. Sp.: 66.
Hemionitis kaulfussii (Kunze) Christenh., comb. nov. 1841.
Basionym: Cheilanthes kaulfussii Kunze, Linnaea 13:
145. 1839.

Hemionitis likiangensis (Ching) Christenh., comb. nov. Hemionitis maderensis (Lowe) Christenh., comb. nov.
Basionym: Aleuritopteris likiangensis Ching, Acta Basionym: Cheilanthes maderensis Lowe, Trans.
Phytotax. Sin. 19: 64. 1981. Cambridge Philos. Soc. 6: 528. 1838.
Hemionitis limitanea (Maxon) Christenh., comb. nov. Hemionitis magdalenensis (Brade) Christenh., comb.
Basionym: Notholaena limitanea Maxon, Amer. Fern & stat. nov. Basionym: Doryopteris subsimplex var.
J. 9: 70. 1919. magdalenensis Brade., Arq. Inst. Biol. Veg. 2: 3. 1935.
Hemionitis lindheimeri (Hook.) Christenh., comb. nov. Hemionitis mairei (Brause) Christenh., comb. nov.
Basionym: Cheilanthes lindheimeri Hook., Sp. Filic. 2: Basionym: Pellaea mairei Brause, Hedwigia 54: 201.
101. 1852. 1914.
Hemionitis lindigii (Mett.) Christenh., comb. nov. Hemionitis majestosa (Yesilyurt) Christenh., comb.
Basionym: Cheilanthes lindigii Mett., Ann. Sci. Nat., nov. Basionym: Doryopteris majestosa Yesilyurt,
Bot., sér 5, 2: 218. 1864. Amer. Fern J. 97: 212. 2008.
Hemionitis linearis (Bonap.) Christenh., comb. nov. Hemionitis malabarica (Madhus. & Jyothi ex
Basionym: Adiantopsis linearis Bonap., Notes Ptérid. Geev.) Christenh., comb. nov. Basionym: Pellaea
10: 185. 1920. malabarica Madhus. & Jyothi ex Geev., Indian Fern
Hemionitis lomariacea (Klotzsch) Christenh., comb. J. 9: 39. 1992.
nov. Basionym: Doryopteris lomariacea Klotzsch, Hemionitis marantae (L.) Christenh., comb. nov.
Linnaea 20: 343. 1847. Basionym: Acrostichum marantae L., Sp. Pl. 2: 1071.
Hemionitis lonchophora (Römer ex Mett.) Christenh., 1753.
comb. nov. Basionym: Pteris lonchophora Römer ex Hemionitis marginata (Kunth) Christenh., comb. nov.
Mett., Abh. Senckenberg. Naturf. Gesell. 3: 48. 1859. Basionym: Cheilanthes marginata Kunth, Nov. Gen.
Hemionitis lonchophylla (R.M.Tryon) Christenh., comb. Sp. (quarto ed.) 1: 22. 1815.
nov. Basionym: Notholaena lonchophylla R.M.Tryon, Hemionitis marlothii (Hieron.) Christenh., comb. nov.
Contr. Gray Herb. 179: 19. 1956. Basionym: Notholaena marlothii Hieron., Bot. Jahrb.
Hemionitis longipila (Baker) Christenh., comb. nov. Syst. 46: 384. 1911.
Basionym: Cheilanthes longipila Baker, Ann. Bot. Hemionitis marsupianthes (Fée) Christenh., comb.
(Oxford) 5. 211. 1891. nov. Basionym: Myriopteris marsupianthes Fée,
Hemionitis longipilosa (Bonap.) Christenh., comb. nov. Mém. Foug. 5: 149. 1852.
Basionym: Pellaea longipilosa Bonap., Notes Ptérid. Hemionitis maxoniana (Mickel) Christenh., comb. nov.
16: 33. 1924. Basionym: Cheilanthes maxoniana Mickel, Mem.
Hemionitis lorentzii (Hieron.) Christenh., comb. nov. New York Bot. Gard. 88: 201. 2004.
Basionym: Pellaea lorentzii Hieron., Bot. Jahrb. Syst. Hemionitis meifolia (D.C.Eaton) Christenh., comb. nov.
22: 392. 1896. Basionym: Cheilanthes meifolia D.C.Eaton, Proc.
Hemionitis lozanoi (Maxon) Christenh., comb. nov. Amer. Acad. Arts & Sci. 18: 185. 1883.
Basionym: Pellaea lozanoi Maxon, Contr. U.S. Natl. Hemionitis membranacea (Davenp.) Christenh., comb.
Herb. 10: 500. 1908, as ’lozani’. nov. Basionym: Pellaea membranacea Davenp., Bot.
Hemionitis ludens (Wall. ex Hook.) Christenh., comb. Gaz. 21: 262. 1896.
nov. Basionym: Pteris ludens Wall. ex Hook., Sp. Filic. Hemionitis mexicana (Davenp.) Christenh., comb.
2: 210. 1858. nov. Basionym: Cheilanthes mexicana Davenp., Bull.
Hemionitis luetzelburgii (Rosenst.) Christenh., comb. Torrey Bot. Cl. 15: 227. 1888.
nov. Basionym: Adiantopsis luetzelburgii Rosenst., Hemionitis michelii (Christ) Christenh., comb. nov.
Repert. Sp. Nov. Regni Veg. 20: 91. 1924. Basionym: Doryopteris michelii Christ, Bull. Acad. Int.
Hemionitis lumholtzii (Maxon & Weath.) Christenh., Géogr. Bot. 20: 14. 1910.
comb. nov. Basionym: Notholaena lumholtzii Maxon Hemionitis mickelii (T.Reeves) Christenh., comb. nov.
& Weath., Contr. Gray Herb. 127: 16. 1939. Basionym: Cheilanthes mickelii T.Reeves, Brittonia
Hemionitis lutea (Cav.) Christenh., comb. nov. 32: 502. 1980.
Basionym: Pteris lutea Cav., Descr. Pl.: 267. 1802. Hemionitis microphylla (Sw.) Christenh., comb. nov.
Hemionitis lyngholmii (Windham) Christenh., comb. Basionym: Adiantum microphyllum Sw., Prodr.: 135.
nov. Basionym: Pellaea lyngholmii Windham, Contr. 1788.
Univ. Michigan Herb. 19: 40. 1993. Hemionitis micropteris (Sw.) Christenh., comb. nov.
Hemionitis madagascariensis (Baker) Christenh., Basionym: Cheilanthes micropteris Sw., Syn. Filic.:
comb. nov. Basionym: Cheilanthes madagascariensis 126. 1806.
Baker, J. Linn. Soc., Bot. 16: 198. 1877.

17
Hemionitis minutula (Sehnem) Christenh., comb. nov. Bull. Torrey Bot. Club 4: 12. 1873.
Basionym: Adiantopsis minutula Sehnem, Fl. Ilustr. Hemionitis nielsii (W.Jacobsen) Christenh., comb. nov.
Catar. 1 (Pteridac.): 73. 1972. Basionym: Cheilanthes nielsii W.Jacobsen, Ferns &
Hemionitis mollis (Kunze) Christenh., comb. nov. Fern Allies S. Africa: 252. 1983.
Basionym: Notholaena mollis Kunze, Linnaea 9: 54. Hemionitis nigricans (Willd.) Christenh., comb. nov.
1834. Basionym: Pteris nigricans Willd., Sp. Pl., ed. 4, 5:
Hemionitis moncloviensis (Baker) Christenh., comb. 396. 1810.
nov. Basionym: Cheilanthes moncloviensis Baker, Hemionitis niphobola (C.Chr.) Christenh., comb. nov.
Ann. Bot. (Oxford) 5: 210. 1891. Basionym: Cheilanthes niphobola C.Chr., Acta Horti
Hemionitis monstraparva (Fay W.Li & Windham) Gothob. 1: 88. 1924.
Christenh., comb. nov. Basionym: Gaga Hemionitis nitida (R.Br.) Christenh., comb. nov.
monstraparva Fay W.Li & Windham, Syst. Bot. 37: Basionym: Pteris nitida R.Br., Prodr. Fl. Nov. Holland.:
858. 2012. 155. 1810.
Hemionitis monticola (Gardner) Christenh., comb. Hemionitis nitidula (Hook.) Christenh., comb. nov.
nov. Basionym: Cheilanthes monticola Gardner, Basionym: Cheilanthes nitidula Hook., Sp. Filic. 2:
Icon. Pl. 5: t. 487. 1842. 112. 1852.
Hemionitis moritziana (Kunze) Christenh., comb. nov. Hemionitis nivea (Poir.) Christenh., comb. nov.
Basionym: Cheilanthes moritziana Kunze, Linnaea Basionym: Pteris nivea Poir. in Lam. Encycl. Méth.,
23: 307. 1850. Bot. 5: 718. 1804.
Hemionitis morla Christenh., nom. nov. Replaced Hemionitis nobilis (T.Moore) Christenh., comb. nov.
synonym: Cheilanthes coriacea Decne., Arch. Mus. Basionym: Litobrochia nobilis T.Moore, Gard. Chron.
Hist. Nat. 2: 190. 1841, not: Hemionitis coriacea 1862(40): 932. 1862.
D.Don. Etymology: Named for the Ancient Morla, a Hemionitis notabilis (Maxon) Christenh., comb. nov.
giant terrapin in The Neverending Story (Ende, 1979) Basionym: Pellaea notabilis Maxon, Contr. U.S. Natl.
Hemionitis mucronata (D.C.Eaton) Christenh., comb. Herb. 10: 500. 1908.
nov. Basionym: Allosorus mucronatus D.C.Eaton, Hemionitis notholaenoides (Desv.) Christenh., comb.
Amer, J. Sci & Arts, ser. 2, 22: 138. 1856. nov. Basionym: Pteris notholaenoides Desv., Mém.
Hemionitis multifida (Sw.) Christenh., comb. nov. Soc. Linn. Paris 6: 299. 1827.
Basionym: Adiantum multifidum Sw., J. Bot. Hemionitis nuda (Ching) Christenh., comb. nov.
(Schrader) 1800(2): 85. 1801. Basionym: Aleruitopteris nuda Ching, Fl. Tsinling. 2:
Hemionitis multipartita (Fée) Christenh., comb. & stat. 207. 1974.
nov. Basionym: Doryopteris raddiana (C.Presl) Fée Hemionitis nudiuscula (R.Br.) Christenh., comb. nov.
var. multipartita Fée, Crypt. Vasc. Brésil 1: 45. 1869. Basionym: Pteris nudiuscula R.Br., Prodr. Fl. Nov.
Hemionitis myriophylla (Desv.) Christenh., comb. nov. Holland.: 155. 1810.
Basionym: Cheilanthes myriophylla Desv., Mag. Hemionitis obducta (Mett. ex Kuhn) Christenh., comb.
Neuesten Entdeck. Gesammten Naturk., Gesell. nov. Basionym: Cheilanthes obducta Mett. ex Kuhn,
Naturf. Freunde Berlin 5. 328. 1811. Linnaea 36: 83. 1869.
Hemionitis myrtillifolia (Mett. ex Kuhn) Christenh., Hemionitis occulta (Sehnem) Christenh., comb. nov.
comb. nov. Basionym: Pellaea myrtillifolia Mett. ex Basionym: Adiantopsis occulta Sehnem, Pesquisas,
Kuhn, Linnaea 36: 85. 1869. Bot. 3: 508. 1959.
Hemionitis namaquensis (Baker) Christenh., comb. Hemionitis opposita (Kaulf.) Christenh., comb. nov.
nov. Basionym: Pellaea namaquensis Baker, J. Bot. Basionym: Cheilanthes opposita Kaulf., Enum. Filic.:
12: 199. 1874. 211. 1824.
Hemionitis nana (Hook.) Christenh., comb. & stat. nov. Hemionitis orbignyana (Mett. ex Kuhn) Christenh.,
Basionym: Pellaea falcata Fée var. nana Hook., Sp. comb. nov. Basionym: Cheilanthes orbignyana Mett.
Filic. 2: 136. 1858. ex Kuhn, Linnaea 36: 82. 1869.
Hemionitis nealleyi (Seaton ex J.M.Coult.) Christenh., Hemionitis ornithopus (Mett.) Christenh., comb. nov.
comb. nov. Basionym: Notholaena nealleyi Seaton Basionym: Pteris ornithopus Mett., Syn. Filic.: 166.
ex J.M.Coult., Contr. U.S. Natl. Herb. 1: 61. 1890. 1867.
Hemionitis neglecta (Maxon) Christenh., comb. nov. Hemionitis ovata (Desv.) Christenh., comb. nov.
Basionym: Notholaena neglecta Maxon, Contr. U.S. Basionym: Pteris ovata Desv., Mém. Soc. Linn. Paris
Natl. Herb. 17: 602. 1916. 6: 301. 1827.
Hemionitis newberryi (D.C.Eaton) Christenh., comb.
nov. Basionym: Notholaena newberryi D.C.Eaton,

Hemionitis pallens (Weath.) Christenh., comb. nov. de Janeiro, Bot. 6: 94. 1929, not: Hemionitis rufa
Basionym: Notholaena pallens Weath., Contr. Gray Sw. Etymology: Perelin, the Night Forest in The
Herb. 179: 78. 1956. Neverending Story (Ende, 1979).
Hemionitis pallida (Baker) Christenh., comb. nov. Hemionitis perfasciculata (Sehnem) Christenh., comb.
Basionym: Pellaea pallida Baker, Syn. Filic. (ed. 2) 2: nov. Basionym: Adiantopsis perfasciculata Sehnem,
478. 1874. Pesquisas, Bot. 13: 21. 1961.
Hemionitis palmeri (Baker) Christenh., comb. nov. Hemionitis perlanata (Pic.Serm.) Christenh., comb.
Basionym: Notholaena palmeri Baker, Hooker’s Icon. nov. Basionym: Notholaena perlanata Pic.-Serm.,
Pl. 17: t. 1678. 1887. Webbia 27: 417. 1972.
Hemionitis pantanalensis (E.L.M.Assis, Ponce & Hemionitis perrieri (J.P.Roux) Christenh., comb. nov.
Labiak) Christenh., comb. nov. Cheilanthes Basionym: Cheilanthes perrieri J.P.Roux, Bothalia 40:
pantanalensis E.L.M.Assis, Ponce & Labiak, Amer. 81. 2010. Replaced synonym: Notholaena lanceolata
Fern J. 98: 203. 2009. Bonap., Notes Ptérid. 5: 65. 1917, not: Hemionitis
Hemionitis papuana (C.Chr.) Christenh., comb. nov. lanceolata L.
Basionym: Cheilanthes papuana C.Chr., Dansk Bot. Hemionitis persica (Bory) Christenh., comb. nov.
Ark. 9: 57. 1937. Basionym: Notholaena persica Bory, Voy. Indes
Hemionitis paradoxa (R.Br.) Christenh., comb. nov. Orient.: 23. 1833.
Basionym: Adiantum paradoxum R.Br., Prodr. Fl. Hemionitis peruviana (Desv.) Christenh., comb. nov.
Nov. Holland. 1: 155. 1810. Basionym: Notholaena peruviana Desv., Mém. Soc.
Hemionitis parryi (D.C.Eaton) Christenh., comb. nov. Linn. Paris 6: 220. 1827.
Basionym: Notholaena parryi D.C.Eaton, Amer. Hemionitis pilifera (R.M.Tryon) Christenh., comb. nov.
Naturalist 9: 351. 1875. Basionym: Notholaena pilifera R.M.Tryon, Contr.
Hemionitis parviloba (Sw.) Christenh., comb. nov. Gray Herb. 179: 79. 1956.
Basionym: Adiantum parvilobum Sw., J. Bot. Hemionitis pilosa (Poir.) Christenh., comb. nov.
(Schrader) 1800(2): 85. 1801. Basionym: Pteris pilosa Poir. in Lam., Encycl. Méth.,
Hemionitis parvisegmenta (M.S.Barker & Hickey) Bot. 5: 717. 1804.
Christenh., comb. nov. Basionym: Adiantopsis Hemionitis platychlamys (Ching) Christenh., comb.
parvisegmenta M.S.Barker & Hickey, Ann. Missouri nov. Basionym: Aleuritopteris platychlamys Ching,
Bot. Gard. 93: 388. 2006. Fl. Tsinling. 2: 207. 1974, as ‘platychalamys’.
Hemionitis patula (Baker) Christenh., comb. nov. Hemionitis poeppigiana (Mett. ex Kuhn) Christenh.,
Basionym: Cheilanthes patula Baker, J. Bot. 26: 225. comb. nov. Basionym: Cheilanthes poeppigiana
1888. Mett. ex Kuhn, Linnaea 36: 84. 1869.
Hemionitis paupercula (Kunze) Christenh., comb. nov. Hemionitis pohliana (Mett.) Christenh., comb.
Basionym: Adiantum pauperculum Kunze, Farrnkr. 2: nov. Basionym: Cheilanthes pohliana Mett., Abh.
65. 1850. Senckenberg. Naturf. Gesell. 3: 67. 1859.
Hemionitis pectiniformis (Baker) Christenh., comb. Hemionitis praetermissa (D.L.Jones) Christenh., comb.
nov. Basionym: Pellaea pectiniformis Baker, Syn. nov. Basionym: Cheilanthes praetermissa D.L.Jones,
Filic. (ed. 2): 147. 1874. Austrobaileya 2: 472. 1988.
Hemionitis pedatoides (Desv.) Christenh., comb. nov. Hemionitis prenticei (Luerss.) Christenh., comb.
Basionym: Pteris pedatoides Desv., Mém. Soc. Linn. nov. Basionym: Cheilanthes prenticei Luerss., Bot.
Paris 6: 293. 1827. Centralbl. 3: 442. 1882.
Hemionitis pellaea Christenh., nom. nov. Replaced Hemionitis pringlei (Davenp.) Christenh., comb. nov.
synonym: Pteris falcata R.Br., Prod. Fl. Nov. Holland.: Basionym: Cheilanthes pringlei Davenp., Bull. Torrey
154. 1810, not: Hemionitis falcata Willd., nor: Pteris Bot. Club 10: 61. 1883.
falcata Sodiro, nom. illeg. Etymology: Named for the Hemionitis pruinata (Kaulf.) Christenh., comb. nov.
former genus Pellaea, now included in Hemionitis. Basionym: Cheilanthes pruinata Kaulf., Enum. Filic.:
Hemionitis pellaeopsis (Mickel) Christenh., comb. 210. 1824.
nov. Basionym: Cheilanthes pellaeopsis Mickel in Hemionitis pteridioides (Reichard) Christenh., comb.
McVaugh, Fl. Novo-Galiciana 17: 241. 1992. nov. Basionym: Polypodium pteridioides Reichard,
Hemionitis peninsularis (Maxon) Christenh., comb. Syst. Pl. 4: 424. 1780.
nov. Basionym: Cheilanthes peninsularis Maxon, Hemionitis pteroides (L.) Christenh., comb. nov.
Contr. U.S. Natl. Herb. 10: 496. 1908. Basionym: Adiantum pteroides L., Mant. Pl.: 130.
Hemionitis perelinae Christenh., nom. nov. Replaced 1767.
synonym: Doryopteris rufa Brade, Bol. Mus. Nac. Rio

19
Hemionitis pulchella (Bory ex Willd.) Christenh., comb. Hemionitis reesii (Jenman) Christenh., comb. nov.
nov. Basionym: Cheilanthes pulchella Bory ex Willd., Basionym: Cheilanthes reesii Jenman, J. Bot. 24:
Sp. Pl. (ed. quarta) 5: 456. 1810, not: Allosorus 267. 1886.
pulchellus M.Martens & Galeotti Hemionitis regnelliana (Mett.) Christenh., comb. nov.
Hemionitis pumilio (R.Br.) Christenh., comb. nov. Basionym: Cheilanthes regnelliana Mett. in Mart. Fl.
Basionym: Notholaena pumilio R.Br., Prod. Fl. Nov. Bras. 1(2): 39. 1870. Replaced synonym: Cheilanthes
Holland.: 146. 1810. flexuosa Baker, Syn. Filic.: 138. 1867, not: Hemionitis
Hemionitis purpusii (T.Reeves) Christenh., comb. nov. flexuosa (Kunze) Christenh.
Basionym: Cheilanthes purpusii T.Reeves, Rhodora Hemionitis regularis (Kunze) Christenh., comb. nov.
84: 293. 1982. Basionym: Adiantum regulare Kunze, Farrnkr. 2: 67.
Hemionitis pygmaea (Ching) Christenh., comb. nov. 1850.
Basionym: Aleuritopteris pygmaea Ching, Acta Hemionitis reynoldsii (F.Muell) Christenh., comb. nov.
Phytotax. Sin. 19: 61. 1981. Basionym: Notholaena reynoldsii F.Muell., Fragm.
Hemionitis pyornkrachzarkii Christenh., nom. nov. Phytogr. Austr. 8: 175. 1874.
Replaced synonym: Pellaea microphylla Fée, Hemionitis riedelii (Baker) Christenh., comb. nov.
Crypt. Vasc. Brésil 1: 43. 1869, not. Hemionitis Basionym: Pellaea riedelii Baker, Ann. Bot. (Oxford)
microphylla (Sw.) Christenh. Etymology: Named for 6: 213. 1891.
the ‘rock biter’ Pyornkrachzark, a character in The Hemionitis rivalis (Sehnem) Christenh., comb. nov.
Neverending Story (Ende, 1979). Basionym: Doryopteris rivalis Sehnem, Pesquisas,
Hemionitis qianguiensis (W.M.Chu & H.G.Zhou) Bot. 13: 25. 1961.
Christenh., comb. nov. Basionym: Cheilanthes Hemionitis robusta (Kuntze) Christenh., comb. nov.
qianguiensis W.M.Chu & H.G.Zhou, Guihaia 12: 321. Basionym: Allosorus robustus Kuntze, Linnaea 10:
1992. 502. 1836.
Hemionitis quadripinnata (Forssk.) Christenh., comb. Hemionitis rosei (Maxon) Christenh., comb. nov.
nov. Basionym: Pteris quadripinnata Forssk., Fl. Basionym: Notholaena rosei Maxon, Contr. U.S. Natl.
Aegypt.-Arab.: 186. 1775. Herb. 16: 59. 1912.
Hemionitis querquobadii Christenh., nom. nov. Hemionitis rosenstockii (Brade) Christenh., comb. nov.
Replaced synonym: Doryopteris pentagona Pic. Basionym: Doryopteris rosenstockii Brade, Bol. Mus.
Serm., Webbia 60: 231. 2005, not: Hemionits Nac. Rio de Janeiro 7: 143. 1931.
pentagona (Schelpe & N.C.Anthony) Christenh. Hemionitis rosulata (C.Chr.) Christenh., comb. nov.
Etymology: Named for Querquobad, the Silver Sage, Basionym: Cheilanthes rosulata C.Chr., Acta Horti
ruler of Amarganth in The Neverending Story (Ende, Gothob. 1: 89. 1924.
1979). (Figure 3E)
Hemionitis rotundifolia (G.Forst.) Christenh., comb.
Hemionitis quezelii (Tardieu) Christenh., comb. nov. nov. Basionym: Pteris rotundifolia G.Forst., Fl. Ins.
Basionym: Negripteris quezelii Tardieu in Quèzel, Austr. Prodr.: 79. 1786.
Mission Bot. Tibesti 92: t. 1(1–5). 1958.
Hemionitis rouxii (Fraser-Jenk. & E.Wollenw.)
Hemionitis quinquelobata (Fée) Christenh., comb. nov. Christenh., comb. nov. Basionym: Aleuritopteris
Basionym: Pellaea quinquelobata Fée, Crypt. Vasc. rouxii Fraser-Jenk. & E.Wollenw., Ferns & Fern-Allies
Brésil 1: 42. 1869. Nepal 1: 244. 2015.
Hemionitis raddiana (C.Presl) Christenh., comb. Hemionitis rufopunctata (Rosenst.) Christenh., comb.
nov. Basionym: Litobrochia raddiana C.Presl, Tent. nov. Basionym: Cheilanthes rufopunctata Rosenst.,
Pterid.: 149. 1836. Meded. Rijks-Herb. 19: 9. 1913.
Hemionitis radiata (L.) Christenh., comb. nov. Hemionitis rupicola (Maxon) Christenh., comb. nov.
Basionym: Adiantum radiatum L., Sp. Pl. 2: 1094. Basionym: Adiantopsis rupicola Maxon, Contr. U.S.
1753. Natl. Herb. 10: 485. 1908.
Hemionitis rawsonii (Pappe) Christenh., comb. nov. Hemionitis sagittifolia (Raddi) Christenh., comb. nov.
Basionym: Notholaena rawsonii Pappe in Pappe & Basionym: Pteris sagittifolia Raddi, Opusc. Sci. 3:
Rawson, Syn. Fil. Afr. Austr.: 42. 1858. 292. 1819.
Hemionitis recurvata (Baker) Christenh., comb. nov. Hemionitis sargentii (Christ) Christenh., comb. nov.
Basionym: Cheilanthes recurvata Baker, J. Bot. 16: Basionym: Gymnopteris sargentii Christ, Bot. Gaz.
299. 1878. 51: 355. 1911.
Hemionitis rediviva (Fée) Christenh., comb. nov. Hemionitis sarmientoi (Ponce) Christenh., comb. nov.
Basionym: Doryopteris rediviva Fée, Crypt. Vasc. Basionym: Cheilanthes sarmientoi Ponce, Amer. Fern
Brésil 2: 30. 1872. J. 79: 131. 1989.

Hemionitis scalariformis (Link-Pérez, Seabolt & Hemionitis solitaria (R.N.Tryon) Christenh., comb. nov.
Ledford) Christenh., comb. nov. Basionym: Basionym: Notholaena solitaria R.M.Tryon, Rhodora
Adiantopsis scalariformis Link-Pérez, Seabolt & 63: 82. 1961.
Ledford, Syst. Bot. 41: 916. 2016. Hemionitis sonorensis (Goodd.) Christenh., comb.
Hemionitis scariosa (Sw.) Christenh., comb. nov. nov. Basionym: Cheilanthes sonorensis Goodd.,
Basionym: Acrostichum scariosum Sw., Syn. Filic.: Muhlenbergia 8: 93. 1912.
16. 1806. Hemionitis speciosa (Ching & S.K.Wu) Christenh.,
Hemionitis schaffneri (E.Fourn.) Christenh., comb. comb. nov. Basionym: Aleuritopteris speciosa Ching
nov. Basionym: Aleuritopteris schaffneri E.Fourn., & S.K.Wu, Acta Phytotax. Sin. 19: 61. 1981.
Bull. Soc. Bot. France 27: 328. 1880. Hemionitis spiculata (Mickel) Christenh., comb. nov.
Hemionitis schimperi (Kunze) Christenh., comb. nov. Basionym: Cheilanthes spiculata Mickel, Mem. New
Basionym: Cheilanthes schimperi Kunze, Farrnkr. 1: York Bot. Gard. 88: 210. 2004.
52. 1840. Hemionitis squamosa (Gillies ex Hook. & Grev.)
Hemionitis schweinfurthii (Hieron.) Christenh., comb. Christenh., comb. nov. Basionym: Cheilanthes
nov. Basionym: Pteridella schweinfurthii Hieron., squamosa Gillies ex Hook. & Grev., Icon. Filic.: t. 151.
Pflanzenw. Ost-Afrikas C: 78. 1895. 1829.
Hemionitis scioana (Chiov.) Christenh., comb. nov. Hemionitis stieri (Rosenst.) Christenh., comb. nov.
Basionym: Mohria scioana Chiov., Atti Reale Accad. Basionym: Doryopteris stieri Rosenst., Hedwigia 46:
Italia, Mem. Cl. Sc. Fis. Mat. Nat. 11: 66. 1940. 86. 1906.
Hemionitis selinoides (Mickel) Christenh., comb. Hemionitis straminea (Ching) Christenh., comb. nov.
nov. Basionym: Cheilanthes selinoides Mickel in Basionym: Pellaea straminea Ching, Bull. Fan Mem.
McVaugh, Fl. Novo-Galiciana 17: 243. 1992. Inst. Biol. 2: 203. 1931.
Hemionitis senae (Baker) Christenh., comb. nov. Hemionitis striata (Desv.) Christenh., comb. nov.
Basionym: Adiantum senae Baker, J. Bot. 23: 217. Basionym: Pteris striata Desv., Mém. Soc. Linn. Paris
1885. 6: 300. 1827.
Hemionitis shensiensis (Ching) Christenh., comb. Hemionitis stuebeliana (Hieron.) Christenh., comb. &
nov. Basionym: Aleuritopteris shensiensis Ching, Fl. stat. nov. Basionym: Pellaea dealbata (Pursh) Prantl
Tsinling. 2: 66, 207. 1974. var. stuebeliana Hieron., Hedwigia 48: 225. 1909.
Hemionitis siamensis (S.K.Wu) Christenh., comb. nov. Hemionitis subargentea (Ching) Christenh., comb. nov.
Basionym: Aleuritopteris siamensis S.K.Wu, Mem. Basionym: Aleuritopteris subargentea Ching, Acta
Fac. Sci. Kyoto Univ., Biol. 8: 152. 1983. Phytotax. Sin. 19: 65. 1981.
Hemionitis sichouensis (Ching & S.K.Wu) Christenh., Hemionitis subdimorpha (C.B.Clarke & Baker)
comb. nov. Basionym: Aleuritopteris sichouensis Christenh., comb. & stat. nov. Basionym: Cheilanthes
Ching & S.K.Wu, Acta Phytotax. Sin. 19: 69. 1981. farinosa (Forssk.) Kaulf. var. subdimorpha C.B.Clarke
Hemionitis sieberi (Kunze) Christenh., comb. nov. & Baker, J. Linn. Soc., Bot. 24: 411. 1888.
Basionym: Cheilanthes sieberi Kunze, Ind. Sem. Hort. Hemionitis subpaleacea (Maxon) Christenh., comb.
Lips. 1839 and Pl. Preiss. (Lehmann) 2: 112. 1847 nov. Basionym: Bommeria subpaleacea Maxon,
(validation). Contr. U.S. Natl. Herb. 17: 169. 1913.
Hemionitis sikanda Christenh., nom. nov. Replaced Hemionitis subsimplex (Fée) Christenh., comb. nov.
synonym: Doryopteris pedatifida Christ, Bull. Basionym: Pellaea subsimplex Fée, Crypt. Vasc. Brésil
Herb. Boissier ser. 2, 2: 546. 1902, not: Hemionitis 1: 44. 1869.
pedatifida Sm. Etymology: Sikanda is the legendary Hemionitis subvillosa (Hook.) Christenh., comb. nov.
sword given to Bastian by Graograman in The Basionym: Cheilanthes subvillosa Hook., Sp. Filic. 2:
Neverending Story (Ende, 1979). 87. 1852.
Hemionitis similis (Ballard) Christenh., comb. nov. Hemionitis sulphurea (Cav.) Christenh., comb. nov.
Basionym: Cheilanthes similis Ballard, Kew Bull. 12: Basionym: Pteris sulphurea Cav., Descr. Pl.: 269.
47. 1957. 1801.
Hemionitis sinuata (Lag. ex Sw.) Christenh., comb. nov. Hemionitis surinamensis (Yesilyurt) Christenh., comb.
Basionym: Acrostichum sinuatum Lag. ex Sw., Syn. nov. Basionym: Doryopteris surinamensis Yesilyurt,
Filic.: 14. 1806. (Figure 3F) Brittonia 60: 63. 2008.
Hemionitis skinneri (Hook.) Christenh., comb. nov. Hemionitis takeuchii (W.H.Wagner) Christenh., comb.
Basionym: Pellaea skinneri Hook., Sp. Filic. 2: 141. nov. Basionym: Cheilanthes takeuchii W.H.Wagner,
1858. Contr. Univ. Michigan Herb. 19: 66. 1993.

21
Hemionitis tamburii (Hook.) Christenh., comb. nov. Hemionitis triphylla (Lam.) Christenh., comb. nov.
Basionym: Pellaea tamburii Hook., Sp. Filic. 2: 134. Basionym: Adiantum triphyllum Lam., Encycl. Meth.,
1858. Bot. 1: 41. 1783.
Hemionitis tenrecorum Christenh., nom. nov. Replaced Hemionitis tripinnata (Baker) Christenh., comb. nov.
synonym: Doryopteris madagascariensis Tardieu, Basionym: Pellaea tripinnata Baker, J. Linn. Soc., Bot.
Fl. Madagasc. 5(1): 148. 1958. Etymology: Named 25: 350. 1889.
for tenrecs (Tenrec ecaudatus) a native Madagascar Hemionitis truncata (Goodd.) Christenh., comb. nov.
mammal. Basionym: Pellaea truncata Goodd., Muhlenbergia
Hemionitis tenuifolia (Burm.f.) Christenh., comb. nov. 8: 94. 1912.
Basionym: Trichomanes tenuifolium Burm.f., Fl. Hemionitis tweediana (Hook.) Christenh., comb. nov.
Indica: 237. 1768. (Figure 4A) Basionym: Cheilanthes tweediana Hook., Sp. Pl. 2:
Hemionitis ternifolia (Cav.) Christenh., comb. nov. 84. 1852.
Basionym: Pteris ternifolia Cav., Descr. Pl.: 266. 1802. Hemionitis urglae Christenh., nom. nov. Replaced
Hemionitis thellungii (Herter) Christenh., comb. nov. synonym: Notholaena rigida Davenp., Gard. & Forest
Basionym: Cheilanthes thellungii Herter, Anal. Mus. 4: 519. 1891, not: Hemionitis rigida Cav. Etymology:
Nac. Montevideo, ser. 2, 1: 360. 1925. Named for Urgl, a gnome married to Engywook in
Hemionitis thwaitesii (Mett. ex Kuhn) Christenh., The Neverending Story (Ende, 1979)
comb. nov. Basionym: Cheilanthes thwaitesii Mett. Hemionitis uyulala Christenh., nom. nov. Replaced
ex Kuhn, Linnaea 36: 82. 1869. synonym: Cheilanthes pteroides Sw., Syn. Filic.:
Hemionitis tibetica (Fraser-Jenk. & Wangdi) Christenh., 128. 1806, not: Hemionitis pteroides (L.) Christenh.
comb. nov. Cheilanthes tibetica Fraser-Jenk. & Etymology: Named for Uyulala the Southern Oracle
Wangdi, Fern Gaz. 18: 218. 2009. in The Neverending Story (Ende, 1979).
Hemionitis tijucana (Brade & Rosenst.) Christenh., Hemionitis valdiviana (Phil.) Christenh., comb. nov.
comb. nov. Basionym: Doryopteris tijucana Brade & Basionym: Cheilanthes valdiviana Phil., Linnaea 29:
Rosenst., Bol. Mus. Nac. Rio de Janeiro 7: 144. 1931. 106. 1857.
Hemionitis timida (Link-Pérez & Hickey) Christenh., Hemionitis vandervekenii (Pic.-Serm.) Christenh.,
comb. nov. Basionym: Adiantopsis timida Link-Pérez comb. nov. Basionym: Aleuritopteris vandervekenii
& Hickey, Syst. Bot. 35: 579. 2011. Pic.-Serm., Bull. Jard. Bot. Natl. Belg. 53: 210. 1983.
Hemionitis timorensis (Alderw.) Christenh., comb. nov. Hemionitis varians (Raddi) Christenh., comb. nov.
Basionym: Pellaea timorensis Alderw., Bull. Dépt. Basionym: Pteris varians Raddi, Opusc. Sci. 3: 292.
Agric. Ind. Néerl. 18: 11. 1908. 1819.
Hemionitis tinaei (Tod.) Christenh., comb. nov. Hemionitis veitchii (Christ) Christenh., comb. nov.
Basionym: Cheilanthes tinaei Tod., Giorn. Sci. Nat. Basionym: Doryopteris veitchii Christ, Bull. Acad. Int.
Econ. Palermo 1: 217. 1866. Géogr. Bot. 16: 134. 1906.
Hemionitis triangularis (Kaulf.) Christenh., comb. nov. Hemionitis velutina (Tardieu & C.Chr.) Christenh.,
Basionym: Gymnogramma triangulare Kaulf., Enum. comb. nov. Basionym: Notholaena velutina Tardieu
Filic.: 73. 1824. & C.Chr., Notul. Syst. (Paris) 6: 167. 1938.
Hemionitis tricholepifera (Pic.-Serm.) Christenh., Hemionitis venusta (Fée) Christenh., comb. nov.
comb. nov. Basionym: Negripteris tricholepifera Pic.- Basionym: Notholaena venusta Brade, Anais Reun.
Serm., Webbia 27: 411. 1972. Sul-Amer. Bot. 2: 7. 1940.
Hemionitis trichomanoides (L.) Christenh., comb. nov. Hemionitis vestita (Desv.) Christenh., comb. nov.
Basionym: Pteris trichomanoides L., Sp. Pl. 2: 1074. Basionym: Notholaena vestita Desv., J. Bot. Appl. 1:
1753. 93. 1813.
Hemionitis trichophylla (Baker) Christenh., comb. nov. Hemionitis villosa (Davenp. ex Maxon) Christenh.,
Basionym: Cheilanthes trichophylla Baker, Ann. Bot. comb. nov. Basionym: Cheilanthes villosa Davenp.
(Oxford) 5: 211. 1891. ex Maxon, Proc. Biol. Soc. Wash. 31: 142. 1918.
Hemionitis trifurcata (Baker) Christenh., comb. nov. Hemionitis vincentii (M.S.Barker & Hickey) Christenh.,
Basionym: Cheilanthes trifurcata Baker, Bull. Misc. comb. nov. Basionym: Adiantopsis vincentii
Inform. Kew 1901(175-177): 144. 1901. M.S.Barker & Hickey, Ann. Missouri Bot. Gard. 93:
398. 2006.
Hemionitis trilobata (J.Prado) Christenh., comb. nov.
Basionym: Doryopteris trilobata J.Prado, Kew Bull. Hemionitis viridis (Forssk.) Christenh., comb. nov.
48: 13. 1993. Basionym: Pteris viridis Forssk., Fl. Aegypt.-Arab.:
186. 1775.

Hemionitis viscida (Davenp.) Christenh., comb. nov. Etymology: Named for Yikka, Bastian’s mule in The
Basionym: Cheilanthes viscida Davenp., Bull. Torrey Neverending Story (Ende, 1979).
Bot. Club 6: 191. 1877. Hemionitis yorii Christenh., nom. nov. Replaced
Hemionitis viscosa (Schuettp. & Windham) Christenh., synonym: Adiantopsis pentagona M.S.Barker &
comb. nov. Basionym: Pentagramma viscosa Hickey, Ann. Missouri Bot. Gard. 93: 392. 2006,
Schuettp. & Windham, Syst. Bot. 40: 642. 2015. not: Hemionits pentagona (Schelpe & N.C.Anthony)
Hemionitis volcanensis (de la Sota) Christenh., comb. Christenh. Etymology: Named for dream picture
nov. Basionym: Cheilanthes volcanensis de la Sota, miner Yor in The Neverending Story (Ende, 1979).
Darwiniana 17: 78. 1972. Hemionitis yunnanensis (Ching) Christenh., comb.
Hemionitis vooshvazool Christenh., nom. nov. nov. Basionym: Pellaea yunnanensis Ching, Acta
Replaced synonym: Pellaea microphylla Mett. Phytotax. Sin. 20: 235. 1982.
ex Kuhn, Linnaea 36: 86. 1869, not. Hemionitis Pteris anopteris Christenh., nom. nov. Replaced
microphylla (Sw.) Christenh. Etymology: Named for a synonym: Anopteris hexagona (L.) C.Chr. subsp.
nighthob, one of the messengers in The Neverending intermedia C.V.Morton, Bull. Jard. Bot. État Bruxelles
Story (Ende, 1979). 27: 583. 1957, not: Pteris intermedia Blume.
Hemionitis welwitschii (Hook. ex Baker) Christenh., Etymology: Named for the former genus Anopteris
comb. nov. Basionym: Cheilanthes welwitschii Hook. now included in Pteris.
ex Baker, Syn. Filic.: 142. 1874.
Hemionitis windhamii (D.M.Benham) Christenh., F23. Aspleniaceae. References: Kato & Darnaedi
comb. nov. Basionym: Astrolepis windhamii (1988), Ohlsen et al. (2014), Perrie et al. (2014),
D.M.Benham, Amer. Fern J. 82: 60. 1992. Almeida et al. (2016), Wei et al. (2018)
Hemionitis woodfordii (C.H.Wright) Christenh., comb. Asplenium pleurosorum Christenh., nom. nov.
nov. Basionym: Cassebeera woodfordii C.H.Wright, Replaced synonym: Grammitis rutifolia R.Br.,
Bull. Misc. Inf. Kew 1908(4): 183. 1908. Prod. Fl. Nov. Holland.: 146. 1810, not: Asplenium
Hemionitis wootonii (Maxon) Christenh., comb. nov. rutifolium Kunze. Etymology: Named for the
Basionym: Cheilanthes wootonii Maxon, Proc. Biol. now defunct genus Pleurosorum, a synonym of
Soc. Washington 31: 146. 1918. Asplenium. (Figures 4C–D)
Hemionitis wrightiana (Hook.) Christenh., comb. nov. Athyrium approximatum (W.M.Chu) Christenh., comb.
Basionym: Pellaea wrightiana Hook., Sp. Filic. 2: 142. nov. Basionym: Cornopteris approximata W.M.Chu,
1858. (Figure 4B) Acta Bot. Yunnan. 3: 342. 1981.
Hemionitis wrightii (Hook.) Christenh., comb. nov. Athyrium badium (Ching) Christenh., comb. nov.
Basionym: Cheilanthes wrightii Hook., Sp. Filic. 2: 87. Basionym: Cornopteris badia Ching, Bull. Fan Mem.
1858. Inst. Biol. Bot. 11: 58. 1941.
Hemionitis xayide Christenh., nom. nov. Replaced Athyrium banajaoense (C.Chr.) Christenh., comb. nov.
synonym: Cheilanthes incisa Kunze, Abh. Basionym: Dryopteris banajaoensis C.Chr., Ind. Filic.,
Senckenberg. Naturf. Gesell. 3: 100. 1859, not: Suppl. 1906–1912: 30. 1913. Replaced synonym:
Hemionitis incisa Blanco. Etymology: Named for Dryopteris tenerrima Copeland, Philipp., J. Sci. Bot.
Xayíde, an evil sorceress in The Neverending Story 4: 111. 1909, not: Dryopteris tenerrima Rosenst.
(Ende, 1979) Athyrium filix-incantatrix Christenh., nom. nov.
Hemionitis yalungensis (H.S.Kung) Christenh., comb. Replaced synonym: Cornopteris latibasis W.M.Chu,
nov. Basionym: Aleuritopteris yalungensis H.S.Kung, Acta Bot. Yunnan. 3: 344. 1981, not: Athyrium
Acta Bot. Yunnan. 5: 356. 1983. latibasis Ching. Etymology: The epithet means
‘witch fern’, one of the common names of this
Hemionitis yavapensis (T.Reeves ex Windham)
species.
Christenh., comb. nov. Basionym: Cheilanthes
yavapensis T.Reeves ex Windham, Contr. Univ. Athyrium majus (W.M.Chu) Christenh., comb. nov.
Michigan Herb. 19: 32. 1993. Basionym: Cornopteris major W.M.Chu, Acta Bot.
Yunnan. 3: 348. 1981.
Hemionitis ygramulorum Christenh., nom. nov.
Replaced synonym: Cassebeera pinnata Kaulf., Athyrium narthex Christenh., nom. nov. Replaced
Enum. Filic.: 217. 1824, not: Hemionitis pinnata synonym: Cornopteris omeiensis Ching, Bull. Fan
Hook.f. Etymology: Named for Ygramul the Many, a Mem. Inst. Biol., n.s. 1: 287. 1949, not: Athyrium
shapeshifter in The Neverending Story (Ende, 1979). omeiense Ching. Etymology: From Classical Greek
νάρθηξ (narthex), meaning a ‘giant fennel’ (Ferula
Hemionitis yikka Christenh., nom. nov. Replaced
communis, Apiaceae) or an ‘atrium’, referring to
synonym: Pellaea rufa A.F.Tryon, Ann. Missouri Bot.
the small indusial pockets that are ‘porch-like’ or
Gard. 42: 101. 1955, not: Hemionitis rufa Sw.
swollen like a fennel bulb.

23
A B
C D
E F
Figure 4: Ferns. A, Hemionitis tenuifolia (Burm.f.) Christenh.; B, Hemionitis wrightiana (Hook.) Christenh.;
C–D, Asplenium pleurosorum Christenh.; E–F, Blechnum cyatheoides (Kaulf.) Christenh.. All images © Maarten
Christenhusz.

Athyrium philippinense (M.Kato) Christenh., comb. 234. 1862.

nov. Basionym: Cornopteris philippinensis M.Kato, Phegopteris fijiensis (K.U.Kramer & E.Zogg) Christenh.,
Acta Phytotax. Geobot. 30: 110. 1979. comb. nov. Basionym: Pseudophegopteris fijiensis
Athyrium pseudofluviale (Ching & W.M.Chu) K.U.Kramer & E.Zogg, Blumea 33: 423. 1988.
Christenh., comb. nov. Basionym: Cornopteris Phegopteris henriquesii (Baker) Christenh., comb. nov.
pseudofluvialis Ching & W.M.Chu, Acta Bot. Yunnan. Basionym: Polypodium henriquesii Baker, Bol. Soc.
3: 345. 1981. Brot. 4: 154. 1886.
Athyrium seramense (M.Kato) Christenh., comb. nov. Phegopteris hirtirachis (C.Chr.) Christenh., comb. nov.
Basionym: Cornopteris seramensis M.Kato, J. Jap. Basionym: Dryopteris hirtirachis C.Chr., Ind. Filic.,
Bot. 61: 233. 1986. Suppl. 1913–1916: 15. 1917.
Athyrium yunnanense Christenh., nom. nov. Replaced Phegopteris kinabaluensis (Holttum) Christenh., comb.
synonym: Cornopteris latiloba Ching, Fl. Xizang. nov. Basionym: Pseudophegopteris kinabaluensis
1: 144. 1983, not: Athyrium latilobum Copel. Holttum, Blumea 17: 16. 1969.
Etymology: Named for Yunnan, a Chinese province Phegopteris microstegia (Hook.) Christenh., comb.
where this species is found. nov. Basionym: Nephrodium microstegium Hook.,
Blechnum cyatheoides (Kaulf.) Christenh., comb. nov. Sp. Filic. 1: 119. 1862.
Basionym: Sadleria cyatheoides Kaulf., Enum. Filic.: Phegopteris persimilis (Baker) Christenh., comb. nov.
162. 1824. (Figures E–F) Basionym: Polypodium persimile Baker, J. Bot. 14:
Blechnum unisorum (Baker) Christenh., comb nov. 344. 1876.
Basionym: Polypodium unisorum Baker, Syn. Filicl.: Phegopteris rectangularis (Zoll.) Christenh., comb.
307. 1867. nov. Basionym: Polypodium rectangulare Zoll., Syst.
Blechnum wagnerianum (D.D.Palmer & T.Flynn) Verz.: 37, 48. 1854.
Christenh., comb. nov. Basionym: Sadleria Phegopteris tibetana (Ching & S.K.Wu) Christenh.,
wagneriana D.D.Palmer & T.Flynn, Pacific Sci. 51: comb. nov. Basionym: Pseudophegopteris tibetana
302. 1997. Ching & S.K.Wu, Fl. Xizang. 1: 164. 1983, not:
Deparia thwaitesii (A.Braun ex Mett.) Christenh., Phegopteris tibetica Ching.
comb. nov. Basionym: Asplenium thwaitesii A.Braun Phegopteris yigongensis (Ching) Christenh., comb.
ex Mett., Abh. Senckenberg. Naturf. Gesell. 6: 183. nov. Basionym: Pseudophegopteris yigongensis
1859. Ching, Fl. Xizang. 1: 165. 1983.
Deparia zanzibarica (Baker) Christenh., comb. nov. Phegopteris zayuensis (Ching & S.K.Wu) Christenh.,
Asplenium zanzibaricum Baker, Ann. Bot. (Oxford) 5: comb. nov. Basionym: Pseudophegopteris zayuensis
311. 1891. Ching & S.K.Wu, Fl. Xizang. 1: 164. 1983.
Diplaziopsis flavoviridis (Alston) Christenh. comb. nov. Thelypteris abbreviata (Ching & K.H.Shing) Christenh.,
Basionym: Diplazium flavoviride Alston, J. Bot. 78: comb. nov. Basionym: Cyclosorus abbreviatus Ching
226. 1940. & K.H.Shing, Jiangxi Sci. 8: 45. 1990.
Macrothelypteris banaensis (Tardieu & C.Chr.) Thelypteris aberrans (Holttum) Christenh., comb.
Christenh., comb. nov. Basionym: Thelypteris nov. Basionym: Pneumatopteris aberrans Holttum,
banaensis Tardieu & C.Chr., Notul. Syst. (Paris) 7: 66. Blumea 33: 401. 1988.
1938.
Thelypteris acutidens (Holttum) Christenh., comb. nov.
Phegopteris andringitrensis (Rakotondr.) Christenh., Basionym: Chingia acutidens Holttum, Kalikasan 3:
comb. nov. Basionym: Pseudophegopteris 17. 1974.
andringitrensis Rakotondr., Adansonia 34: 232.
Thelypteris adenopelta (Holttum) Christenh., comb.
2012.
nov. Basionym: Christella adenopelta Holttum, Kew
Phegopteris aubertii (Desv.) Christenh., comb. nov. Bull. 31: 322. 1976.
Basionym: Polypodium aubertii Desv., Mém. Soc.
Thelypteris alcasidii (Holttum) Christenh., comb. nov.
Linn. Paris 6: 243. 1827.
Basionym: Sphaerostephanos alcasidii Holttum, Fl.
Phegopteris brevipes (Ching & S.K.Wu) Christenh., Males., Ser. 2, Pterid. l: 503. 1982.
comb. nov. Basionym: Pseudophegopteris brevipes
Thelypteris alpina (Holttum) Christenh., comb. nov.
Ching & S.K.Wu, Fl. Xizang. 1: 163. 1983.
Basionym: Sphaerostephanos alpinus Holttum, Fl.
Phegopteris cyclocarpa (Holttum) Christenh., comb. Males., Ser. 2, Pterid. l: 488. 1982.
nov. Basionym: Pseudophegopteris cyclocarpa
Thelypteris alticola (Holttum) Christenh., comb. nov.
Holttum, Blumea 13: 131. 1965.
Basionym: Sphaerostephanos alticola Holttum, Fl.
Phegopteris dianae (Hook.) Christenh., comb. nov. Males., Ser. 2, Pterid. l: 484. 1982.
Basionym: Polypodium dianae Hook., Sp. Filic. 4:

25
A B
C D
Figure 5: Ferns. A, Thelypteris dodo Christenh.; B–C, Thelypteris tahitensis (Holttum) Christenh.; and D,
Campyloneurum crassifolium (L.) Christenh.. All images © Maarten Christenhusz.

Thelypteris amphitricha (Holttum) Christenh., comb. Ching ex Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 328.
nov. Basionym: Pronephrium amphitrichum 1999, not: Thelypteris submarginalis (Langsd. &
Holttum, Fl. Males., Ser. 2, Pterid. l: 521. 1982. Fisch.) Small. Etymology: Named for the Baggins
Thelypteris angiensis (Holttum) Christenh., comb. family of Hobbiton (Tolkien, 1937–1955).
nov. Basionym: Plesioneuron angiense Holttum, Fl. Thelypteris balabacensis (P.M.Zamora & Co)
Males., Ser. 2, Pterid. l: 406. 1982. Christenh., comb. nov. Basionym: Pronephrium
Thelypteris angmarica Christenh., nom. nov. Replaced balabacensis P.M.Zamora & Co, Nat. Appl. Sci. Bull.,
synonym: Parathelypteris trichochlamys Ching ex Univ. Philipp. 32: 45. 1980.
K.H.Shing, Fl. Reipubl. Popularis Sin. 4: 320. 1999, Thelypteris balrog Christenh., nom. nov. Replaced
not: Thelypteris trichochlamys (Holttum) Christenh. synonym: Plesioneuron saxicola M.Kato, Bull. Natl.
Etymology: Named for Angmar, a fictional kingdom Mus. Nat. Sci., Tokyo, B 33: 23. 2007, not: Thelypteris
in Middle-earth, at the north end of the Misty saxicola (Sw.) C.F.Reed. Etymology: Named for a
Mountains (Tolkien, 1937–1955). fictional menacing creature in Tolkien’s (1937–1955)
Thelypteris angustibasis (Holttum) Christenh., comb. Middle-earth legendarium.
nov. Basionym: Sphaerostephanos angustibasis Thelypteris basicurtata (Holttum) Christenh., comb.
Holttum, Fl. Males., Ser. 2, Pterid. l: 476. 1982. nov. Basionym: Pneumatopteris basicurtata
Thelypteris angusticaudata (Holttum) Christenh., Holttum, Blumea 21: 309. 1973.
comb. nov. Basionym: Pneumatopteris Thelypteris batulantensis (Holttum) Christenh., comb.
angusticaudata Holttum, Blumea 21: 308. 1973. nov. Basionym: Sphaerostephanos batulantensis
Thelypteris angustipinna (Ching) Christenh., comb. Holttum, Fl. Males., Ser. 2, Pterid. l: 473. 1982.
& stat. nov. Basionym: Cyclosorus truncatus (Poir.) Thelypteris beleriandica Christenh., nom. nov.
Farw. var. angustipinnus Ching, Bull. Fan Mem. Inst. Replaced synonym: Goniopteris bradei Salino,
Biol. Bot. 8: 218. 1938. Brittonia 68: 448. 2016. Etymology: Named for
Thelypteris aragornii Christenh., nom. nov. Replaced Beleriand, a region in Middle-earth (Tolkien, 1937–
synonym: Pronephrium medogensis Y.X.Lin, Fl. 1955).
Reipubl. Popularis Sin. 4: 351. 1999, not: Thelypteris Thelypteris benoiteana (Gaudich.) Christenh. comb.
medogensis (Ching & S.K.Wu) Christenh. Etymology: nov. Basionym: Polystichum benoiteanum Gaudich.,
Named for Aragorn II, chieftain of the Dúnedain of Voy. Uranie, Bot.: 331. 1828.
Arnor and member of the Fellowship of the Ring Thelypteris bewaniensis (Holttum) Christenh. comb.
(Tolkien, 1937–1955). nov. Basionym: Chingia bewaniensis Holttum, Fl.
Thelypteris arenicola (Holttum) Christenh., comb. Males., Ser. 2, Pterid. l: 393. 1982.
nov. Basionym: Mesophlebion arenicola Holttum, Thelypteris bilboi Christenh., nom. nov. Replaced
Blumea 22: 228. 1975. synonym: Pneumatopteris imbricata Holttum,
Thelypteris arwenae Christenh., nom. nov. Replaced Blumea 21: 322. 1973, not: Thelypteris imbricata
synonym: Pseudocyclosorus stramineus Ching ex (Liebm.) C.F.Reed. Etymology: Named for Bilbo
Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 329. 1999, not: Baggins of Hobbiton, title character of The Hobbit
Thelypteris straminea (Baker) C.F.Reed. Etymology: (Tolkien, 1937).
Named for elf Arwen Undómiel or Evenstar (Tolkien, Thelypteris birii (R.D.Dixit & Balkr.) Christenh., comb.
1937–1955). nov. Basionym: Pronephrium birii R.D.Dixit & Balkr.,
Thelypteris auctipinna (Holttum) Christenh., comb. Indian Fern J. 7: 20. 1990.
nov. Basionym: Pneumatopteris auctipinna Holttum, Thelypteris bombadilii Christenh., nom. nov. Replaced
Blumea 21: 310. 1973. synonym: Metathelypteris glandulifera Ching ex
Thelypteris aureoglandulifera (Ching) Christenh., Shing, Fl. Reipubl. Popularis Sin. 4: 321. 1999,
comb. nov. Basionym: Cyclosorus aureoglandulifer not: Thelypteris glandulifera (Brack.) C.F.Reed.
Ching, Fl. Reipubl. Popularis Sin. 4: 345. 1999. Etymology: Named for Tom Bombadil, supporting
Thelypteris aureoglandulosa (Ching & K.H.Shing) character in Tolkien’s (1954, 1955, 1962) Middle-
Christenh., comb. nov. Basionym: Cyclosorus earth legendarium.
aureoglandulosus Ching & KH.Shing, Bull. Bot. Res., Thelypteris borealis (H.Hara) Christenh., comb. & stat.
Harbin 3(3): 4. 1983. nov. Basionym: Dryopteris nipponica (Franch. & Sav.)
Thelypteris australis (Holttum) Christenh., comb. nov. C.Chr. var. borealis H.Hara, Bot. Mag (Tokyo) 48: 695.
Basionym: Chingia australis Holttum, Kew Bull. 41: 1934.
518. 1986. Thelypteris boridensis (Holttum) Christenh., comb.
Thelypteris bagginsiorum Christenh., nom. nov. nov. Basionym: Pneumatopteris boridensis Holttum,
Replaced synonym: Pseudocyclosorus submarginalis Fl. Males., Ser. 2, Pterid. l: 434. 1982.

27
Thelypteris boromir Christenh., nom. nov. Replaced Thelypteris caulescens (Holttum) Christenh., comb.
synonym: Trigonospora angustifrons Sledge, nov. Basionym: Sphaerostephanos caulescens
Bull. Brit. Mus. (Nat. Hist.), Bot. 8: 23. 1981, not: Holttum, Fl. Males., Ser. 2, Pterid. l: 472. 1982.
Thelypteris angustifrons (Miq.) Ching. Etymology: Thelypteris cavaleriei (H.Lév.) Christenh., comb. nov.
Named for Boromir, brother of Faramir and Basionym: Christella cavaleriei H.Lév., Fl. Kouy-
supporting character in Tolkien’s (1937–1955) Tchéou: 474. 1915.
Middle-earth legendarium. Thelypteris celebornii Christenh., nom. nov. Replaced
Thelypteris braithwaitei (Holttum) Christenh., comb. synonym: Pronephrium clarkei Sarn.Singh &
nov. Basionym: Sphaerostephanos braithwaitei Panigrahi, Ferns & Fern-Allies Arunachal Pradesh 2:
Holttum, Allertonia 1: 206. 1977. 726. 2005, not: Thelypteris clarkei (Bedd.) C.F.Reed.
Thelypteris brauseanum (Holttum) Christenh., comb. Etymology: Named for the elf Celeborn (pronounced
nov. Basionym: Pronephrium brauseanum Holttum, with a hard c as in ‘cat’), from the Middle-earth
Blumea 20: 107. 1972. legendarium (Tolkien, 1954–1955).
Thelypteris brevipes (Ching) Christenh., comb. nov. Thelypteris centrochinensis (Ching ex Y.X.Lin)
Basionym: Cyclosorus brevipes Ching, Fl. Reipubl. Christenh., comb. nov. Basionym: Leptogramma
Popularis Sin. 4: 336. 1999. centrochinensis Ching ex Y.X.Lin, Fl. Reipubl.
Thelypteris bryanii (C.Chr.) Christenh., comb. nov. Popularis Sin. 4: 323. 1999.
Basionym: Dryopteris bryanii C.Chr., Bull. Bishop Thelypteris ceramica (Alderw.) Christenh., comb. nov.
Mus. 177: 89. 1943. Basionym: Phegopteris ceramica Alderw., Bull. Dépt.
Thelypteris bulusanica (Holttum) Christenh., comb. Agric. Indes Néerl. 18: 15. 1908.
nov. Basionym: Haplodictyum bulusanicum Holttum, Thelypteris changbaishanensis (Ching ex K.H.Shing)
Kalikasan 2: 61. 1973. Christenh., comb. nov. Basionym: Parathelypteris
Thelypteris burrowsiorum (N.R.Crouch) Christenh., changbaishanensis Ching ex K.H.Shing, Fl. Reipubl.
comb. nov. Basionym: Metathelypteris Popularis Sin. 4: 319. 1999.
burrowsiorum N.R.Crouch, Bothalia 41: 193. 2011. Thelypteris cheesmaniae (Holttum) Christenh., comb.
Thelypteris burundensis (Pic.-Serm.) Christenh., comb. nov. Basionym: Pneumatopteris cheesmaniae
nov. Basionym: Christella burundensis Pic.-Serm., Holttum, Blumea 21: 307. 1973.
Bull. Jard. Bot. Natl. Belg. 53: 277. 1983. Thelypteris chingii (Z.Y.Liu) Christenh., comb. nov.
Thelypteris buwaldae (Holttum) Christenh., comb. Basionym: Cyclosorus chingii Z.Y.Liu, Bull. Bot. Res.,
nov. Basionym: Pronephrium buwaldae Holttum, Harbin 3: 25. 1983.
Blumea 20: 115. 1972. Thelypteris clavata (K.H.Shing) Christenh., comb. nov.
Thelypteris caii (Ching) Christenh., comb. nov. Basionym: Cyclosorus clavatus K.H.Shing, Fl. Reipubl.
Basionym: Cyclosorus caii Ching, Fl. Reipubl. Popularis Sin. 4: 350. 1999. Replaced synonym:
Popularis Sin. 4: 346. 1999. Cyclosorus cuneatus Ching ex K.H.Shing, Fl. Reipubl.
Popularis Sin. 4: 347. 1999; not: Thelypteris cuneata
Thelypteris camarinensis (Holttum) Christenh., comb.
(C.Chr.) C.F.Reed.
nov. Basionym: Nannothelypteris camarinensis
Holtt., Kalikasan 5: 119. 1976. Thelypteris clavipilosa (Holttum) Christenh., comb.
nov. Basionym: Chingia clavipilosa Holttum,
Thelypteris camerounensis (Holttum) Christenh.,
Kalikasan 3: 23. 1974.
comb. nov. Basionym: Pseudocyclosorus
camerounensis Holttum, J. S. Afr. Bot. 40: 137. 1974. Thelypteris comorensis (Holttum) Christenh., comb.
nov. Basionym: Pneumatopteris comorensis
Thelypteris caoshanensis (Ching ex K.H.Shing)
Holttum, Blumea 21: 318. 1973.
Christenh., comb. nov. Basionym: Parathelypteris
caoshanensis Ching ex K.H.Shing, Fl. Reipubl. Thelypteris contracta (Ching) Christenh., comb. nov.
Popularis Sin. 4: 321. 1999. Basionym: Basionym: Cyclosorus contractus Ching,
Fl. Reipubl. Popularis Sinicae 4: 192. 1999.
Thelypteris caroli (Holttum) Christenh., comb. nov.
Basionym: Mesophlebion caroli Holttum, Blumea Thelypteris convergens (Holttum) Christenh., comb.
22: 227. 1975. nov. Basionym: Sphaerostephanos convergens
Holttum, Fl. Males., Ser. 2, Pterid. l: 477. 1982.
Thelypteris carrii (Holttum) Christenh., comb. nov.
Basionym: Sphaerostephanos carrii Holttum, Fl. Thelypteris costularisora (Ching ex K.H.Shing)
Males., Ser. 2, Pterid. l: 463. 1982. Christenh., comb. nov. Basionym: Cyclogramma
costularisora Ching ex K.H.Shing, Ann. Bot. Fenn.
Thelypteris cartilagidens (P.M.Zamora & Co)
51: 407. 2014, and Fl. Reipubl. Popularis Sin. 4: 322.
Christenh., comb. nov. Basionym: Sphaerostephanos
1999.
cartilagidens P.M.Zamora & Co, Nat. Appl. Sci. Bull.,
Univ. Philipp. 32: 43. 1980.

Thelypteris crenulata (Holttum) Christenh., comb. Thelypteris draco Christenh., nom. nov. Replaced
nov. Basionym: Pronephrium crenulatum Holttum, synonym: Nephrodium acrostichoides Desv., Mém.
Blumea 20: 123. 1972. Soc. Linn. Paris 6: 255. 1827, not: Thelypteris
Thelypteris croftii (Holttum) Christenh., comb. nov. acrostichoides (Michx.) Nieuwl. Etymology: Draco
Basionym: Plesioneuron croftii Holttum, Fl. Males. meaning dragon, a mythological creature found in
Ser. 2, Pterid. l: 411. 1982. many stories and legends around the world.
Thelypteris cystodioides (Holttum) Christenh., comb. Thelypteris dryas (Holttum) Christenh., comb. nov.
nov. Basionym: Plesioneuron cystodioides Holttum, Basionym: Plesioneuron dryas Holttum, Blumea 22:
Fl. Males., Ser. 2, Pterid. l: 405. 1982. 247. 1975.
Thelypteris damingshanensis (Ching ex Y.X.Lin) Thelypteris dulitensis (Holttum) Christenh., comb. nov.
Christenh., comb. nov. Basionym: Pseudocyclosorus Basionym: Mesophlebion dulitense Holttum, Blumea
damingshanensis Ching ex Y.X.Lin, Fl. Reipubl. 22: 229. 1975.
Popularis Sin. 4: 330. 1999. Thelypteris dulongjiangensis (W.M.Chu) Christenh.,
Thelypteris darainensis (Rakotondr.) Christenh., comb. comb. nov. Basionym: Pseudocyclosorus
nov. Basionym: Christella darainensis Rakotondr., dulongjiangensis W.M.Chu, Acta Bot. Yunnan.,
Adansonia, sér. 3, 34: 224. 2012. Suppl. 5: 44. 1992.
Thelypteris daymaniana (Holttum) Christenh., comb. Thelypteris eburnea (Holttum) Christenh., comb. nov.
nov. Basionym: Sphaerostephanos daymanianus Basionym: Pneumatopteris eburnea Holttum, Fl.
Holttum, Fl. Males., Ser. 2, Pterid. l: 451. 1982. Males., Ser. 2, Pterid. l: 434. 1982.
Thelypteris deficiens (Holttum) Christenh., comb. Thelypteris efogensis (Holttum) Christenh., comb. nov.
nov. Basionym: Pneumatopteris deficiens Holttum, Basionym: Sphaerostephanos efogensis Holttum, Fl.
Blumea 21: 321. 1973. Males., Ser. 2, Pterid. l: 482. 1982.
Thelypteris dehuaensis (Y.X.Lin) Christenh., comb. nov. Thelypteris egenolfioides (Holttum) Christenh., comb.
Basionym: Pseudocyclosorus dehuaensis Y.X.Lin, Fl. nov. Basionym: Pneumatopteris egenolfioides
Reipubl. Popularis Sin. 4: 326. 1999. Holttum, Blumea 21: 306. 1973.
Thelypteris delicata (R.S.Fern. & Salino) Christenh., Thelypteris ekutiensis (Holttum) Christenh., comb. nov.
comb. nov. Basionym: Meniscium delicatum Basionym: Sphaerostephanos ekutiensis Holttum, Fl.
R.S.Fern. & Salino, Phytotaxa 273: 176. 2016. Males., Ser. 2, Pterid. l: 483. 1982.
Thelypteris deltoideofrons (Ching ex W.M.Chu Thelypteris elrondii Christenh., nom. nov. Replaced
& S.G.Lu) Christenh., comb. nov. Basionym: synonym: Meniscium divergens R.S.Fern. & Salino,
Metathelypteris deltoideofrons Ching ex W.M.Chu & Phytotaxa 184: 2. 2014, not: Thelypteris divergens
S.G.Lu, Fl. Yunnan. 20: 720. 2006. (Rosenst.) C.F.Reed. Etymology: Named for Elrond
Half-elven, Lord of Rivendell in the Middle-earth
Thelypteris dimidiolobata (Holttum) Christenh., comb.
legendarium (Tolkien, 1937–1955).
nov. Basionym: Sphaerostephanos dimidiolobatus
Holttum, Fl. Males., Ser. 2, Pterid. l: 459. 1982. Thelypteris emeiensis (Y.X.Lin) Christenh., comb. nov.
Basionym: Glaphyropteridopsis emeiensis Y.X.Lin, Fl.
Thelypteris diminuta (Copel.) Christenh., comb. nov.
Reipubl. Popularis Sin. 4: 324. 1999.
Basionym: Dryopteris diminuta Copel., Philipp. J. Sci.
40: 298. 1929. Thelypteris eminens (Baker) Christenh., comb. nov.
Basionym: Nephrodium eminens Baker, J. Bot. 18:
Thelypteris dissitifolia (Holttum) Christenh., comb.
213. 1880.
nov. Basionym: Stegnogramma dissitifolia Holttum,
Fl. Males., Ser. 2, Pterid. 1: 543. 1981. Thelypteris endertii (C.Chr.) Christenh., comb. nov.
Basionym: Dryopteris endertii C.Chr., Dansk Bot. Ark.
Thelypteris diversiloba (C.Presl) Christenh., comb.
9: 60. 1937.
nov. Basionym: Nephrodium diversilobum C.Presl,
Epimel. Bot.: 47. 1851. Thelypteris endorica Christenh., nom. nov. Replaced
synonym: Cyclosorus serratus Copel. Philipp. J. Sci.
Thelypteris doctersii (Holttum) Christenh., comb. nov.
81: 36. 1952, not: Thelypteris serrata (Cav.) Alston.
Basionym: Plesioneuron doctersii Holttum, Blumea
Etymology: Named for Endor, the Quenya name of
22: 246. 1975.
Middle-earth (Tolkien, 1937–1955).
Thelypteris dodo Christenh., nom. nov. Replaced
Thelypteris eowynae Christenh., nom. nov. Replaced
synonym: Aspidium elatum Bojer ex Willd. Sp. Pl.
synonym: Nephrodium motleyanum Hook., Syn.
(ed. quarta) 5: 285. 1810, not: Thelypteris elata
Filic.: 266. 1867, not: Thelypteris motleyana
Schelpe. Etymology: Named for the dodo (Raphus
(Christ) Holttum. Etymology: Named for Éowyn, a
cucullatus), an extinct bird of Mauritius, where this
noblewoman of Rowan in Middle-earth (Tolkien,
species of fern is found. (Figure 5A)
1954–1955).

29
Thelypteris eriocarpa (Ching) Christenh., comb. nov. Thelypteris foxworthyi (Holttum) Christenh., comb.
Basionym: Glaphyropteridopsis eriocarpa Ching, nov. Basionym: Sphaerostephanos foxworthyi
Acta Phytotax. Sin. 8: 321. 1963. Holttum, Fl. Males. Ser. 2, Pterid. l: 475. 1982.
Thelypteris erwinii (Holttum) Christenh., comb. nov. Thelypteris frodoi Christenh., nom. nov. Replaced
Basionym: Sphaerostephanos erwinii Holttum, Fl. synonym: Trigonospora loyalii Panigrahi & Sarn.
Males., Ser. 2, Pterid. l: 504. 1982. Singh, Ferns & Fern-Allies Arunachal Pradesh 2: 772.
Thelypteris excisa (Holttum) Christenh., comb. nov. 2005, not: Thelypteris loyalii (Holttum) Fraser-Jenk..
Basionym: Pseudocyclosorus excisus Holttum, Etymology: Named for Frodo Baggins of Hobbiton,
Blumea 13: 133. 1965. member of the Fellowship of the Ring and bearer of
the One Ring (Tolkien, 1937–1955).
Thelypteris exsculpta (Baker) Christenh., comb. nov.
Basionym: Acrostichum exsculptum Baker, J. Bot. 26: Thelypteris fuchsii (Holttum) Christenh., comb. nov.
326. 1888. Basionym: Plesioneuron fuchsii Holttum, Blumea 22:
236. 1975.
Thelypteris falcatiloba (Holttum) Christenh., comb.
nov. Basionym: Mesophlebion falcatilobum Holttum, Thelypteris fugongensis (Y.X.Lin) Christenh., comb.
Fl. Males., Ser. 2, Pterid. l: 382. 1982. nov. Basionym: Pseudocyclosorus fugongensis
Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 327. 1999.
Thelypteris fangornica Christenh., nom. nov. Replaced
synonym: Dryopteris stegnogramme C.Chr. var. Thelypteris furcatovenulosa (Y.X.Lin) Christenh., comb.
asplenoides C.Chr., Acta Horti Gothob. 1: 56. nov. Basionym: Pseudocyclosorus furcatovenulosus
1924, not: Thelypteris asplenioides (Sw.) Proctor. Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 330. 1999.
Etymology: Named for Farngorn (or Beardtree), an Thelypteris galadrielae Christenh., nom. nov.
ancient forest located in Middle-earth and home of Replaced synonym: Parathelypteris caudata Ching
Treebeard the Ent (Tolkien, 1937–1955). ex K.H.Shing, Fl. Reipubl. Popularis Sin. 4: 320.
Thelypteris faramir (Ching ex Y.X.Lin) Christenh., 1999, not: Thelypteris caudata (Ching) C.F.Reed.
nom. nov. Replaced synonym: Pseudocyclosorus Etymology: Named for royal elf Galadriel, Lady of
angustipinnus Ching ex Y.X.Lin, Fl. Reipubl. Popularis Lórien (Tolkien, 1937–1955).
Sin. 4: 329. 1999, not: Thelypteris angustipinna Thelypteris gandalfii Christenh., nom. nov. Replaced
(Ching) Christenh. Etymology: Named for Faramir, synonym: Trigonospora glandulosa Sledge, Bull. Brit.
second son of Denethor II of Gondor in Middle-earth Mus. (Nat. Hist.), Bot. 8: 23. 1981, not: Thelypteris
(Tolkien, 1954–1955). glandulosa (Desv.) Proctor. Etymology: Named for
Thelypteris fideleae (Rakotondr.) Christenh., comb. the wizard Gandalf of the Istari order and leader of
nov. Basionym: Pronephrium fideleae Rakotondr., the Fellowship of the Ring (Tolkien, 1937–1955).
Adansonia, sér. 3, 34: 226. 2012, as ‘fidelei’ Thelypteris gaudichaudii (Holttum) Christenh., comb.
Thelypteris filix-femina Christenh., nom. nov. Replaced nov. Basionym: Sphaerostephanos gaudichaudii
synonym: Glaphyropteridopsis jinfushanensis Ching Holttum, Kew Bull. 34: 228. 1979.
& Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 325. 1999, Thelypteris giluwensis (Holttum) Christenh., comb.
not: Thelypteris jinfoshanensis (Ching & Z.Y.Liu) nov. Basionym: Pronephrium giluwense Holttum, Fl.
Christenh. Note and etymology: the basionym is an Males., Ser. 2, Pterid. l: 520. 1982.
orthographic variant and if maintained will cause Thelypteris gimlii Christenh., nom. nov. Replaced
confusion, hence a new name, based on the delicate synonym: Goniopteris subdimorpha Salino,
leaves of this species, was chosen. It translates as Phytotaxa 255: 253. 2015, not: Thelypteris
female fern fern, a synonym of Thelypteris which subdimorpha (Copel.) C.F.Reed. Etymology: Named
means the same in Greek. for dwarf warrior Gimli, son of Glóin, member of the
Thelypteris flavoviridis (Holttum) Christenh., comb. Fellowship of the Ring (Tolkien, 1937–1955).
nov. Basionym: Sphaerostephanos flavoviridis Thelypteris glorfindelii Christenh., nom. nov. Replaced
Holttum, Fl. Males., Ser. 2, Pterid. l: 450. 1982. synonym: Nephrodium debile Baker, J. Bot. 18:
Thelypteris florencei (A.R.Sm. & Lorence) Christenh., 212. 1880, not: Thelypteris debilis (Mett.) C.F.Reed.
comb. nov. Basionym: Cyclosorus florencei A.R.Sm. Etymology: Named for the elf Glorfindel, Lord of
& Lorence, PhytoKeys 4: 40. 2011. Gondolin in Middle-earth (Tolkien, 1937–1955).
Thelypteris foliolosa (Holttum) Christenh., comb. nov. Thelypteris gollum Christenh., nom. nov. Replaced
Basionym: Sphaerostephanos foliolosus Holttum, Fl. synonym: Pseudocyclosorus obliquus Ching ex
Males., Ser. 2, Pterid. l: 488. 1982. Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 331. 1999, not:
Thelypteris formosa (Nakaike) Christenh., comb. Thelypteris obliqua (Holttum) Christenh. Etymology:
nov. Basionym: Parathelypteris formosa Nakaike, J. Named for the stoor hobbit Sméagol, who was
Nippon Fernist Club 3 (Suppl. 2): 137. 2004. corrupted by the One Ring and was later named

Gollum after his habit of making a gutteral noise in Finnish), fictional electric characters in the
(Tolkien, 1937–1955). Moominvalley stories of Tove Jansson (1945–1970).
Thelypteris gondorica Christenh., nom. nov. Replaced Thelypteris hekouensis (Ching ex Y.X.Lin) Christenh.,
synonym: Glaphyropteridopsis mollis Ching & comb. nov. Basionym: Pronephrium hekouensis
Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 324. 1999, not: Ching ex Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 353.
Thelypteris mollis (Mett.) R.M.Tryon. Etymology: 1999.
Named for the fictional kingdom of Gondor in Thelypteris hellwigensis (Holttum) Christenh., comb.
Middle-earth (Tolkien, 1937–1955). nov. Basionym: Sphaerostephanos hellwigensis
Thelypteris gongshanensis (Y.X.Lin) Christenh., comb. Holttum, Fl. Males., Ser. 2, Pterid. l: 500. 1982.
nov. Basionym: Pseudocyclosorus gongshanensis Thelypteris helmsdeepensis Christenh., nom. nov.
Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 328. 1999. Replaced synonym: Cyclosorus oppositipinnus
Thelypteris gracilis (Ching ex Y.X.Lin) Christenh., Ching & Z.Y.Liu, Bull. Bot. Res., Harbin 4: 15. 1984,
comb. nov. Basionym: Pronephrium gracilis Ching ex not Thelypteris oppositipinna (Alderw.) Ching.
Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 352. 1999. Etymology: Named for Helms Deep, a fictional land
Thelypteris grandescens (Holttum) Christenh., comb. in Middle-earth (Tolkien (1945–1955).
nov. Basionym: Sphaerostephanos grandescens Thelypteris hemuli Christenh., nom. nov. Replaced
Holttum, Fl. Males., Ser. 2, Pterid. l: 465. 1982. synonym: Cyclosorus macrophyllus Ching & Z.Y.Liu,
Thelypteris grandissima (Ching & K.H.Shing) Bull. Bot. Res., Harbin 4: 16. 1984, not: Thelypteris
Christenh., comb. nov. Basionym: Cyclosorus macrophylla (Kunze) C.V.Morton. Etymology: Named
grandissimus Ching & K.H.Shing, Fl. Fujianica 1: 599. for a Hemulen (Hemuli in Finnish), a fictional
1982. character in the Moominvalley stories of Tove
Jansson (1945–1970).
Thelypteris grima Christenh., nom. nov. Replaced
synonym: Phegopteris nervosa Fée, Mém. Foug. Thelypteris hendersonii (Holttum) Christenh., comb.
5: 244. 1850, not: Thelypteris nervosa (Klotzsch) nov. Basionym: Sphaerostephanos hendersonii
R.M.Tryon. Etymology: Named for Grima the Holttum, Fl. Males., Ser. 2, Pterid. l: 493. 1982.
Wormtongue, a fictional malicious character in Thelypteris hernaezii (Holttum) Christenh., comb. nov.
Tolkien’s (1954–1955) Middle-earth legendarium. Basionym: Sphaerostephanos hernaezii Holttum, Fl.
Thelypteris grokei Christenh., nom. nov. Replaced Males., Ser. 2, Pterid. l: 459. 1982.
synonym: Cyclogramma tibetica Ching & S.K.Wu, Thelypteris hirsuta (Kunze ex Mett.) Christenh., comb.
Fl. Xizang. 1: 157. 1983, not: Thelypteris tibetica nov. Basionym: Aspidium hirsutum Kunze ex Mett.,
(Ching) Fraser-Jenk. Etymology: Named for the Abh. Senckenberg. Naturf. Gesell. 2: 391. 1858.
Groke (English), a fictional ghost-like creature in the Thelypteris hirtipes (K.H.Shing & C.F.Zhang) Christenh.,
Moominvalley stories of Tove Jansson (1945–1970). comb. nov. Basionym: Cyclosorus hirtipes K.H.Shing
Thelypteris guangxianensis (Ching ex Y.X.Lin) & C.F.Zhang, Fl. Reipubl. Popularis Sin. 4: 345. 1999.
Christenh., comb. nov. Basionym: Pseudocyclosorus Thelypteris hoalensis (Holttum) Christenh., comb. nov.
guanxianensis Ching ex Y.X.Lin, Fl. Reipubl. Popularis Basionym: Sphaerostephanos hoalensis Holttum, Fl.
Sin. 4: 331. 1999. Males., Ser. 2, Pterid. l: 488. 1982.
Thelypteris guangxiensis (Y.X.Lin) Christenh., comb. Thelypteris hobbitonica Christenh., nom. nov. Replaced
nov. Basionym: Pseudocyclosorus guangxiensis synonym: Cyclosorus sledgei Panigrahi & Sarn.Singh,
Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 327. 1999. Ferns & Fern-Allies Arunachal Pradesh 2: 704. 2005,
Thelypteris gymnorachis (Holttum) Christenh., comb. not: Thelypteris sledgei Fraser-Jenk. Etymology:
nov. Basionym: Sphaerostephanos gymnorachis Named for Hobbiton, a fictional village in the Shire of
Holttum, Fl. Males., Ser. 2, Pterid. l: 479. 1982. Middle-earth (Tolkien, 1937–1955).
Thelypteris haisuli Christenh., nom. nov. Replaced Thelypteris hobbitorum Christenh., nom. nov. Replaced
synonym: Sphaerostephanos pilosissimus Holttum, synonym: Nephrodium zeylanicum Fée, Mém. Foug.
Fl. Males., Ser. 2, Pterid. l: 504. 1982, not: Thelypteris 10: 42. 1865, not: Thelypteris zeylanica Ching.
pilosissima C.V.Morton. Etymology: Named for Etymology: Named for the hobbits, a fictional race
Haisuli (Finnish), a foul-smelling creature in the of halflings that live in Middle-earth (Tolkien, 1937–
Moominvalley stories of Tove Jansson (1945–1970). 1955).
Thelypteris hattivattiorum Christenh., nom. nov. Thelypteris hochreutineri (C.Chr.) Christenh., comb.
Replaced synonym: Cyclosorus multisorus Ching, Fl. nov. Basionym: Dryopteris hochreutineri C.Chr.,
Reipubl. Popularis Sin. 4: 334. 1999, not: Thelypteris Annuaire Conserv. Jard. Bot. Genève 15–16: 188.
multisora (C.Chr. ex C.Chr. & Holttum) C.F.Reed. 1912.
Etymology: Named for the Hattifatteners (Hattivatti

31
Thelypteris holttumii (Sarn.Singh & Panigrahi) Thelypteris jermyi (Holttum) Christenh., comb. nov.
Christenh., comb. nov. Basionym: Cyclosorus Basionym: Pneumatopteris jermyi Holttum, Blumea
holttumii Sarn.Singh & Panigrahi, Ferns & Fern-Allies 21: 310. 1973.
Arunachal Pradesh 2: 682. 2005. Thelypteris jijiangensis (Ching ex Y.X.Lin) Christenh.,
Thelypteris hopeana (Baker) Christenh., comb. nov. comb. nov. Basionym: Pseudocyclosorus jijiangensis
Basionym: Nephrodium hopeanum Baker, Syn. Filic. Ching ex Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 330.
ed. 2: 494. 1874. 1999.
Thelypteris hosokawana Christenh., nom. nov. Thelypteris jinfoshanensis (Ching & Z.Y.Liu) Christenh.,
Replaced synonym: Glaphyropteris rupicola Hosok., comb. nov. Basionym: Leptogramma jinfoshanensis
not: Thelypteris rupicola (C.Chr.) Ching. Etymology: Ching & Z.Y.Liu, Bull. Bot. Res., Harbin 4: 17. 1984.
This species was named for Takahide Hosokawa who Thelypteris jinghongensis (Ching ex K.H.Shing)
first described it. Christenh., comb. nov. Basionym: Cyclosorus
Thelypteris huishuiensis (Ching ex Y.X.Lin) Christenh., jinghongensis Ching ex K.H.Shing, Fl. Reipubl.
comb. nov. Basionym: Leptogramma huishuiensis Popularis Sin. 4: 217. 1999.
Ching ex Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 323. Thelypteris johannae (Holttum) Christenh., comb. nov.
1999. Basionym: Pseudocyclosorus johannae Holttum, J. S.
Thelypteris humbertii (Holttum) Christenh., comb. African Bot. 40: 138. 1974.
nov. Basionym: Pneumatopteris humbertii Holttum, Thelypteris kalkmanii (Holttum) Christenh., comb. nov.
Blumea 21: 313. 1973. Basionym: Sphaerostephanos kalkmanii Holttum, Fl.
Thelypteris humilis (Holttum) Christenh., comb. nov. Males., Ser. 2, Pterid. l: 454. 1982.
Basionym: Sphaerostephanos humilis Holttum, Fl. Thelypteris kerintjiensis (Holttum) Christenh., comb.
Males., Ser. 2, Pterid. l: 468. 1982. nov. Basionym: Pneumatopteris kerintjiensis
Thelypteris hypolarix Christenh., nom. nov. Replaced Holttum, Blumea 21: 312. 1973.
synonym: Pseudocyclosorus emeiensis Ching ex Thelypteris kiauensis (C.Chr.) Christenh., comb. nov.
Y.X.Lin, Fl. Reipubl. Pop. Sin. 4: 330. 1999, not: Basionym: Dryopteris kiauense C.Chr., Dansk Bot.
Thelypteris emeiensis (Y.X.Lin) Christenh. Etymology: Ark. 9: 64. 1937.
This species grows in larch (Larix) forests, hence the Thelypteris kjellbergii (Holttum) Christenh., comb.
name, hypo meaning ‘under’. nov. Basionym: Pronephrium kjellbergii Holttum,
Thelypteris immucosa (Holttum) Christenh., comb. Blumea 20: 117. 1972.
nov. Basionym: Sphaerostephanos immucosus Thelypteris koordersii (Christ) Christenh., comb. nov.
Holttum, Blumea 33: 404. 1988. Basionym: Aspidium koordersii Christ, Ann. Jard. Bot.
Thelypteris inaequilobata (Holttum) Christenh., comb. Buitenzorg 15: 128. 1897.
nov. Basionym: Nannothelypteris inaequilobata Thelypteris kostermansii (Holttum) Christenh., comb.
Holttum, Kalikasan 2: 67. 1973. nov. Basionym: Plesioneuron kostermansii Holttum,
Thelypteris incisa (Holttum) Christenh., comb. nov. Fl. Males., Ser. 2, Pterid. l: 406. 1982.
Basionym: Pneumatopteris incisa Holttum, Blumea Thelypteris krameri (Sarn.Singh & Panigrahi)
21: 317. 1973. Christenh., comb. nov. Basionym: Cyclosorus
Thelypteris indrapurae (Holttum) Christenh., comb. krameri Sarn.Singh & Panigrahi, Ferns & Fern-Allies
nov. Basionym: Sphaerostephanos indrapurae Arunachal Pradesh 2: 687. 2005.
Holttum, Fl. Males., Ser. 2, Pterid. l: 475. 1982. Thelypteris kundipensis (Holttum) Christenh., comb.
Thelypteris isildurii Christenh., nom. nov. Replaced nov. Basionym: Plesioneuron kundipense Holttum,
synonym: Nephrodium angustifolium C.Presl, Blumea 22: 247. 1975.
Epimel. Bot.: 48. 1851, not: Thelypteris angustifolia Thelypteris lamii (Holttum) Christenh., comb. nov.
(Willd.) Proctor. Etymology: Named for Isildur, son of Basionym: Sphaerostephanos lamii Holttum, Fl.
Elendil of Gondor, who cut the One Ring of Sauron’s Males., Ser. 2, Pterid. l: 455. 1982.
hand, a fictional character in the Middle-earth
Thelypteris lastreoides (C.Presl) Christenh., comb. nov.
legendarium (Tolkien, 1937–1955).
Basionym: Pronephrium lastreoides C.Presl, Epimel.
Thelypteris isomorpha (Holttum) Christenh., comb. Bot.: 259. 1851.
nov. Basionym: Sphaerostephanos isomorphus
Thelypteris latisquamata (Holttum) Christenh., comb.
nov. Basionym: Pneumatopteris latisquamata
Thelypteris japenensis (Holttum) Christenh., comb. Holttum, Blumea 21: 305. 1973.
nov. Basionym: Pneumatopteris japenensis Holttum,
Fl. Males., Ser. 2, Pterid. l: 426. 1982.

Thelypteris lawakii (Holttum) Christenh., comb. Thelypteris marojejyensis (Rakotondr.) Christenh.,

nov. Basionym: Pneumatopteris lawakii Holttum, comb. nov. Basionym: Pronephrium marojejyensis
Fl. Males., Ser. 2, Pterid. l: 430. 1982. Replaced Rakotondr., Adansonia 34: 230. 2012.
synonym: Polystichum truncatum Gaudich., not: Thelypteris marquesica (Holttum) Christenh., comb.
Thelypteris truncata (Poir.) K.Iwats. nov. Basionym: Plesioneuron marquesicum Holttum,
Thelypteris legolas Christenh., nom. nov. Replaced Allertonia 1: 192. 1977.
synonym: Stegnogramma crenata Holttum, Blumea Thelypteris medlerae (W.N.Takeuchi) Christenh.,
33: 404. 1998, not: Thelypteris crenata A.R.Sm. comb. nov. Basionym: Pneumatopteris medlerae
& Lellinger. Etymology: Named for woodland elf W.N.Takeuchi, Blumea 52: 154. 2007.
Legolas, member of the Fellowship of the Ring Thelypteris medogensis (Ching & S.K.Wu) Christenh.,
(Tolkien, 1937–1955). comb. nov. Basionym: Cyclosorus medogensis Ching
Thelypteris lenormandii (C.Chr) Christenh., comb. nov. & S.K.Wu, Fl. Xizang. 1: 171. 1983.
Basionym: Dryopteris lenormandii C.Chr., Index Filic.: Thelypteris medusella (Holttum) Christenh., comb.
274. 1905. nov. Basionym: Plesioneuron medusella Holttum, Fl.
Thelypteris lindleyi (W.N.Takeuchi) Christenh., comb. Males., Ser. 2, Pterid. l: 400. 1982.
nov. Basionym: Amphineuron lindleyi W.N.Takeuchi, Thelypteris melanorachis (Holttum) Christenh., comb.
Harvard Pap. Bot. 10: 105. 2005. nov. Basionym: Sphaerostephanos melanorachis
Thelypteris linearis (Ching & K.H.Shing ex Y.X.Lin) Holttum, Fl. Males., Ser. 2, Pterid. l: 457. 1982.
Christenh., comb. nov. Basionym: Pseudocyclosorus Thelypteris menadensis (Holttum) Christenh., comb.
linearis Ching & K.H.Shing ex Y.X.Lin, Fl. Reipubl. nov. Basionym: Sphaerostephanos menadensis
Popularis Sin. 4: 328. 1999. Holttum, Fl. Males., Ser. 2, Pterid. l: 469. 1982.
Thelypteris lithophila (Holttum) Christenh., comb. Thelypteris mengeniana (Holttum) Christenh., comb.
nov. Basionym: Pneumatopteris lithophila Holttum, nov. Basionym: Sphaerostephanos mengenianus
Fl. Males., Ser. 2, Pterid. l: 422. 1982. Holttum, Fl. Males., Ser. 2, Pterid. l: 491. 1982.
Thelypteris longqishanensis (K.H.Shing) Christenh., Thelypteris merryi Christenh., nom. nov. Replaced
comb. nov. Basionym: Cyclosorus longqishanensis synonym: Stegnogramma aspidioides Blume, Enum
K.H.Shing, Fl. Reipubl. Popularis Sin. 4: 219. 1999. Pl. Jav. 173. 1828, not: Thelypteris aspidioides
Thelypteris lorzingii (Holttum) Christenh., comb. nov. (Willd.) R.M.Tryon. Etymology: Named for Meriadoc
Basionym: Chingia lorzingii Holttum, Fl. Males., Ser. (Merry) Brandybuck, a hobbit who joined the
2, Pterid. l: 393. 1982. Fellowship of the Ring (Tolkien, 1937–1955).
Thelypteris lucbanii (Holttum) Christenh., comb. nov. Thelypteris michaelis (Holttum) Christenh., comb.
Basionym: Sphaerostephanos lucbanii Holttum, Fl. nov. Basionym: Pneumatopteris michaelis Holttum,
Males., Ser. 2, Pterid. l: 463. 1982. Blumea 21: 313. 1973.
Thelypteris lushanensis (Ching ex Y.X.Lin) Christenh., Thelypteris micropaleata (Holttum) Christenh., comb.
comb. nov. Basionym: Pseudocyclosorus lushanensis nov. Basionym: Pneumatopteris micropaleata
Ching ex Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 328. Holttum, Blumea 21: 319. 1973.
1999. Thelypteris micropinnata (Holttum) Christenh., comb.
Thelypteris lushuiensis (Y.X.Lin) Christenh., comb. nov. nov. Basionym: Pronephrium micropinnatum
Basionym: Pseudocyclosorus lushuiensis Y.X.Lin, Fl. Holttum, Blumea 20: 108. 1972.
Reipubl. Popularis Sin. 4: 327. 1999. Thelypteris middle-earthica Christenh., nom. nov.
Thelypteris maguanensis (Ching ex K.H.Shing) Replaced synonym: Christella pacifica Holttum, Kew
Christenh., comb. nov. Basionym: Cyclogramma Bull. 31: 320. 1976, not: Thelypteris pacifica C.F.Reed.
maguanensis Ching ex K.H.Shing, Fl. Reipubl. Etymology: Named for the fictional continent of
Popularis Sin. 4: 322. 1999. Middle-earth in Tolkien’s legendarium (1937–1955).
Thelypteris major (Copel.) Christenh., comb. nov. Thelypteris millarae (Holttum) Christenh., comb. nov.
Basionym: Haplodictyum majus Copel., Elmer’s Basionym: Pronephrium millarae Holttum, Blumea
Leaflets 9: 3110. 1920. 20: 115. 1972.
Thelypteris makassarica (Holttum) Christenh., comb. Thelypteris minahassae (Holttum) Christenh., comb.
nov. Basionym: Sphaerostephanos makassaricus nov. Basionym: Pronephrium minahassae Holttum,
Holttum, Fl. Males., Ser. 2, Pterid. l: 498. 1982. Blumea 20: 119. 1972.
Thelypteris marattioides (Alston) Christenh., comb. Thelypteris minasmorgulensis Christenh., nom. nov.
nov. Basionym: Dryopteris marattioides Alston, J. Replaced synonym: Cyclosorus castaneus A.R.Sm. &
Bot. 78: 227. 1940. Lorence, PhytoKeys 4: 39. 2011, not: Thelypteris

33
castanea (Tagawa) Ching. Etymology: Named for Pl. Javae 2: 148. 1828, not: Thelypteris affinis (C.Presl)
Minas Morgul (Tower of Black Sorcery) a fictional C.V.Morton. Etymology: Named for the Moomins (or
city in Gondor (Tolkien, 1937–1955). Mumins in Swedish), main characters in the Moomin
Thelypteris minastirithensis Christenh., nom. nov. valley stories by Tove Janssen (1945–1970).
Replaced synonym: Cyclosorus christii Copel., Fern Thelypteris muminmammae Christenh., nom. nov.
Fl. Philipp. 2: 362. 1960, not: Thelypteris christii Replaced synonym: Cyclosorus elatus Ching ex
(C.Chr.) C.F.Reed. Etymology: Named for Minas Tirith K.H.Shing, Fl. Reipubl. Popularis Sin. 4: 342. 1999,
(Tower of Guard), a fictional city and the heavily not: Thelypteris elata Schelpe, nor: Cyclosorus
fortified capital of Gondor (Tolkien, 1937–1955). elatus (Mett. ex Kuhn) Alston, nom. illeg. Etymology:
Thelypteris mindorensis (Holttum) Christenh., comb. Named for Moominmamma (Muminmamma in
nov. Basionym: Sphaerostephanos mindorensis Swedish), mother of the main character in the
Holttum, Kalikasan 4: 54. 1975. Moominvalley stories by Tove Janssen (1945–1970).
Thelypteris mingchegensis (Ching) Christenh., comb. Thelypteris murkelensis (M.Kato) Christenh., comb.
nov. Basionym: Dictyocline mingchegensis Ching, nov. Basionym: Plesioneuron murkelense M.Kato,
Acta Phytotax. Sin. 8: 334. 1963. Bull. Natl. Mus. Nat. Sci., Tokyo, B 33: 20. 2007.
Thelypteris mingendensis (Gilli) Christenh., comb. Thelypteris my Christenh., nom. nov. Replaced
nov. Basionym: Lastrea mingendensis Gilli, Ann. synonym: Aspidium glandulosum Blume, Enum.
Naturhist. Mus. Wien 81: 24. 1978. Pl. Javae 2: 144. 1828, not: Thelypteris glandulosa
(Desv.) Proctor. Etymology: Named for Little My, a
Thelypteris minuscula (Maxon) Christenh., comb. nov.
mischievous little girl in the Moominvalley stories by
Basionym: Dryopteris minuscula Maxon, Bull. Misc.
Tove Janssen (1945–1970).
Inform. Kew 1932: 135. 1932.
Thelypteris nakaikeana (R.D.Dixit & Balkrishna)
Thelypteris mjobergii (Holttum) Christenh., comb. nov.
Christenh., comb. nov. Basionym: Pronephrium
Basionym: Sphaerostephanos mjobergii Holttum, Fl.
nakaikeanum R.D.Dixit & Balkrishna, Indian Fern J.
Males., Ser. 2, Pterid. l: 494. 1982.
7: 2. 1990.
Thelypteris modesta (Holttum) Christenh., comb. nov.
Thelypteris nakaikei (Holttum) Christenh., comb. nov.
Basionym: Christella modesta Holttum, J. S. African
Basionym: Sphaerostephanos nakaikei Holttum, Fl.
Bot. 40: 144. 1974.
Thelypteris mordorica Christenh., nom. nov.
Thelypteris nanxiensis (Ching ex K.H.Shing) Christenh.,
Replaced synonym: Dictyoclina sagittifolia Ching,
comb. nov. Basionym: Cyclosorus nanxiensis Ching
Acta Phytotax. Sin. 8: 335. 1963, not: Thelypteris
ex K.H.Shing, Fl. Reipubl. Popularis Sin. 4: 343. 1999.
sagittifolia (Blume) C.F.Reed. Etymology: Named
for the fictional land Mordor (Black Land) in Thelypteris nazgulorum Christenh., nom. nov.
southwestern Middle-earth (Tolkien, 1937–1955). Replaced synonym: Parathelypteris chingii K.H.Shing
& J.F.Cheng, Jiangxi Sci. 8: 44. 1990, not: Thelypteris
Thelypteris moriana Christenh., nom. nov. Replaced
chingii (Z.Y.Liu) Christenh. Etymology: Named for the
synonym: Cyclosorus attenuatus Ching ex K.H.Shing,
Nazgûl, Ringwraiths or Black Riders, nine servants
Fl. Reipubl. Sin. 4: 341. 1999. Replaced synonym:
of Sauron in Tolkien’s (1954–1955) Middle-earth
Cyclosorus mollissimus Ching ex K.H.Shing, Fl.
legendarium.
Reipubl. Sin. 4: 347. 1999, not: Thelypteris attenuata
(Kuntze) C.V.Morton, nor: Thelypteris mollissima Thelypteris neohibernica Christenh., nom. nov.
(Kunze) Thapa. Etymology: Named for a fictional, Replaced synonym: Christella minima Holttum, Kew
large underground tunnel complex carved out of the Bull. 31: 304. 1976, not: Thelypteris minima A.R.Sm.
Misty Mountains in Middle-earth (Tolkien, 1937– & M.Kessler. Etymology: Named for New Ireland,
1955). where this species is found.
Thelypteris morotaiensis (Holttum) Christenh., comb. Thelypteris nigrescens (Ching ex Shing) Christenh.,
nov. Basionym: Sphaerostephanos morotaiensis comb. nov. Basionym: Parathelypteris nigrescens
Holttum, Fl. Males., Ser. 2, Pterid. l: 453. 1982. Ching ex K.H.Shing, Fl. Reipubl. Popularis Sin. 4: 321.
1999.
Thelypteris moseleyi (Holttum) Christenh., comb. nov.
Basionym: Sphaerostephanos moseleyi Holttum, Fl. Thelypteris nigricans (Salino & T.E.Almeida) Christenh.,
Males., Ser. 2, Pterid. l: 460. 1982. comb. nov. Basionym: Goniopteris nigricans Salino &
T.E.Almeida, PhytoKeys 57: 41. 2015.
Thelypteris muluensis (Holttum) Christenh., comb.
nov. Basionym: Sphaerostephanos muluensis Thelypteris norrisii (Rosenst.) Christenh., comb. nov.
Holttum, Fl. Males., Ser. 2, Pterid. l: 499. 1982. Basionym: Dryopteris norrisii Rosenst., Meded. Rijks-
Herb. 31: 8. 1917.
Thelypteris muminorum Christenh., nom. nov.
Replaced synonym: Aspidium affine Blume, Enum.

Thelypteris novae-brittaniae (Holttum) Christenh., Thelypteris paraochthodes (Ching ex K.H.Shing &

comb. nov. Basionym: Sphaerostephanos novae- J.E.Cheng) Christenh., comb. nov. Basionym:
britanniae Holttum, Fl. Males., Ser. 2, Pterid. l: 477. Pseudocyclosorus paraochthodes Ching ex K.H.Shing
1982. & J.E.Cheng, Jiangxi Sci. 8(3): 43. 1990.
Thelypteris novae-caledoniae (Holttum) Christenh., Thelypteris paraphysophora (Alderw.) Christenh.,
comb. nov. Basionym: Pneumatopteris novae- comb. nov. Basionym: Dryopteris paraphysophora
caledoniae Holttum, Blumea 21: 313. 1973. Alderw., Bull. Jard. Bot. Buitenzorg 2: 143. 1920.
Thelypteris nudisora (Holttum) Christenh., comb. nov. Thelypteris parksii (Ballard) Christenh., comb. nov.
Basionym: Sphaerostephanos nudisorus Holttum, Fl. Basionym: Dryopteris parksii F.Ballard, Bull. Misc.
Males., Ser. 2, Pterid. l: 476. 1982. Inform. Kew 1937(6): 347. 1937.
Thelypteris numenorica Christenh., nom. nov. Thelypteris parvifolia (Ching) Christenh., comb. nov.
Replaced synonym: Glaphyropteridopsis villosa Basionym: Cyclosorus parvifolius Ching, Fl. Fujianica
Ching & W.M.Chu, Fl. Reipubl. Popularis Sin. 4: 1: 155. 1982.
325. 1999, not: Thelypteris villosa (Link) C.F.Reed. Thelypteris parviloba (Ching & K.H.Shing ex K.H.Shing)
Etymology: Named for Númenor or Westernesse, a Christenh., comb. nov. Basionym: Cyclosorus
fictional, large island to the west of Middle-earth in parvilobus Ching, Fl. Reipubl. Popularis Sin. 4: 338.
Tolkien (1937–1955). 1999.
Thelypteris obliqua (Holttum) Christenh., comb. nov. Thelypteris patentipinna (Holttum) Christenh., comb.
Basionym: Pneumatopteris obliqua Holttum, Blumea nov. Basionym: Pneumatopteris patentipinna
21: 309. 1973. Holttum, Blumea 21: 307. 1973.
Thelypteris omatiana (Holttum) Christenh., comb. nov. Thelypteris pauciloba (Ching ex K.H.Shing) Christenh.,
Basionym: Sphaerostephanos omatianus Holttum, comb. nov. Basionym: Parathelypteris pauciloba
Fl. Males., Ser. 2, Pterid. l: 483. 1982. Ching ex K.H.Shing, Fl. Reipubl. Popularis Sin. 4: 321.
Thelypteris orientalis (Ching ex K.H.Shing) Christenh., 1999.
comb. nov. Basionym: Cyclosorus orientalis Ching ex Thelypteris paucipaleata (Holttum) Christenh., comb.
K.H.Shing, Fl. Reipubl. Popularis Sin. 4: 332. 1999. nov. Basionym: Chingia paucipaleata Holttum,
Thelypteris orizabae (Fée) Christenh., comb. nov. Kalikasan 3: 29. 1974.
Basionym: Aspidium orizabae Fée ex E.Fourn., Thelypteris pauciserrata (Ching & C.F.Zhang)
Mexic. Pl. 1: 95. 1872. Christenh., comb. nov. Basionym: Cyclosorus
Thelypteris pallida (Ching & W.M.Chu) Christenh., pauciserratus Ching & C.F.Zhang, Bull. Bot. Res.,
comb. nov. Basionym: Glaphyropteridopsis pallida Harbin 3: 8. 1983.
Ching & W.M.Chu, Fl. Reipubl. Popularis Sin. 4: 325. Thelypteris peekelii (Alderw.) Christenh., comb. nov.
1999. Basionym: Dryopteris peekelii Alderw., Bull. Dépt.
Thelypteris pandeorum Christenh., nom. nov. Replaced Agric. Indes Néerl. 18: 7. 1908.
synonym: Pronephrium kumaunicum P.C.Pande & Thelypteris peltata (Alderw.) Christenh., comb. nov.
H.C.Pande, Ill. Fern Fl. Kumaon Himalaya 1: 180. Basionym: Dryopetris peltata Alderw., Bull. Jard. Bot.
2003, not: Thelypteris ×kumaunica (Holttum) Fraser- Buitenzorg ser. 2, 16: 12. 1914.
Jenk. Etymology: Named for Indian botanists P.C. Thelypteris pendens (D.D.Palmer) Christenh.,
Pande and H.C. Pande, who first described this comb. nov. Basionym: Pneumatopteris pendens
species. D.D.Palmer, Amer. Fern J. 95: 81. 2005.
Thelypteris panigrahii Christenh., nom. nov. Replaced Thelypteris peramelensis (Holttum) Christenh., comb.
synonym: Metathelypteris krameri Sarn.Singh & nov. Basionym: Pronephrium peramelense Holttum,
Panigrahi, Ferns & Fern-Allies Arunachal Pradesh Blumea 20: 115. 1972.
2: 716. 2005, not: Thelypteris krameri (Sarn.Singh
Thelypteris peregrinii Christenh., nom. nov. Replaced
& Panigrahi) Christenh. Etymology: Named for
synonym: Sphaerostephanos exindusiatus Holttum,
Gopinath Panigrahi, who first described this species.
Fl. Males., Ser. 2, Pterid. l: 463. 1982, not: Thelypteris
Thelypteris papuana (Holttum) Christenh., comb. exindusiata W.H.Wagner. Etymology: Named for the
nov. Basionym: Pneumatopteris papuana Holttum, hobbit Peregrin Took or Pippin, who together with
Blumea 21: 311. 1973. his cousin Merry Brandybuck joined the Fellowship
Thelypteris paracuminata (Ching ex K.H.Shing & of the Ring in Tolkien (1937–1955).
J.F.Cheng) Christenh., comb. nov. Basionym: Thelypteris pergamacea (Holttum) Christenh., comb.
Cyclosorus paracuminatus Ching ex K.H.Shing & nov. Basionym: Pneumatopteris pergamacea
J.F.Cheng, Jiangxi Sci 8(3): 46. 1990. Holttum, Blumea 21: 315. 1973.

35
Thelypteris perrigida (Alderw.) Christenh., comb. nov. Thelypteris pseudofalciloba (W.M.Chu) Christenh.,
Basionym: Phegopteris perrigida Alderw., Bull. Jard. comb. nov. Basionym: Pseudocyclosorus
Bot. Buitenzorg ser. 2, XVI: 27. 1914. pseudofalcilobus W.M.Chu, Acta Bot. Yunnan., Suppl.
Thelypteris persquamifera (Alderw.) Christenh., comb. 5: 46. 1992.
nov. Basionym: Dryopteris persquamifera Alderw., Thelypteris pseudomegaphylla (Alderw.) Christenh.,
Bull. Jard. Bot. Buitenzorg 16: 10. 1914. comb. nov. Basionym: Dryopteris pseudomegaphylla
Thelypteris petiolulata (Ching ex K.H.Shing) Christenh., Alderw., Malay. Ferns Suppl.: 180. 1917.
comb. nov. Basionym: Metathelypteris petiolulata Thelypteris pseudorepens (Ching ex K.H.Shing)
Ching ex K.H.Shing, Fl. Reipubl. Popularis Sin. 4: 321. Christenh., comb. nov. Basionym: Pseudocyclosorus
1999. pseudorepens Ching ex K.H.Shing ex Ching, Acta
Thelypteris phanerophlebia (Baker) Christenh., comb. Phytotax. Sin. 3l: 571. 1993.
nov. Basionym: Nephrodium phanerophlebium Thelypteris psilophylla (Holttum) Christenh., comb.
Baker, Syn. Filic., ed. 2: 494. 1874. nov. Basionym: Pneumatopteris psilophylla Holttum,
Thelypteris platyloba (Holttum) Christenh., comb. Fl. Males., Ser. 2, Pterid. l: 427. 1982.
nov. Basionym: Plesioneuron platylobum Holttum, Thelypteris pullei (Holttum) Christenh., comb. nov.
Blumea 22: 246. 1975. Basionym: Plesioneuron pullei Holttum, Blumea 22:
Thelypteris plurivenosa (Holttum) Christenh., comb. 236. 1975.
nov. Basionym: Sphaerostephanos plurivenosus Thelypteris pullenii (Holttum) Christenh., comb. nov.
Holttum, Fl. Males., Ser. 2, Pterid. l: 496. 1982. Basionym: Sphaerostephanos pullenii Holttum, Fl.
Thelypteris polisiana (Holttum) Christenh., comb. nov. Males., Ser. 2, Pterid. l: 480. 1982.
Basionym: Sphaerostephanos polisianus Holttum, Thelypteris pycnosora (C.Chr.) Christenh., comb. nov.
Kalikasan 4: 65. 1975. Basionym: Dryopteris pycnosora C.Chr., Bull. Bishop
Thelypteris polyotica Christenh., nom. nov. Replaced Mus. 177: 83. 1943.
synonym: Christella multiauriculata Punetha, J. Thelypteris pygmaea (Ching & C.F.Zhang) Christenh.,
Indian Bot. Soc. 69: 177. 1990, not: Thelypteris comb. nov. Basionym: Cyclosorus pygmaeus Ching &
multicauriculata (Copel.) C.F.Reed. Etymology: From C.F.Zhang, Bull. Bot. Res., Harbin 3: 5. 1983.
Greek, meaning ‘many eared’ a translantion of the Thelypteris qingchengensis (Y.X.Lin) Christenh., comb.
Latin epithet. nov. Basionym: Pseudocyclosorus qingchengensis
Thelypteris posthumi (Holttum) Christenh., comb. nov. Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 329. 1999.
Basionym: Sphaerostephanos posthumi Holttum, Fl. Thelypteris qinlingensis (Ching ex K.H.Shing)
Males., Ser. 2, Pterid. l: 493. 1982, as ‘posthumii’, but Christenh., comb. nov. Basionym: Parathelypteris
named for Posthumus, hence only one i is needed. qinlingensis Ching ex K.H.Shing, Fl. Reipul. Popularis
Thelypteris potamia (Holttum) Christenh., comb. nov. Sin. 4: 320. 1999.
Basionym: Sphaerostephanos potamios Holttum, Fl. Thelypteris queenslandica (Holttum) Christenh., comb.
Males., Ser. 2, Pterid. l: 502. 1982. nov. Basionym: Amphineuron queenslandicum
Thelypteris pricei (Holttum) Christenh., comb. nov. Holttum, Kew Bull. 41: 518. 1986.
Basionym: Chingia pricei Holttum, Kalikasan 3: 22. Thelypteris rechingeri (Holttum) Christenh., comb.
1974. nov. Basionym: Parathelypteris rechingeri Holttum,
Thelypteris pseudoacuminata (Panigrahi & Sarn. Allertonia 1: 192. 1977.
Singh) Christenh., comb. nov. Basionym: Cyclosorus Thelypteris recondita (Holttum) Christenh., comb. nov.
pseudoacuminatus Panigrahi & Sarn.Singh, Ferns & Basionym: Sphaerostephanos reconditus Holttum,
Fern-Allies Arunachal Pradesh 2: 696. 2005. Fl. Males., Ser. 2, Pterid. l: 472. 1982.
Thelypteris pseudoarida (Ching ex K.H.Shing) Thelypteris reineckei (C.Chr.) Christenh., comb. nov.
Christenh., comb. nov. Basionym: Cyclosorus Basionym: Dryopteris reineckei C.Chr., Bull. Bishop
pseudoaridus Ching ex K.H.Shing, Fl. Reipubl. Mus. 177: 90. 1943.
Popularis Sin. 4: 349. 1999. Thelypteris rex Christenh., nom. nov. Replaced
Thelypteris pseudobalansae (Panigrahi & Sarn.Sing) synonym: Pronephrium macrophyllum Ching ex
Christenh., comb. nov. Basionym: Cyclosorus Y.X.Lin, Fl. Reipubl. Pop. Sin. 4: 352. 1999, not:
pseudobalansae Panigrahi & Sarn.Singh, Ferns & Thelypteris macrophylla (Kunze) C.V.Morton.
Fern-Allies Arunachal Pradesh 2: 699. 2005. Etymology: Named the king, rex, due to is large leave
Thelypteris pseudocuneata (Ching ex K.H.Shing) size.
Christenh., comb. nov. Basionym: Cyclosorus Thelypteris richardsii (Baker) Christenh., comb. nov.
pseudocuneatus Ching ex K.H.Shing, Fl. Reipubl. Basionym: Nephrodium richardsii Baker, Syn. Filic.,
Popularis Sin. 4: 348. 1999. ed. 2: 494. 1874.

Thelypteris rigidiloba (Holttum) Christenh., comb. Thelypteris scandens (Holttum) Christenh., comb. nov.
nov. Basionym: Plesioneuron rigidilobum Holttum, Basionym: Sphaerostephanos scandens Holttum,
Blumea 22: 248. 1975. Allertonia l: 211. 1977.
Thelypteris rivendellensis Christenh., nom. nov. Thelypteris scopulorum (Holttum) Christenh., comb.
Replaced synonym: Leptogramma celebica Ching, nov. Basionym: Pronephrium scopulorum Holttum,
Sinensia 7: 99. 1936, not: Thelypteris celebica Fl. Males., Ser. 2, Pterid. l: 532. 1982.
(Baker) C.F.Reed. Etymology: Named for the fictional Thelypteris seidleri (Salino) Christenh., comb. nov.
Elven realm of Rivendell in Middle-earth (Tolkien, Basionym: Goniopteris seidleri Salino, Phytotaxa
1937–1955). 255: 249. 2016.
Thelypteris rohanica Christenh., nom. nov. Replaced Thelypteris semicordata (Holttum) Christenh., comb.
synonym: Glaphyropteridopsis glabrata Ching & nov. Basionym: Sphaerostephanos semicordatus
W.M.Chu ex Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: Holttum, Fl. Males., Ser. 2, Pterid. l: 486. 1982.
325. 1999, not: Thelypteris glabrata (Mett. ex Kuhn) Thelypteris semimetralis (Holttum) Christenh., comb.
Tardieu. Etymology: Named for the fictional kingdom nov. Basionym: Sphaerostephanos semimetralis
of Rohan in Middle-earth (Tolkien, 1937–1955). Holttum, Fl. Males., Ser. 2, Pterid. l: 486. 1982.
Thelypteris royenii (Holttum) Christenh., comb. nov. Thelypteris septempedalis (Alston) Christenh., comb.
Basionym: Plesioneuron royenii Holttum, Blumea 22: nov. Basionym: Dryopteris septempedalis Alston, J.
240. 1975. Bot. 78: 227. 1940.
Thelypteris rufescens (Holttum) Christenh., comb. Thelypteris seramensis (M.Kato) Christenh., comb.
nov. Basionym: Mesophlebion rufescens Holttum, nov. Basionym: Coryphopteris seramensis M.Kato,
Blumea 22: 230. 1975. Bull. Natl. Mus. Nat. Sci., Tokyo, B 33: 18. 2007.
Thelypteris salinoi (I.O.Moura & L.C.Moura) Christenh., Thelypteris setosa (Y.X.Lin) Christenh., comb. nov.
comb. nov. Basionym: Goniopteris salinoi I.O.Moura Basionym: Pronephrium setosum Y.X.Lin, Fl. Reipubl.
& L.C.Moura, Brittonia 68: 452. 2016. Popularis Sin. 4: 352. 1999.
Thelypteris samwisei Christenh., nom. nov. Replaced Thelypteris shadowfax Christenh., nom. nov. Replaced
synonym: Stegnogramma latipinna Ching ex synonym: Nephrodium canum Baker, Syn. Filic.: 267.
Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 350. 1999, not: 1867, not: Thelypteris cana (J.Sm.) Ching. Etymology:
Thelypteris latipinna (Benth.) K.Iwats. Etymology: Named for the fictional horse Shadowfax of Rohan,
Named for the hobbit Samwise Gamgee, a fictional chief of the Mearas in the Middle-earth legendarium
character who joined the Fellowship of the Ring in (Tolkien, 1937–1955).
Tolkien’s (1937–1955) legendarium.
Thelypteris shelob Christenh., nom. nov. Replaced
Thelypteris sandsii (Holttum) Christenh., comb. nov. synonym: Pneumatopteris dilatata Holttum, Blumea
Basionym: Plesioneuron sandsii Holttum, Fl. Males., 33: 401. 1988, not: Thelypteris dilatata House.
Ser. 2, Pterid. 2: 407. 1982. Etymology: Named for Shelob, a fictional giant
Thelypteris santomasii (Holttum) Christenh., comb. spider in the Middle-earth legendarium (Tolkien,
nov. Basionym: Sphaerostephanos santomasii 1937–1955).
Holttum, Kalikasan 4: 62 (1975) Thelypteris shimenensis (K.H.Shing & C.M.Zhang)
Thelypteris sarasinorum (Holttum) Christenh., comb. Christenh., comb. nov. Basionym: Cyclosorus
nov. Basionym: Sphaerostephanos sarasinorum shimenensis K.H.Shing & C.M.Zhang, Keys Vasc. Pl.
Holttum, Fl. Males., Ser. 2, Pterid. l: 498. 1982. Wuling Mts.: 565. 1995.
Thelypteris saruman Christenh., nom. nov. Replaced Thelypteris shirensis Christenh., nom. nov. Replaced
synonym: Pronephrium hirsutum Ching ex Y.X.Lin, Fl. synonym: Christella nana Holttum, Kew Bull. 31:
Reipubl. Popularis Sin. 4: 351. 1999, as ‘Pronrphrium 304. 1976, not: Thelypteris nana A.Rojas. Etymology:
hirsutum’, not: Thelypteris hirsuta (Kunze ex Mett.) Named for the Shire, a fictional land in Middle-earth
Christenh. Etymology: Named for the wizard (Tolkien, 1937–1955).
Saruman the White, a fictional character in the Thelypteris shuangbaiensis (Ching ex Y.X.Lin)
Middle-earth legendarium (Tolkien 1937–1955). Christenh., comb. nov. Basionym: Pseudocyclosorus
Thelypteris sauron Christenh., nom. nov. Replaced shuangbaiensis Ching ex Y.X.Lin, Fl. Reipubl. Popularis
synonym: Lastrea attenuata Brack., U.S. Expl. Sin. 4: 327. 1999.
Exped., Filic. 16: 193. 1854, not: Thelypteris Thelypteris sibeliana (Holttum) Christenh., comb.
attenuata (Kuntze) C.V.Morton. Etymology: Named nov. Basionym: Pneumatopteris sibelana Holttum,
for fictional character Sauron the Necromancer, the Blumea 21: 301. 1973.
main antagonist in the Middle-earth legendarium
(Tolkien, 1937–1955).

37
Thelypteris sichuanensis (Y.X.Lin) Christenh., comb. Thelypteris stresemannii (Holttum) Christenh., comb.
nov. Basionym: Glaphyropteridopsis sichuanensis nov. Basionym: Sphaerostephanos stresemannii
Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 131, 324. 1999, Holttum, Fl. Males., Ser. 2, Pterid. l: 490. 1982.
as ‘sichanensis’. Thelypteris subacuta (Ching) Christenh., comb. nov.
Thelypteris ×silmarillion Christenh., nom. nov. Basionym: Cyclosorus subacutus Ching, Fl. Fujianica
Replaced synonym: Pronephrium ×interruptum 1: 598. 1982.
Holttum, Blumea 33: 403. 1988, not: Thelypteris Thelypteris subcalcarata (Alderw.) Christenh., comb.
interrupta (Willd.) B.C.Stone. Etymology: Named & stat. nov. Basionym: Phegopteris totta Mett. var.
for The Silmarillion, a collection of mythopoeic subcalcarata Alderw., Malay. Ferns Suppl. 1: 515.
works by English writer J. R. R. Tolkien, edited and 1916.
published posthumously by his son, Christopher Thelypteris subcanescens (Holttum) Christenh., comb.
Tolkien (1977). nov. Basionym: Sphaerostephanos subcanescens
Thelypteris sinensis (Ching & W.M.Chu) Christenh., Holttum, Blumea 33: 404. 1988.
comb. nov. Basionym: Craspedosorus sinensis Ching Thelypteris subcaudipinna (Sarn.Singh & Panigrahi)
& W.M.Chu, Acta Phytotax. Sin. 16: 26. 1978. Christenh., comb. nov. Basionym: Trigonospora
Thelypteris sinica (Ching ex Y.X.Lin) Christenh., comb. subcaudipinna Sarn.Singh & Panigrahi, Ferns & Fern-
nov. Basionym: Leptogramma sinica Ching ex Y.X.Lin, Allies Arunachal Pradesh 2: 777. 2005.
Fl. Reipubl. Popularis Sin. 4: 323. 1999. Thelypteris subcordata (Holttum) Christenh., comb.
Thelypteris sinodentata (Ching Z.Y.Liu) Christenh., nov. Basionym: Sphaerostephanos subcordatus
comb. nov. Basionym: Cyclosorus sinodentatus Holttum, Fl. Males., Ser. 2, Pterid. l: 469. 1982.
Ching & Z.Y.Liu, Bull. Bot. Res., Harbin 4: 13. 1984. Thelypteris subdentata (Holttum) Christenh., comb.
Thelypteris smeagol Christenh., nom. nov. Replaced nov. Basionym: Christella subdentata Holttum, Kew
synonym: Cyclosorus intermedius W.C.Shieh & Bull. 31: 335. 1976.
J.L.Tsai, J. Sci. Engin. 24: 8. 1987, not: Thelypteris Thelypteris subfalciloba (K.H.Shing ex Ching)
intermedia (Muhl. ex Willd.) House. Etymology: Christenh., comb. nov. Basionym: Pseudocyclosorus
Named for fictional character Sméagol, a Stoor subfalcilobus K.H.Shing ex Ching, Acta Phytotax. Sin.
Hobbit of the River Folk, who came upon the One 31: 571. 1993.
Ring (my precious), which corrupted him into
Thelypteris subglabra (Holttum) Christenh., comb.
Gollum (Tolkien, 1937–1955).
nov. Basionym: Plesioneuron subglabrum HoIttum,
Thelypteris smithii (Salino) Christenh., comb. nov. Blumea 22: 243. 1975.
Basionym: Goniopteris smithii Salino, Phytotaxa 255:
Thelypteris subjuncta (Baker) Christenh., comb. nov.
251. 2016.
Basionym: Nephrodium subjunctum Baker, Ann. Bot.
Thelypteris snufkin Christenh., nom. nov. Replaced (Oxford) 5: 321. 1891.
synonym: Chingia urens Holttum, Kalikasan 3: 28.
Thelypteris subopposita (Holttum) Christenh., comb.
1974, not: Thelypteris urens (Rosenst.) C.F.Reed.
nov. Basionym: Sphaerostephanos suboppositus
Etymology: Named for Snufkin (English), Moomin’s
best friend in the Moominvalley stories by Tove
Jansson (1945–1970). Thelypteris subornatipes (Sarn.Singh & Panigrahi)
Christenh., comb. nov. Basionym: Pseudocyclosorus
Thelypteris solsonica (Holttum) Christenh., comb.
subornatipes Sarn.Singh & Panigrahi, Ferns & Fern-
nov. Basionym: Pronephrium solsonicum Holttum,
Allies Arunachal Pradesh 2: 746. 2005.
Kalikasan 3: 197. 1974.
Thelypteris subterminalis (Holttum) Christenh., comb.
Thelypteris soluta (Holttum) Christenh., comb. nov.
nov. Basionym: Plesioneuron subterminale Holttum,
Basionym: Sphaerostephanos solutus Holttum, Fl.
Blumea 22: 241. 1975.
Thelypteris subtruncata (Bory) Christenh., comb.
Thelypteris splendens (Ching) Christenh., comb. nov.
nov. Basionym: Polypodium subtruncatum Bory in
Basionym: Glaphyropteridopsis splendens Ching,
Bélanger, Voy. Indes Or. 2: 32. 1833.
Acta Phytotax. Sin. 8: 322. 1963.
Thelypteris sudestica (Holttum) Christenh., comb. nov.
Thelypteris squamatella (Holttum) Christenh., comb.
Basionym: Sphaerostephanos sudesticus Holttum, Fl.
nov. Basionym: Sphaerostephanos squamatellus
Thelypteris sulawesiensis (K.Iwats.) Christenh., comb.
Thelypteris stenura (Holttum) Christenh., comb. nov.
& stat. nov. Basionym: Pronephrium rubicundum
Basionym: Plesioneuron stenura Holttum, Fl. Males.,
(Alderw.) Holttum subsp. sulawesiense K.lwats., Acta
Ser. 2, Pterid. l: 408. 1982.
Phytotax. Geobot. 28: 162. 1977.

Thelypteris sumbawensis (C.Chr.) Christenh., comb. Blumea 22: 240. 1975.

nov. Basionym: Dryopteris sumbawensis C.Chr., Thelypteris treebeardii Christenh., nom. nov. Replaced
Hedwigia 74: 231. 1934. synonym: Pneumatopteris sandwicensis Brack., U.S.
Thelypteris tahitensis (Holttum) Christenh., comb. Expl. Exped., Filic. 16: 26. 1854, not: Thelypteris
nov. Basionym: Plesioneuron tahitense Holttum, sandwicensis (Hook. & Arn.) Fosb. Etymology:
Blumea 22: 241. 1975. (Figures 5B–C) Named for the Ent Treebeard (or Fangorn), a fictional
Thelypteris tandikatensis (Alderw.) Christenh., comb. character in the Middle-earth legendarium (Tolkien,
nov. Basionym: Dryopteris tandikatensis Alderw., 1954–1955).
Bull. Jard. Bot. Buitenzorg Ser. 2, XI: 11. 1913. Thelypteris triangularis (R.S.Fern. & Salino) Christenh.,
Thelypteris tarningensis (Ching) Christenh., comb. comb. nov. Basionym: Meniscium triangularis
nov. Basionym: Cyclosorus tarningensis Ching, Wuyi R.S.Fern. & Salino, Phytotaxa 184: 4. 2014.
Sci. J. 1: 4. 1981. Thelypteris trichochlamys (Holttum) Christenh., comb.
Thelypteris telefominica (Holttum) Christenh., comb. nov. Basionym: Sphaerostephanos trichochlamys
nov. Basionym: Sphaerostephanos telefominicus Holttum, Fl. Males., Ser. 2, Pterid. l: 465. 1982.
Holttum, Fl. Males., Ser. 2, Pterid. l: 499. 1982. Thelypteris trimetralis (Holttum) Christenh., comb.
Thelypteris tenerior (Holttum) Christenh., comb. nov. nov. Basionym: Sphaerostephanos trimetralis
Basionym: Chingia tenerior Holttum, Fl. Males., Ser. Holttum, Fl. Males., Ser. 2, Pterid. l: 451. 1982.
2, Pterid. l: 395. 1982. Thelypteris tsoi (Ching) Christenh., comb. nov.
Thelypteris tenggerensis (Holttum) Christenh., comb. Basionym: Pseudocyclosorus tsoi Ching, Fl. Fujianica
nov. Basionym: Pseudocyclosorus tenggerensis 1: 619. 1991.
Holttum, Reinwardtia 8: 500. 1974. Thelypteris uaniensis (Holttum) Christenh., comb. nov.
Thelypteris thailandica (S.Linds.) Christenh., comb. Basionym: Sphaerostephanos uaniensis Holttum, Fl.
nov. Basionym: Cyclosorus thailandicus S.Linds., Males., Ser. 2, Pterid. l: 494. 1982.
Thai Forest Bull., Bot. 36: 47. 2008. Thelypteris unita (Kunze) Christenh., comb. nov.
Thelypteris theodenii Christenh., nom. nov. Replaced Basionym: Gymnogramma unita Kunze, Linnaea 18:
synonym: Pseudocyclosorus caudatus Holttum, 115. 1844.
Blumea 13: 133. 1965, not: Thelypteris caudata Thelypteris usambarensis (Holttum) Christenh., comb.
(Ching) C.F.Reed. Etymology: Named for Théoden, nov. Basionym: Pneumatopteris usambarensis
King of Rohan, a fictional character in the Middle- Holttum, Blumea 21: 312. 1973.
earth legendarium (Tolkien, 1945–1955). Thelypteris valinorica Christenh., nom. nov. Replaced
Thelypteris thysanoides (Holttum) Christenh., comb. synonym: Goniopteris rudis Ridl., Trans. Linn. Soc.
nov. Basionym: Pronephrium thysanoides Holttum, London, Bot. 9: 259. 1916, not: Thelypteris rudis
Fl. Males., Ser. 2, Pterid. l: 525. 1982. (Kunze) Proctor. Etymology: Named for the mythical
Thelypteris tibangensis (C.Chr.) Christenh., comb. nov. land Valinor, the Undying Lands, in Middle-earth
Basionym: Dryopteris tibangensis C.Chr., Dansk Bot. (Tolkien, 1954–1955).
Ark. 9: 66. 1937. Thelypteris vandervekenii (Pic.Serm.) Christenh.,
Thelypteris tildeniae (Holttum) Christenh., comb. nov. comb. nov. Basionym: Metathelypteris vandervekenii
Basionym: Amphineuron tildeniae Holttum, Blumea Pic.Serm., Bull. Jard. Bot. Belg. 53: 272. 1983.
23: 214. 1977. Thelypteris vaupelii (C.Chr.) Christenh., comb. nov.
Thelypteris timorensis (Holttum) Christenh., comb. Basionym: Dryopteris vaupelii C.Chr., Bull. Bishop
nov. Basionym: Christella timorensis Holttum, Kew Mus. 177: 89. 1943.
Bull. 31: 316. 1976. Thelypteris veitchii (Holttum) Christenh., comb. nov.
Thelypteris tobaica (Holttum) Christenh., comb. nov. Basionym: Sphaerostephanos veitchii Holttum,
Basionym: Pneumatopteris tobaica Holttum, Blumea Allertonia 1: 204. 1977.
21: 316. 1973. Thelypteris versteeghii (Holttum) Christenh., comb.
Thelypteris torrentis (Ching ex Y.X.Lin) Christenh., nov. Basionym: Pneumatopteris versteeghii Holttum,
comb. nov. Basionym: Pseudocyclosorus torrentis Blumea 21: 321. 1973.
Ching ex Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 326. Thelypteris walkeri (Holttum) Christenh., comb. nov.
1999. Basionym: Pneumatopteris walkeri Holttum, Blumea
Thelypteris trachyphylla (Holttum) Christenh., comb. 21: 323. 1973.
nov. Basionym: Pronephrium trachyphyllum Thelypteris wauensis (Holttum) Christenh., comb. nov.
Holttum, Fl. Males., Ser. 2, Pterid. l: 525. 1982. Basionym: Sphaerostephanos wauensis Holttum, Fl.
Thelypteris translucens (Holttum) Christenh., comb. Males., Ser. 2, Pterid. l: 505. 1982.
nov. Basionym: Plesioneuron translucens Holttum, Thelypteris wenzhouensis (K.H.Shing & C.F.Zhang)

39
Christenh., comb. nov. Basionym: Cyclosorus Christenh., comb. nov. Basionym: Niphidium
wenzhouensis K.H.Shing & C.F.Zhang, Fl. Reipubl. albopunctatissimum Lellinger, Amer. Fern J. 62: 109.
Popularis Sin. 4: 234. 1999. 1972.
Thelypteris windischii (Salino) Christenh., comb. nov. Campyloneurum americanum (Hook.) Christenh.,
Basionym: Goniopteris windischii Salino, Phytotaxa comb. nov. Basionym: Polypodium americanum
255: 255. 2016. Hook., Sp. Filic. 5: 54. 1864.
Thelypteris woitapensis (Holttum) Christenh., comb. Campyloneurum anocarpos (Kunze) Christenh., comb.
nov. Basionym: Sphaerostephanos woitapensis nov. Basionym: Polypodium anocarpos Kunze,
Holttum, Fl. Males., Ser. 2, Pterid. l: 483. 1982. Linnaea 9: 40. 1834.
Thelypteris womersleyi (Holttum) Christenh., comb. Campyloneurum carinatum (Lellinger) Christenh.,
nov. Basionym: Pronephrium womersleyi Holttum, comb. nov. Basionym: Niphidium carinatum
Blumea 20: 108. 1972. Lellinger, Amer. Fern J. 62: 113. 1972.
Thelypteris woodlarkensis (Copel.) Christenh., comb. Campyloneurum crassifolium (L.) Christenh., comb.
nov. Basionym: Cyathea woodlarkensis Copel., nov. Basionym: Polypodium crassifolium L., Sp. Pl. 2:
Philipp. J. Sci., C 9: 1. 1914. 1083. 1753. (Figure 5D)
Thelypteris wulingshanensis (C.M.Zhang) Christenh., Campyloneurum longifolium (Cav.) Christenh., comb.
comb. nov. Basionym: Cyclosorus wulingshanensis nov. Basionym: Polypodium longifolium Cav., Descr.
C.M.Zhang, Keys Vasc. Pl. Wuling Mts: 567. 1995. Pl.: 245. 1802.
Thelypteris wuyishanica (Ching) Christenh., comb. Campyloneurum macbridei (Lellinger) Christenh.,
nov. Basionym: Metathelypteris wuyishanica Ching, comb. nov. Basionym: Niphidium macbridei
Wuyi Sci. J. 1: 5. 1981. Lellinger, Amer. Fern J. 62: 117. 1972.
Thelypteris xingwenensis (Ching ex Y.X.Lin) Christenh., Campyloneurum mortonianum (Lellinger) Christenh.,
comb. nov. Basionym: Stegnogramma xingwenensis comb. nov. Basionym: Niphidium mortonianum
Ching ex Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 350. Lellinger, Amer. Fern J. 62: 115. 1972.
1999. Campyloneurum nidulare (Rosenst.) Christenh., comb.
Thelypteris xinpingensis (Ching ex Y.X.Lin) Christenh., & stat. nov. Basionym: Polypodium crassifolium L.
comb. nov. Basionym: Pseudocyclosorus xinpingensis var. nidulare Rosenst., Repert. Sp. Nov. Regni Veg.
Ching ex Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 326. 22: 18. 1925.
1999, as ‘xinpingesis’. Campyloneurum rufosquamatum (Lellinger)
Thelypteris yahanensis (Ching ex Y.X.Lin) Christenh., Christenh., comb. nov. Basionym: Niphidium
comb. nov. Basionym: Leptogramma yahanensis rufosquamatum Lellinger, Amer. Fern J. 62: 115.
Ching ex Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 324. 1972.
1999. Campyloneurum vittaria (Mett.) Christenh., comb.
Thelypteris yamawakii (H.Ito) Christenh., comb. nov. nov. Basionym: Polypodium vittaria Mett., Filic.
Basionym: Cyclosorus yamawakii H.Ito, J. Jap. Bot. Lechl. 1: 8. 1856.
27: 340. 1952. Draconopteris grandidentata (Ces) Christenh., comb.
Thelypteris yuanjiangensis (Ching) Christenh., comb. nov. Basionym: Polypodium dilatatum Liebm. var.
nov. Basionym: Cyclosorus yuanjiangensis Ching, Fl. grandidentatum Ces., Atti Accad. Sci. Fis. Nat. Napoli
Reipubl. Popularis Sin. 4: 203. 1999. 7: 27. 1876.
Thelypteris yunguiensis (Ching & Y.X.Lin) Christenh., Dryopteris robur Christenh., nom. nov. Replaced
comb. nov. Basionym: Pronephrium yunguiensis synonym: Aspidium triste Blume, Enum. Pl. Javae 2:
Ching & Y.X.Lin, Fl. Reipubl. Popularis Sin. 4: 352. 169. 1828, not: Dryopteris tristis (Kunze) Kuntze, nor:
1999. Dryopteris stenolepis (Baker) C.Chr. Etymology: From
Thelypteris zayuensis (Ching & S.K.Wu) Christenh., Latin, robur, strength.
comb. nov. Basionym: Pseudocyclosorus zayuensis Grammitis abebaion (Alderw.) Christenh., comb. nov.
Ching & S.K.Wu, Fl. Xizang. 1: 168. 1983. Basionym: Polypodium abebaion Alderw., Nova
Thelypteris zhangii (K.H.Shing) Christenh., comb. nov. Guinea 14: 45. 1924.
Basionym: Cyclosorus zhangii K.H.Shing, Fl. Reipubl. Grammitis acrosora (A.Rojas) Christenh., comb. nov.
Popularis Sin. 4: 231. 1999. Basionym: Cochlidium acrosorum A.Rojas, Mét. Ecol.
Sist. 2(2): 14. 2007.
F24. Polypodiaceae. References: Ranker et al. (2004), Grammitis aethiopica (Pic.Serm.) Christenh., comb.
Labiak & Moran (2018). nov. Basionym: Xiphopteris aethiopica Pic.Serm.,
Webbia 27: 450. 1972.
Campyloneurum albopunctatissimum (Lellinger)
Grammitis affinis (Labiak) Christenh., comb. nov.

Basionym: Lellingeria affinis Labiak, Fl. Neotrop. Hook.

Monogr. 111: 14. 2013, not Grammitis affinis Wall., Grammitis asiatica (Copel.) Christenh., comb. nov.
nom. inval. nom. nud.) Basionym: Calymmodon asiaticus Copel., Philipp. J.
Grammitis alansmithii (A.Rojas) Christenh., comb. Sci. 38: 154. 1929.
nov. Basionym: Melpomene alansmithii A.Rojas, Grammitis asthenophylla (L.E.Bishop ex A.R.Sm.)
Revista Biol. Trop. 49: 444. 2001. Christenh., comb. nov. Basionym: Ceradenia
Grammitis albicans (Lehnert) Christenh., comb. nov. asthenophylla L.E.Bishop ex A.R.Sm., Novon 3: 182.
Basionym: Melpomene albicans Lehnert, Amer. Fern 1993.
J. 98: 216. 2009. Grammitis aulaeifolia (L.E.Bishop ex A.R.Sm.)
Grammitis alboglandulosa (Bonap.) Christenh., comb. Christenh., comb. nov. Basionym: Ceradenia
nov. Basionym: Polypodium alboglandulosum aulaeifolia L.E.Bishop ex A.R.Sm., Novon 3: 182.
Bonap., Notes Ptérid. 10: 186. 1920. 1993.
Grammitis allocota (Alderw.) Christenh., comb. nov. Grammitis auroseiomena (L.E.Bishop) Christenh.,
Basionym: Polypodium allocotum Alderw., Nova comb. nov. Basionym: Ceradenia auroseiomena
Guinea 14: 47. 1924. L.E.Bishop, Amer. Fern J. 79: 18. 1989.
Grammitis alternidens (Ces.) Christenh., comb. nov. Grammitis axioche Christenh., nom. nov. Replaced
Basionym: Polypodium alternidens Ces., Atti Accad. synonym: Ascogrammitis tungurahuae Sundue,
Sci. Fis. Mat. Napoli 7: 25. 1876. Brittonia 62: 391. 2010, not: Grammitis tunguraguae
Grammitis amplisora (Labiak) Christenh., comb. nov. (Rosenst.) C.V.Morton. Etymology: Ἀξιόχη, a nymph
Basionym: Lellingeria amplisora Labiak, Fl. Neotrop. in Greek mythology.
Monogr. 111: 16. 2013. Grammitis ayopayana (M.Kessler & A.R.Sm.)
Grammitis anazalea (Sundue & Lehnert) Christenh., Christenh., comb. nov. Basionym: Ceradenia
comb. nov. Basionym: Melpomene anazalea Sundue ayopayana M.Kessler & A.R.Sm., Organisms
& Lehnert, Amer. Fern J. 98: 209. 2009. Diversity Evol. 8: 167. 2008.
Grammitis ancestralis (Copel.) Christenh., comb. Grammitis balslevii Christenh., nom. nov. Replaced
nov. Basionym: Prosaptia ancestralis Copel., Leafl. synonym: Lellingeria paramicola Labiak, Fl. Neotrop.
Philipp. Bot. 3: 385. 1910. Monogr. 111: 72. 2013, not: Grammitis paramicola
L.E.Bishop. Etymology: Named for Henrik Balslev,
Grammitis anjanaharibensis (Rakotondr.) Christenh.,
who collected the type specimen of this species in
comb. nov. Basionym: Zygophlebia anjanaharibensis
Ecuador.
Rakotondr., Taxon 55: 147. 2006.
Grammitis barathrophylla (Baker) Christenh., comb.
Grammitis antillensis (Proctor) Christenh., comb. nov.
nov. Basionym: Polypodium barathrophyllum Baker,
Basionym: Grammitis phlegmaria (J.Sm.) Proctor var.
J. Bot. 29: 107. 1891.
antillensis Proctor, Rhodora 68: 467. 1966.
Grammitis beleensis (Copel.) Christenh., comb. nov.
Grammitis antipodalis (Copel.) Christenh., comb. nov.
Basionym: Ctenopteris beleensis Copel., Univ. Calif.
Basionym: Xiphopteris antipodalis Copel., Univ. Calif.
Publ. Bot. 18: 225. 1942.
Publ. Bot. 18: 224. 1942.
Grammitis bipinnatifidum (Baker) Christenh., comb.
Grammitis aoide Christenh., nom. nov. Replaced
nov. Basionym: Polypodium bipinnatifidum Baker, J.
synonym: Lellingeria pumila Labiak, Brittonia 52:
Bot. 28: 109. 1890.
249. 2000, not: Grammitis pumila Sw., nor: G.
pumila (C.Chr.) Stolze. Etymology: After Ἀοιδή was Grammitis bivascularis (M.T.Murillo & A.R.Sm.)
was the Boeotian Muse of voice and song in Ancient Christenh., comb. nov. Basionym: Luisma
Greek mythology. bivascularis M.T.Murillo & A.R.Sm., Novon 13: 313.
2003.
Grammitis arche Christenh., nom. nov. Replaced
synonym: Ceradenia maxoniana L.E.Bishop, Amer. Grammitis blechnifrons (Hayata) Christenh., comb.
Fern J. 79: 24. 1979, not: Grammitis maxoniana nov. Basionym: Polypodium decrescens Christ var.
Lellinger. Etymology: After Ἀρχή, the Muse of Origins blechnifrons Hayata, Icon. Pl. Formosan. 4: 245.
in Ancient Greek mythology. 1914.
Grammitis arthrothrix (L.E.Bishop & A.R.Sm.) Grammitis boivinii (Mett. ex Kuhn) Christenh., comb.
Christenh., comb. nov. Basionym: Ceradenia nov. Basionym: Polypodium boivinii Mett. ex Kuhn,
arthrothrix L.E.Bishop & A.R.Sm., Ann. Missouri Bot. Filic. Afr.: 146. 1868.
Gard. 77: 255. 1990. Grammitis bolobensis (Brause) Christenh., comb.
Grammitis ascensionensis (Hieron.) Christenh., comb. nov. Basionym: Polypodium bolobense Brause, Bot.
nov. Basionym: Polypodium ascensionense Hieron., Jahrb. Syst. 49: 38. 1912.
Hedwigia 44. 93. 1905, not Grammitis ascensionis Grammitis brachyphlebia (Baker) Christenh., comb.

41
nov. Basionym: Polypodium brachyphlebium Baker, Amer. Fern J. 98: 219. 2009.
Ann. Bot. (Oxford) 8: 129. 1894. Grammitis carrascoensis (M.Kessler & A.R.Sm.)
Grammitis bradeana (Labiak) Christenh., comb. nov. Christenh., comb. nov. Basionym: Lellingeria
Basionym: Terpsichore bradeana Labiak, Brittonia carrascoensis M.Kessler & A.R.Sm., Organisms
52: 251. 2000. Diversity Evol. 8: 169. 2008.
Grammitis brasiliense (Rosenst.) Christenh., comb. & Grammitis celebica (Blume) Christenh., comb. nov.
stat. nov. Basionym: Polypodium tenuiculum Fée var. Basionym: Polypodium celebicum Blume, Enum. Pl.
brasiliense Rosenst., Hedwigia 56: 370. 1915. Javae 2: 127. 1828.
Grammitis brenesii (A.Rojas) Christenh., comb. nov. Grammitis ceylanica (Parris) Christenh., comb. nov.
Basionym: Lellingeria brenesii A.Rojas, Lankesteriana Basionym: Prosaptia ceylanica Parris, Fern Gaz. 16:
7: 553. 2007. 242. 2002.
Grammitis brevipes (C.V.Morton) Christenh., comb. Grammitis circumvallata (Rosenst.) Christenh.,
nov. Basionym: Grammitis sodiroi var. brevipes comb. nov. Basionym: Polypodium circumvallatum
C.V.Morton, Phytologia 22: 79. 1971. Rosenst., Repert. Spec. Nov. Regni Veg. 12: 178.
Grammitis brevipubens (Labiak) Christenh., comb. 1913.
nov. Basionym: Stenogrammitis brevipubens Labiak, Grammitis clathrata (Sundue & M.Kessler) Christenh.,
Amer. Fern J. 102: 161. 2012. comb. nov. Basionym: Terpsichore clathrata Sundue
Grammitis brevivenosa (Alderw.) Christenh., comb. & M.Kessler, Organisms Diversity Evol. 8: 163. 2008.
nov. Basionym: Polypodium brevivenosum Alderw., Grammitis clavigera (A.R.Sm. ex Sundue) Christenh.,
Bull. Jard. Bot. Buitenzorg Ser. 2, 28: 40. 1918. comb. nov. Basionym: Ascogrammitis clavigera
Grammitis brooksiae (Alderw.) Christenh., comb. nov. A.R.Sm. ex Sundue, Brittonia 62: 373. 2010.
Basionym: Polypodium brooksiae Alderw., Bull. Jard. Grammitis cleio Christenh., nom. nov. Replaced
Bot. Buitenzorg Ser. 2, 23: 19. 1916. synonym: Lellingeria rupestris Parris, Kew Bull.
Grammitis burbidgei (Baker) Christenh., comb. nov. 57: 429. 2002, not: Grammitis rupestris Parris.
Basionym: Polypodium burbidgei Baker, J. Bot. 17: Etymology: Κλειώ, the muse of history in Ancient
42. 1879. Greek mythology.
Grammitis callichore Christenh., nom. nov. Replaced Grammitis cocosensis (A.Rojas) Christenh., comb. nov.
synonym: Galactodenia pumila Sundue & Labiak, Basionym: Terpsichore cocosensis A.Rojas, Brenesia
Syst. Bot. 37: 345. 2012, not: Grammitis pumila Sw., 45-46: 47. 1997.
nor: G. pumila (C.Chr.) Stolze. Etymology: Καλλιχόρη, Grammitis colombiensis (Sundue) Christenh., comb.
the nymph that nursed Dionysos in Greek mythology. nov. Basionym: Ascogrammitis colombiensis
Grammitis calliope Christenh., nom. nov. Replaced Sundue, Brittonia 62: 374. 2010.
synonym: Ctenopteris bipinnata Copel., Univ. Calif. Grammitis comorensis (Baker) Christenh., comb. nov.
Publ. Bot. 18: 225. 1942, not: Grammitis bipinnata Basionym: Polypodium comorense Baker, J. Bot. 15:
Stolze. Etymology: Καλλιόπη, the muse of epic 72. 1877.
poetry and eloquence in Ancient Greek mythology. Grammitis comosa (L.E.Bishop) Christenh., comb. nov.
Grammitis callophylla (C.H.Wright) Christenh., comb. Basionym: Ceradenia comosa L.E.Bishop, Amer. Fern
nov. Basionym: Polypodium callophyllum C.H.Wright, J. 79: 16. 1989.
Bull. Misc. Inf. Kew 1909: 362. 1909. Grammitis congregatifolia (Alderw.) Christenh.,
Grammitis calolepis (Labiak) Christenh., comb. nov. comb. nov. Basionym: Polypodium congregatifolium
Basionym: Lellingeria calolepis Labiak, Fl. Neotrop. Alderw., Nova Guinea 14: 47. 1924.
Monogr. 111: 32. 2013. Grammitis connellii (Baker) Christenh., comb. nov.
Grammitis canescens (A.Rojas) Christenh., comb. nov. Basionym: Polypodium connellii Baker, Trans. Linn.
Basionym: Terpsichore canescens A.Rojas, Mét. Ecol. Soc. London, Bot. ser. 2, 6: 82. 1901.
Sist. 3: 14. 2008. Grammitis contacta (Copel.) Christenh., comb. nov.
Grammitis capillata (Brause) Christenh., comb. nov. Basionym: Ctenopteris contacta Copel., Philipp. J.
Basionym: Polypodium capillatum Brause, Bot. Sci. 84: 447. 1956.
Jahrb. Syst. 49: 39. 1912. Grammitis contigua (C.Presl) Christenh., comb. nov.
Grammitis capillipes (C.Chr.) Christenh., comb. nov. Basionym: Prosaptia contigua C.Presl, Tent. Pterid.:
Basionym: Polypodium capillipes C.Chr., Brittonia 2: 166. 1836.
310. 1937. Grammitis costaricensis (Rosenst.) Christenh., comb.
Grammitis caput-gorgonis (Lehnert) Christenh., comb. nov. Basionym: Polypodium lherminieri Fée var.
nov. Basionym: Melpomene caput-gorgonis Lehnert, costaricense Rosenst., Repert. Spec. Nov. Regni Veg.

22: 17. 1925. Basionym: Polypodium duriusculum Christ, Verh.

Grammitis cryptocarpa (Copel.) Christenh., comb. nov. Nat. Ges. Basel 11: 213. 1895.
Basionym: Prosaptia cryptocarpa Copel., Philipp. J. Grammitis ecostata (Sodiro) Christenh., comb. nov.
Sci. 1(Suppl. 2): 158. 1906. Basionym: Polypodium ecostatum Sodiro, Crypt.
Grammitis ctenoidea (Brause) Christenh., comb. nov. Vasc. Quit. :308. 1893.
Basionym: Polypodium ctenoideum Brause, Bot. Grammitis elastica (Bory ex Willd.) Christenh., comb.
Jahrb. Syst. 56: 188. 1920. nov. Basionym: Polypodium elasticum Bory ex Willd.,
Grammitis curtisii (Baker) Christenh., comb. nov. Sp. Pl., ed. 4, 5: 183. 1810.
Basionym: Polypodium curtisii Baker, J. Bot. 19: 367. Grammitis emersonii (C.Presl) Christenh., comb. nov.
1881. Basionym: Prosaptia emersonii C.Presl, Tent. Pterid.:
Grammitis davalliacea (Copel.) Christenh., comb. nov. 166. 1836.
Basionym: Prosaptia davalliacea Copel., Univ. Calif. Grammitis engleriana (Brause) Christenh., comb. nov.
Publ. Bot. 12: 404. 1931. Basionym: Davallia engleriana Brause, Bot. Jahrb.
Grammitis decrescens (Christ) Christenh., comb. nov. Syst. 49: 27.1912.
Basionym: Polypodium decrescens Christ, Ann. Jard. Grammitis enterosoroides (Christ) Christenh., comb.
Bot. Buitenzorg II, 4: 35. 1904. nov. Basionym: Polypodium enterosoroides Christ,
Grammitis deltodon (Baker) Christenh., comb. nov. Bull. Herb. Boissier ser. 2, 7: 260. 1907.
Basionym: Polypodium deltodon Baker, J. Linn. Soc., Grammitis epaleata (Parris) Christenh., comb. nov.
Bot. 15: 419. 1876. Basionym: Ctenopteris epaleata Parris, Fern Gaz. 16:
Grammitis deltoideophylla (Baker) Christenh., comb. 239. 2001.
nov. Basionym: Polypodium deltoideophyllum Baker, Grammitis erato Christenh., nom. nov. Replaced
J. Bot. 14: 345. 1876. synonym: Polypodium conforme Brack., U.S. Expl.
Grammitis depressa (C.Chr.) Christenh., comb. nov. Exped., Filic. 16: 4. 1854, not: Grammitis conformis
Basionym: Polypodium depressum C.Chr., Index Filic.: J.Sm. Etymology: Ἐρατώ, the muse of love poetry in
522. 1906. Ancient Greek mythology.
Grammitis dercetis Christenh., nom. nov. Replaced Grammitis esmeraldensis Christenh., nom. nov.
synonym: Lellingeria bishopii Labiak, Fl. Neotrop. Replaced synonym: Lellingeria smithii Labiak, Fl.
Monogr. 111: 24. 2013, not: Grammitis bishopii Neotrop. Monogr. 111: 90. 2013, not: Grammitis
Stolze. Etymology: ∆ερκετις, a nymph in Greek smithii (A.Rojas) Christenh. Etymology: Named
mythology. for Esmeraldas in Ecuador, where the type of this
species was collected.
Grammitis devolii (S.J.Moore, Parris & W.L.Chiou)
Christenh., comb. nov. Basionym: Xiphopterella Grammitis esquiveliana (A.Rojas) Christenh., comb.
devolii S.J.Moore, Parris & W.L.Chiou, Bot. Stud. nov. Basionym: Terpsichore esquiveliana A.Rojas,
(Taipei) 54: 3. 2013. Revista Biol. Trop. 49: 448. 2001.
Grammitis devoluta (Baker) Christenh., comb. nov. Grammitis eunoe Christenh., nom. nov. Replaced
Basionym: Polypodium devolutum Baker, J. Linn. synonym: Ceradenia clavipila L.E.Bishop ex M.Kessler
Soc., Bot. 15: 419. 1876. & A.R.Sm., Organisms Diversity Evol. 8: 167. 2008,
not: Grammitis clavipila Parris. Etymology: Εὐνοε, a
Grammitis diaphana (Moguel & M.Kessler) Christenh.,
nymph and daughter of the river god Sangarius in
comb. nov. Basionym: Terpsichore diaphana Moguel
Greek Mythology.
& M.Kessler, Phytotaxa 2: 38. 2009.
Grammitis euterpe Christenh., nom. nov. Replaced
Grammitis dilatata (Sundue & M.Kessler) Christenh.,
synonym: Polypodium villosissimum var. majus
comb. nov. Basionym: Terpsichore dilatata Sundue
Reimers, not: Grammitis major (Copel.) C.V.Morton.
& M.Kessler, Organisms Diversity Evol. 8: 163. 2008.
Etymology: Eὐτέρπη, the muse of music and lyric
Grammitis doniana (Spreng.) Christenh., comb. nov. poetry in Ancient Greek mythology.
Basionym: Polypodium donianum Spreng., Syst. Veg.
Grammitis eximia (Copel.) Christenh., comb. nov.
(ed. 16) 4: 54. 1827.
Basionym: Ctenopteris eximia Copel., Philipp. J. Sci.
Grammitis dryas Christenh., nom. nov. Replaced 81: 112. 1953.
synonym: Ctenopteris brassii Copel., Univ. Calif. Publ.
Grammitis fawcettii (Jenman) Christenh., comb. nov.
Bot. 18: 225. 1942, not: Grammitis brassii Copel.
Basionym: Enterosora fawcettii Jenman, Gard.
Etymology: Δρύας, prince of Thrace, who was killed
Chron. ser. 3, 18: 62. 1894.
by his father King Lycourgos (c 820 BC), who mistook
him for a grapevine, a divine plant to Dionysos, a cult Grammitis fayae Christenh., nom. nov. Replaced
that King Lycurgus attempted to eradicate. synonym: Lellingeria pendulina A.R.Sm. & R.C.Moran,
Amer. Fern J. 81: 81. 1991, not: Grammitis pendulina
Grammitis duriuscula (Christ) Christenh., comb. nov.

43
(Sw.) Proctor. Etymology: Named for Alice Fay, nov. Basionym: Zygophlebia goodmanii Rakotondr.,
collector of the type specimen. Taxon 55: 145. 2006.
Grammitis flagellata (Lehnert) Christenh., comb. nov. Grammitis gordonii (S.B.Andrews) Christenh., comb.
Basionym: Melpomene flagellata Lehnert, Amer. nov. Basionym: Ctenopteris gordonii S.B.Andrews,
Fern J. 98: 222. 2009. Austrobaileya 1: 12. 1977.
Grammitis flagelliformis (Brause) Christenh., comb. Grammitis govidjoaensis (Brause) Christenh., comb.
nov. Basionym: Polypodium flagelliforme Brause, nov. Basionym: Polypodium govidjoaense Brause,
Bot. Jahrb. Syst. 56: 187. 1920. Bot. Jahrb. Syst. 49: 41. 1912.
Grammitis flagellipinnata (M.Kessler & A.R.Sm.) Grammitis gracillima (Copel.) Christenh., comb.
Christenh., comb. nov. Basionym: Lellingeria nov. Basionym: Polypodium gracillimum Copel. in
flagellipinnata M.Kessler & A.R.Sm., Organisms Perkins, Fragm. Fl. Philipp.: 189. 1905.
Diversity Evol. 8: 169. 2008. Grammitis grammitidiphylla (Copel.) Christenh., comb.
Grammitis flavovirens (Alston) Christenh., comb. nov. nov. Basionym: Calymmodon grammitidiphyllus
Basionym: Polypodium flavovirens Alston, J. Bot. 78: Copel., Univ. Calif. Publ. Bot. 18: 224. 1942.
228. 1940. Grammitis grata (Fée) Christenh., comb. nov.
Grammitis flexibilis (Labiak) Christenh., comb. nov. Basionym: Polypodium gratum Fée, Crypt. Vasc.
Basionym: Lellingeria flexibilis Labiak, Fl. Neotrop. Brésil 1: 242. 1869.
Monogr. 111: 43. 2013. Grammitis guanacastensis (A.Rojas) Christenh., comb.
Grammitis foersteri (Rosenst.) Christenh., comb. nov. nov. Basionym: Lellingeria guanacastensis A.Rojas,
Basionym: Polypodium bipinnatifidum Baker var. Brenesia 45-46: 37. 1996.
foersteri Rosenst., Repert. Spec. Nov. Regni Veg. Grammitis hainanensis (Parris) Christenh., comb. nov.
Beih. 12: 179. 1913. Basionym: Oreogrammitis hainanensis Parris, Fl.
Grammitis forsythiana (Baker) Christenh., comb. nov. China 2-3: 841. 2013.
Basionym: Polypodium forsythianum Baker, Bull. Grammitis halconensis (Copel.) Christenh., comb. nov.
Misc. Inform. Kew 1897: 300. 1897. Basionym: Polypodium halconense Copel., Philipp. J.
Grammitis fusca (Copel.) Christenh., comb. nov. Sci., C 2: 138. 1907.
Basionym: Ctenopteris fusca Copel., Univ. Calif. Publ. Grammitis hecistophylla (Copel.) Christenh., comb.
Bot. 18: 225. 1942. nov. Basionym: Polypodium hecistophyllum Copel.,
Grammitis fuscata (Blume) Christenh., comb. nov. Philipp. J. Sci. 56: 477. 1935.
Basionym: Polypodium fuscatum Blume, Enum. Pl. Grammitis helike Christenh., nom. nov. Replaced
Javae 2: 129. 1828. synonym: Prosaptia alata Christ., Ann. Jard. Bot.
Grammitis fuscopilosa (F.Muell. & Baker) Christenh., Buitenzorg II 5: 127. 1905, not Grammitis alata
comb. nov. Basionym: Polypodium fuscopilosum (Blume) C.V.Morton. Etymology: Ἑλίκη, a nymph that
F.Muell. & Baker, J. Bot. 25: 169. 1887. nurtured the god Zeus in infancy on Crete.
Grammitis gabonensis (Parris) Christenh., comb. nov. Grammitis hieronymusii (C.Chr.) Christenh., comb. nov.
Basionym: Ctenopterella gabonensis Parris, Fern Basionym: Polypodium hieronymusii C.Chr., Index
Gaz. 19: 89. 2013. Filic.: 533. 1906.
Grammitis geluensis (Rosenst.) Christenh., comb. nov. Grammitis hildebrandtii (Hieron.) Christenh., comb.
Basionym: Polypodium geluense Rosenst., Repert. nov. Basionym: Polypodium hildebrandtii Hieron.,
Spec. Nov. Regni Veg. 5: 374. 1908. Hedwigia 44: 91. 1905.
Grammitis glabra (M.Kessler & A.R.Sm. ) Christenh., Grammitis hirsuta (A.R.Sm. & R.C.Moran) Christenh.,
comb. nov. Basionym: Ceradenia glabra M.Kessler & comb. nov. Basionym: Lellingeria hirsuta A.R.Sm. &
A.R.Sm., Organisms Diversity Evol. 8: 167. 2008. R.C.Moran, Amer. Fern J. 81: 80. 1991.
Grammitis glandulifera (A.Rojas) Christenh., comb. Grammitis hornei (Baker) Christenh., comb. nov.
nov. Basionym: Terpsichore glandulifera A.Rojas, Basionym: Polypodium hornei Baker, J. Bot. 17: 298.
Lankesteriana 6: 96. 2006. 1879.
Grammitis glandulosa (J.Sm.) Christenh., comb. nov. Grammitis huancabambensis (Lehnert) Christenh.,
Basionym: Ctenopteris glandulosa J.Sm., Hist. Filic. comb. nov. Basionym: Melpomene huancabambensis
185. 1875. Lehnert, Amer. Fern J. 98: 224. 2009.
Grammitis glaziovii (Baker) Christenh., comb. nov. Grammitis humbertii (C.Chr.) Christenh., comb. nov.
Basionym: Polypodium glaziovii Baker, Ann. Bot. Basionym: Polypodium humbertii C.Chr., Arch. Bot.,
(Oxford) 5. 465. 1891. Bull. Mens. 2: 215. 1928.
Grammitis goodmanii (Rakotondr.) Christenh., comb. Grammitis hymenophylloides (Parris) Christenh.,

comb. nov. Basionym: Ctenopteris hymenophylloides Basionym: Polypodium kjellbergii C.Chr., Bot. Jahrb.
Parris, Kew Bull. 59: 221. 2004. Syst. 66: 63. 1933.
Grammitis immersa (Brownlie) Christenh., comb. nov. Grammitis kookenamae (Jenman) Christenh., comb.
Basionym: Ctenopteris immersa Brownlie, Nova nov. Basionym: Polypodium kookenamae Jenman,
Hedwigia, Beih. 55: 352. 1977. Timehri 5: 215. 1886.
Grammitis inconspicua (Blume) Christenh., comb. nov. Grammitis kunstleri (Parris) Christenh., comb. nov.
Basionym: Polypodium inconspicuum Blume, Enum. Basionym: Oreogrammitis kunstleri Parris, Gard.
Pl. Javae 2: 130. 1828. Bull. Singapore 58: 254. 2007.
Grammitis integripaleata (Copel.) Christenh., comb. Grammitis labiakii Christenh., nom. nov. Replaced
nov. Basionym: Ctenopteris integripaleata Copel., synonym: Lellingeria jimenezii Labiak, Fl. Neotrop.
Univ. Calif. Publ. Bot. 18: 225. 1942. Monogr. 111: 53. 2013, not: Grammitis jimenezii
Grammitis intermedia (Ching) Christenh., comb. nov. (M.Kessler & A.R.Sm.) Christenh. Etymology: Named
Basionym: Polypodium urceolare var. intermedium for Paulo Labiak, who first described this species.
Ching, Bull. Dept. Biol. Sun Yatsen Univ. 6: 32. 1933. Grammitis laxifolia (A.R.Sm. & R.C.Moran) Christenh.,
Grammitis intonsa (L.E.Bishop ex León-Parra & comb. nov. Basionym: Lellingeria laxifolia A.R.Sm. &
Mostacero) Christenh., comb. nov. Basionym: R.C.Moran, Amer. Fern J. 81: 81. 1991.
Ceradenia intonsa L.E.Bishop ex León-Parra & Grammitis lehnertii Christenh., nom. nov. Replaced
Mostacero, Fl. Colombia 29: 38. 2012. synonym: Melpomene jimenezii Lehnert, Amer. Fern
Grammitis itatiaiensis (Labiak & Condack) Christenh., J. 98: 227. 2009, not: Grammitis jimenezii (M.Kessler
comb. nov. Basionym: Ceradenia itatiaiensis Labiak & A.R.Sm.) Christenh. Etymology: Named for Marcus
& Condack, Novon 18: 495. Lehnert, who first described this species.
Grammitis itatimensis (C.Chr.) Christenh., comb. nov. Grammitis leonae Christenh., nom. nov. Replaced
Basionym: Polypodium itatimense C.Chr., Index Filic., synonym: Terpsichore youngii B.León & A.R.Sm.,
Suppl. 3: 151. 1934. Amer. Fern J. 93: 84. 2003, not: Grammitis youngii
Stolze. Etymology: Named for Blanca León, who first
Grammitis javanica (Copel. ) Christenh., comb. nov.
described this species.
Basionym: Polypodium javanicum Copel., Philipp. J.
Sci., C 8: 142. 1913. Grammitis lepida (Brause) Christenh., comb. nov.
Basionym: Polypodium lepidum Brause, Notizbl. Bot.
Grammitis jimenezii (M.Kessler & A.R.Sm.) Christenh.,
Gart. Berlin-Dahlem 8: 139. 1922.
comb. nov. Basionym: Ceradenia jimenezii M.Kessler
& A.R.Sm., Organisms Diversity Evol. 8: 168. 2008. Grammitis leucolepis (Gilbert) Christenh., comb. nov.
Basionym: Polypodium leucolepis Gilbert, Bull.
Grammitis kalawayae (M.Kessler & A.R.Sm.)
Torrey Bot. Club 24: 260. 1897.
Christenh., comb. nov. Basionym: Ceradenia
kalawayae M.Kessler & A.R.Sm., Organisms Diversity Grammitis leucosora (Bojer ex Hook.) Christenh.,
Evol. 8: 168. 2008. comb. nov. Basionym: Polypodium leucosorum Bojer
ex Hook., Icon. Pl. 10: sub. t. 942–943. 1854.
Grammitis kanashiroi (Nakai) Christenh., comb. nov.
Basionym: Prosaptia kanashiroi Nakai in Yamam., Grammitis leysii (Baker) Christenh., comb. nov.
Suppl. Ic. Pl. Formos. V: 6. 1932. Basionym: Polypodium leysii Baker, J. Bot. 17: 66.
1879.
Grammitis kessleri Christenh., nom. nov. Replaced
synonym: Terpsichore nana Sundue & M.Kessler, Grammitis livida (Mett.) Christenh., comb. nov.
Organisms Diversity Evol. 8: 164. 2008, not: Basionym: Polypodium lividum Mett., Ann. Mus. Bot.
Grammitis nana Fée, nor: Grammitis nana Brack. Lugd.-Bat. 2. 221. 1866.
Etymology: Named for Michael Kessler, who first Grammitis lobulata (A.Rojas) Christenh., comb. nov.
described this species. Basionym: Terpsichore lobulata A.Rojas, Mét. Ecol.
Grammitis khaoluangensis (Tagawa & K.Iwats.) Sist. 3: 14. 2008.
Christenh., comb. nov. Basionym: Xiphopteris Grammitis longeattenuata (D.M.Forero & J.Murillo)
khaoluangensis Tagawa & K.Iwats., Acta Phytotax. Christenh., comb. nov. Basionym: Lellingeria
Geobot. 23: 177. 1969. longeattenuata D.M.Forero & J.Murillo, Caldasia 32:
Grammitis khasyana (Hook.) Christenh., comb. nov. 321. 2010.
Basionym: Polypodium khasyanum Hook., Icon. Pl. Grammitis longicaulis (Sundue & M.Kessler)
10: t. 949. 1854. Christenh., comb. nov. Basionym: Terpsichore
Grammitis kirkii (Parris) Christenh., comb. nov. longicaulis Sundue & M.Kessler, Organisms Diversity
Basionym: Terpsichore kirkii Parris, Kew Bull. 57: Evol. 8: 163. 2008.
431. 2002. Grammitis longiceps (Rosenst.) Christenh., comb. nov.
Grammitis kjellbergii (C.Chr.) Christenh., comb. nov. Basionym: Polypodium longiceps Rosenst., Repert.

45
Spec. Nov. Regni Veg. 12: 177. 1913. Ancient Greek mythology.
Grammitis longipilosa (C.Chr.) Christenh., comb. nov. Grammitis michaelium (Lehnert) Christenh., comb.
Basionym: Polypodium longipilosum C.Chr., Bot. nov. Basionym: Melpomene michaelium Lehnert,
Tidsskr. 25: 78. 1903. Amer. Fern J. 98: 229. 2009, as ‘michaelis’. Note: This
Grammitis longisetosa (Hook.) Christenh., comb. nov. is named for two people called Michael, and hence it
Basionym: Polypodium longisetosum Hook., Sp. Filic. needs to be in the 3rd declension plural.
4: 225. 1864. Grammitis microcystis (L.E.Bishop & A.R.Sm.)
Grammitis loxensis (Sundue) Christenh., comb. nov. Christenh., comb. nov. Basionym: Ceradenia
Basionym: Ascogrammitis loxensis Sundue, Brittonia microcystis L.E.Bishop & A.R.Sm., Ann. Missouri Bot.
62: 380. 2010. Gard. 77: 255. 1990.
Grammitis luetzelburgii (Rosenst.) Christenh., comb. Grammitis microlepis (Rosenst. ) Christenh., comb.
nov. Basionym: Polypodium luetzelburgii Rosenst., nov. Basionym: Polypodium blepharodes Maxon var.
Repert. Spec. Nov. Regni Veg. 20: 93. 1924. microlepis Rosenst., Repert. Spec. Nov. Regni Veg.
22: 14. 1925.
Grammitis macra (Copel.) Christenh., comb. nov.
Basionym: Polypodium macrum Copel. in Perkins, Grammitis micropecten (C.Chr.) Christenh., comb. nov.
Fragm. Fl. Philipp.: 189. 1905. Basionym: Polypodium oosorum var. micropecten
C.Chr., Dansk Bot. Ark. 7: 153. 1932.
Grammitis macrorhyncha (Baker) Christenh., comb.
nov. Basionym: Polypodium macrorhynchum Baker, Grammitis micropora (Copel. ) Christenh., comb. nov.
J. Bot. 18: 370. 1880. Basionym: Ctenopteris micropora Copel., Philipp. J.
Sci. 81: 44. 1952.
Grammitis madagascariensis (Rakotondr. & Rouhan)
Christenh., comb. nov. Basionym: Leucotrichum Grammitis militaris (Maxon) Christenh., comb. nov.
madagascariense Rakotondr. & Rouhan, Syst. Bot. Basionym: Polypodium militare Maxon, Contr. Gray
37: 335. 2012. Herb. 165: 71. 1947.
Grammitis madidiensis (M.Kessler & A.R.Sm.) Grammitis minuta (Blume) Christenh., comb. nov.
Christenh., comb. nov. Basionym: Ceradenia Basionym: Polypodium minutum Blume, Enum. Pl.
madidiensis M.Kessler & A.R.Sm., Organisms Javae 2: 130. 1828.
Diversity Evol. 8: 168. 2008. Grammitis minutissima (J.W.Moore) Christenh.,
Grammitis maidenii (Watts) Christenh., comb. nov. comb. nov. Basionym: Polypodium minutissimum
Basionym: Polypodium maidenii Watts, Proc. Linn. J.W.Moore, Bull. Bernice P. Bishop Mus. 102: 11.
Soc. New South Wales 39: 793. 1915. 1933.
Grammitis malaccana (Baker) Christenh., comb. nov. Grammitis mneme Christenh., nom. nov. Replaced
Basionym: Polypodium malaccanum Baker, Ann. Bot. synonym: Polypodium subspathulatum Brack.,
(Oxford) 8: 129. 1894. U.S. Expl. Exped., Filic. 16: 3. 1854, not: Grammitis
subspathulata Farw. Etymology: Μνήμη was
Grammitis malayensis (Parris) Christenh., comb. nov.
the Boeotian Muse of Memory in Ancient Greek
Basionym: Oreogrammitis malayensis Parris, Gard.
mythology.
Bull. Singapore 58: 254. 2007.
Grammitis mnemosyne Christenh., nom. nov.
Grammitis marginalis (León-Parra) Christenh., comb.
Replaced synonym: Ceradenia setosa M.Kessler
nov. Basionym: Ceradenia marginalis León-Parra, Fl.
& A.R.Sm., Organisms Diversity Evol. 8: 168.
Colombia 29: 53. 2012.
2008, not: Grammitis setosa (Kaulf.) C.Presl.
Grammitis marialta (L.E.Bishop ex León-Parra) Etymology: Μνημοσύνη, mother of the Muses and
Christenh., comb. nov. Basionym: Ceradenia the personification of memory in Ancient Greek
marialta L.E.Bishop ex León-Parra, Fl. Colombia 29: mythology.
56. 2012.
Grammitis mollicoma (Nees & Blume) Christenh.,
Grammitis melete Christenh., nom. nov. Replaced comb. nov. Basionym: Polypodium mollicomum
synonym: Cochlidium repandum L.E.Bishop, Amer. Nees & Blume, Nova Acta Phys.-Med. Acad. Caes.
Fern J. 68: 83. 1978, not: Grammitis repanda Kunze Leop.-Carol. Nat. Cur. 11: 121. 1823.
ex Mett. Etymology: Μελέτη was the Boeotian
Grammitis monosora (Moguel & M.Kessler) Christenh.,
Muse of thought and meditation in Ancient Greek
comb. nov. Basionym: Terpsichore monosora Moguel
mythology.
& M.Kessler, Phytotaxa 2: 40. 2009.
Grammitis melpomene Christenh., nom. nov. Replaced
Grammitis moorei (Parris & Ralf Knapp) Christenh.,
synonym: Melpomene vulcanica Lehnert, Amer. Fern
comb. nov. Basionym: Radiogrammitis moorei Parris
J. 98: 245. 2009, not: Grammitis vulcanica Blume.
& Ralf Knapp, Fl. China 2-3: 842. 2013.
Etymology: Μελπομένη, the muse of tragedy in
Grammitis moranii (Labiak) Christenh., comb. nov.

Basionym: Lellingeria moranii Labiak, Fl. Neotrop. Brade & Rosenst., Arq. Inst. Biol. Veg. 2: 3. 1935.
Monogr. 111: 68. 2013. Grammitis pectinata (T.Moore) Christenh., comb. nov.
Grammitis multicaudata (Copel.) Christenh., comb. Basionym: Prosaptia pectinata T.Moore, Index Filic.:
nov. Basionym: Polypodium multicaudatum Copel., 298. 1861.
Philipp. J. Sci. 1(Suppl. 2): 160. 1906. Grammitis pediculata (Baker) Christenh., comb. nov.
Grammitis murudensis (Copel.) Christenh., comb. nov. Basionym: Polypodium pediculatum Baker, Syn. Filic.
Basionym: Polypodium murudense Copel., Philipp. J. (Hooker & Baker): 455. 1868.
Sci., C 12: 61. 1917. Grammitis pendens (Rosenst.) Christenh., comb. nov.
Grammitis muscoides (Copel.) Christenh., comb. nov. Basionym: Polypodium pendens Rosenst., Repert.
Basionym: Polypodium muscoides Copel., Leafl. Spec. Nov. Regni Veg. 12: 177. 1913.
Philipp. Bot. 3: 839. 1910. Grammitis pensilis (Ridl.) Christenh., comb. nov.
Grammitis musgraviana (Baker) Christenh., comb. Basionym: Polypodium pensile Ridl., Trans. Linn. Soc.
nov. Basionym: Polypodium musgravianum Baker, J. London, Bot. 9: 262. 1916.
Bot. 28: 108. 1890. Grammitis perplexa (Parris) Christenh., comb. nov.
Grammitis narinensis (León-Parra) Christenh., comb. Basionym: Ctenopteris perplexa Parris, Fern Gaz. 16:
nov. Basionym: Ceradenia narinensis León-Parra, Fl. 201. 2001.
Colombia 29: 65. 2012. Grammitis personata (Lehnert) Christenh., comb. nov.
Grammitis negrosensis (Copel.) Christenh., comb. nov. Basionym: Melpomene personata Lehnert, Amer.
Basionym: Polypodium negrosense Copel., Leafl. Fern J. 98: 237. 2009.
Philipp. Bot. 2: 409. 1908. Grammitis phloiocharis (L.E.Bishop) Christenh., comb.
Grammitis nervata (A.Rojas) Christenh., comb. nov. nov. Basionym: Ceradenia phloiocharis L.E.Bishop,
Basionym: Cochlidium nervatum A.Rojas, Mét. Ecol. Amer. Fern J. 79: 19. 1989.
Sist. 2: 14. 2007. Grammitis pilipalaea (M.Kessler & A.R.Sm.)
Grammitis nhatrangensis (C.Chr. & Tardieu) Christenh., Christenh., comb. nov. Basionym: Ceradenia
comb. nov. Basionym: Ctenopteris nhatrangensis pilipalaea M.Kessler & A.R.Sm., Organisms Diversity
C.Chr. & Tardieu, Notul. Syst. (Paris) 8: 181. 1939. Evol. 8: 168. 2008.
Grammitis nutans (Blume) Christenh., comb. nov. Grammitis pilipecten (L.E.Bishop ex M.Kessler
Basionym: Polypodium nutans Blume, Enum. Pl. & A.R.Sm.) Christenh., comb. nov. Basionym:
Javae 2: 128. 1828. Ceradenia pilipecten L.E.Bishop ex M.Kessler &
Grammitis nymphe Christenh., nom. nov. Replaced A.R.Sm., Organisms Diversity Evol. 8: 168. 2008.
synonym: Xiphopterella gracilis Parris, Gard. Bull. Grammitis pinnatifida (C.Presl) Christenh., comb.
Singapore 58: 250. 2007, not: Grammitis gracilis nov. Basionym: Prosaptia pinnatifida C.Presl, Tent.
(Hook.) Stolze. Etymology: νύμφη, a minor nature Pterid.: 166. 1836.
deity in Greek mythology, usually associated with a Grammitis pirrensis (A.R.Sm.) Christenh., comb. nov.
land formation or natural phenomenon. Basionym: Terpsichore pirrensis A.R.Sm., Novon 3:
Grammitis occidentalis (Lehnert) Christenh., comb. 483. 1993.
nov. Basionym: Melpomene occidentalis Lehnert, Grammitis polita (Parris) Christenh., comb. nov.
Amer. Fern J. 98: 231. 2009. Basionym: Oreogrammitis polita Parris, Gard. Bull.
Grammitis oxapampensis (Sundue) Christenh., comb. Singapore 58: 266. 2007.
nov. Basionym: Ascogrammitis oxapampensis Grammitis polyhymnia Christenh., nom. nov. Replaced
Sundue, Brittonia 62: 382. 2010. synonym: Plectopteris gracilis Fée, Mém. Foug. 5:
Grammitis paradoxa (Lehnert) Christenh., comb. nov. 230. 1852, not: Grammitis gracilis (Hook.) Stolze.
Basionym: Melpomene paradoxa Lehnert, Amer. Etymology: Πολυύμνια, the muse of sacred poetry
Fern J. 98: 235. 2009. and hymns in Ancient Greek mythology.
Grammitis parrisiae (Sundue & Labiak) Christenh., Grammitis polymorpha (Copel.) Christenh., comb. nov.
comb. nov. Basionym: Galactodenia parrisiae Basionym: Prosaptia polymorpha Copel., Philipp. J.
Sundue & Labiak, Syst. Bot. 37: 345. 2012. Sci., C 2: 136. 1907.
Grammitis paucipinnata (Parris) Christenh., comb. Grammitis polytricha (Copel.) Christenh., comb. nov.
nov. Basionym: Lellingeria paucipinnata Parris, Kew Basionym: Ctenopteris polytricha Copel., Univ. Calif.
Bull. 57: 428. 2002. Publ. Bot. 18: 225. 1942.
Grammitis paulistana (Brade & Rosenst.) Christenh., Grammitis pradoana Christenh., nom. nov. Replaced
comb. nov. Basionym: Polypodium paulistanum synonym: Lellingeria longifolia Labiak, Fl. Neotrop.
Monogr. 111: 59. 2013, not Grammitis longifolia
Blume. Etymology: Named for Jefferson Prado, for

47
his contributions to Brazilian pteridology. R.Y.Hirai & J.Prado, Taxon 60: 1132. 2011.
Grammitis praeceps (Sundue & M.Kessler) Christenh., Grammitis sabrinae Christenh., nom. nov. Replaced
comb. nov. Basionym: Terpsichore praeceps Sundue synonym: Enterosora bishopii A.Rojas, Lankesteriana
& M.Kessler, Organisms Diversity Evol. 8: 164. 2008. 6: 9. 2006, not: Grammitis bishopii Stolze. Etymology:
Grammitis preslii (Fée) Christenh., comb. nov. Named for Hafren (Sabrina in Latin), a legendary
Basionym: Prosaptia preslii Fée, Mém. Foug., 5. Gen. British princess (daughter of King Locrin) who was
Filic.: 324. 1850. drowned in the River Severn.
Grammitis pseudospiralis (Alderw.) Christenh., comb. Grammitis sarawakensis (Parris) Christenh., comb.
nov. Basionym: Polypodium pseudospirale Alderw., nov. Basionym: Ctenopteris sarawakensis Parris in
Bull. Jard. Bot. Buitenzorg, Ser. 2, 1: 29. 1911. Hooker's Icon. Pl. 40: t. 3986. 1990.
Grammitis pseudotenuicula (Labiak) Christenh., comb. Grammitis satyros Christenh., nom. nov. Replaced
nov. Basionym: Lellingeria pseudotenuicula Labiak, synonym: Prosaptia bipinnata C.Presl, Tent. Pterid.:
Fl. Neotrop. Monogr. 111: 84. 2013. 166. 1836, not: Grammitis bipinnata Stolze.
Etymology: Σάτυρος, a member of a troup of male
Grammitis pseudotrichomanoides (Hayata)
companions of Dionysos, god of wine and harvest,
Christenh., comb. nov. Basionym: Polypodium
who spend their time drinking and chasing nymphs
pseudotrichomanoides Hayata, Icon. Pl. Formosan.
in Ancient Greek mythology.
4: 251. 1914.
Grammitis schlechteri (Christ) Christenh., comb. nov.
Grammitis pubipes (Copel.) Christenh., comb. nov.
Basionym: Prosaptia schlechteri Christ, Nachtr. Fl.
Basionym: Prosaptia pubipes Copel., Bull. Bernice P.
Schutzgeb. Südsee [Schumann & Lauterbach]: 41.
Bishop Mus. 93: 13. 1932.
1905.
Grammitis pullei (Alston) Christenh., comb. nov.
Grammitis sclerophylla (Alderw.) Christenh., comb.
Basionym: Polypodium pullei Alston, J. Bot. 78: 229.
nov. Basionym: Polypodium sclerophyllum Alderw.,
1940.
Nova Guinea 14: 45. 1924.
Grammitis pulogensis (Copel.) Christenh., comb. nov.
Grammitis secunda (Ridl.) Christenh., comb. nov.
Basionym: Polypodium pulogense Copel., Philipp. J.
Basionym: Polypodium secundum Ridl., Trans. Linn.
Sci., C 6: 148. 1911
Soc. London, Bot. 9: 262. 1916.
Grammitis purpurascens (Nadeaud) Christenh., comb.
Grammitis seemannii (J.Sm.) Christenh., comb. nov.
nov. Basionym: Polypodium purpurascens Nadeaud,
Basionym: Cryptosorus seemannii J.Sm., Bonplandia
Énum. Pl. Tahiti: 26. 1873.
9: 262. 1861.
Grammitis reclinata (Brack.) Christenh., comb. nov.
Grammitis semicrypta (Copel.) Christenh., comb. nov.
Basionym: Polypodium reclinatum Brack., U.S. Expl.
Basionym: Prosaptia semicrypta Copel., Philipp. J.
Exped., Filic. 16: 11. 1854.
Sci., C 9: 231. 1914.
Grammitis reineckei (Christ) Christenh., comb. nov.
Grammitis semilunaris (Moguel & M.Kessler)
Basionym: Prosaptia reineckei Christ, Nachtr. Fl.
Christenh., comb. nov. Basionym: Terpsichore
Schutzgeb. Südsee [Schumann & Lauterbach]: 41.
semilunaris Moguel & M.Kessler, Phytotaxa 2: 42.
1905.
2009.
Grammitis repandula (Mett. ) Christenh., comb. nov.
Grammitis serriformis (Christ) Christenh., comb. nov.
Basionym: Polypodium repandulum Mett., Abh.
Basionym: Prosaptia serriformis Christ, Bull. Herb.
Senckenberg. Naturf. Ges. 2: 50. 1856.
Boissier 6: 143. 1898.
Grammitis rhodocarpa (Copel. ) Christenh., comb.
Grammitis sesquipinnata (Copel.) Christenh., comb.
nov. Basionym: Ctenopteris rhodocarpa Copel., Univ.
nov. Basionym: Ctenopteris sesquipinnata Copel.,
Calif. Publ. Bot. 18: 225. 1942.
Univ. Calif. Publ. Bot. 18: 225. 1942.
Grammitis rojasii Christenh., nom. nov. Replaced
Grammitis setulifera (Alderw.) Christenh., comb. nov.
synonym: Lellingeria pinnata A.Rojas, Lankesteriana
Basionym: Polypodium setuliferum Alderw., Bull.
6: 95. 2006, not: Grammitis pinnata (Hook.) F.Muell.
Jard. Bot. Buitenzorg Ser. 2, 16: 32. 1914.
Etymology: Named for Alexander Rojas, who first
described this species. Grammitis shawii (Copel.) Christenh., comb. nov.
Basionym: Polypodium shawii Copel., Philipp. J. Sci.,
Grammitis rosenstockii (Copel.) Christenh., comb. nov.
C 9: 6. 1914.
Basionym: Prosaptia rosenstockii Copel., Univ. Calif.
Publ. Bot. 12. 404. 1931. Grammitis simacensis (Rosenst.) Christenh., comb.
nov. Basionym: Polypodium simacense Rosenst.,
Grammitis rupicola (R.Y.Hirai & J.Prado) Christenh.,
Repert. Spec. Nov. Regni Veg. 25: 60. 1928.
comb. nov. Basionym: Moranopteris rupicola
Grammitis similis (M.Kessler & A.R.Sm. ) Christenh.,
comb. nov. Basionym: Ceradenia similis M.Kessler

& A.R.Sm., Organisms Diversity Evol. 8: 168. 2008. Bernice P. Bishop Mus. 93: 12. 1932.
Grammitis simplex (R.Y.Hirai & J.Prado) Christenh., Grammitis subglabra (M.Kato & Parris) Christenh.,
comb. nov. Basionym: Moranopteris simplex R.Y.Hirai comb. nov. Basionym: Prosaptia contigua C.Presl var.
& J.Prado, Taxon 60: 1135. 2011. subglabra M.Kato & Parris, J. Fac. Sci. Univ. Tokyo,
Grammitis sinica (Christ) Christenh., comb. nov. Bot. 15: 119. 1992.
Basionym: Polypodium sinicum Christ, Bull. Herb. Grammitis subhamatopilosa (Alderw.)
Boissier 7: 3. 1899. Christenh., comb. nov. Basionym: Polypodium
Grammitis sinohirtella (Parris) Christenh., comb. nov. subhamatopilosum Alderw., Bull. Jard. Bot.
Basionym: Oreogrammitis sinohirtella Parris, Fl. Buitenzorg ser. 3, 5: 220. 1922.
China 2-3: 842. 2013. Grammitis subimpressa (Copel.) Christenh., comb.
Grammitis sinuata (T.Moore) Christenh., comb. nov. nov. Basionym: Ctenopteris subimpressa Copel.,
Basionym: Gymnogramma sinuata T.Moore, Ann. Philipp. J. Sci. 84: 463. 1956.
Bot. (Oxford) 5: 483. 1891. Synonym: Enterosora Grammitis subminuta (Alderw.) Christenh., comb.
asplenioides L.E.Bishop, Syst. Bot. 17: 358. 1992, nov. Basionym: Polypodium subminutum Alderw.,
not: Grammitis asplenioides C.Presl Malayan Ferns : 598. 1909.
Grammitis sklenarii (Lehnert) Christenh., comb. nov. Grammitis subnuda (Mett. ex Kuhn) Christenh., comb.
Basionym: Melpomene sklenarii Lehnert, Amer. Fern nov. Basionym: Polypodium subnudum Mett. ex
J. 98: 241. 2009. Kuhn, Linnaea 36: 130. 1869.
Grammitis smithii (A.Rojas) Christenh., comb. nov. Grammitis subpinnata (Baker) Christenh., comb. nov.
Basionym: Terpsichore smithii A.Rojas, Mét. Ecol. Basionym: Polypodium subpinnatum Baker, J. Linn.
Sist. 3: 15. 2008. Soc., Bot. 15: 419. 1876.
Grammitis sparsipilosa (Holttum) Christenh., comb. Grammitis subrepandula (Christ) Christenh., comb.
nov. Basionym: Ctenopteris sparsipilosa Holttum, nov. Basionym: Polypodium subrepandulum Christ,
Gard. Bull. Singapore 11: 268. 1947. Ann. Jard. Bot. Buitenzorg II. 5: 119. 1905.
Grammitis spathulata (A.R.Sm.) Christenh., comb. Grammitis subsecundodissecta (Zoll.) Christenh., comb.
nov. Basionym: Terpsichore spathulata A.R.Sm., nov. Basionym: Polypodium subsecundodissectum
Novon 3: 483. 1993. Zoll., Syst. Verz. 38, 48. 1854.
Grammitis spixiana (Mett.) Christenh., comb. nov. Grammitis subtriangularis (Alderw.) Christenh., comb.
Basionym: Polypodium spixianum Mett., Abh. nov. Basionym: Polypodium subtriangulare Alderw.,
Senckenberg. Naturf. Ges. 2: 57. 1856. Bull. Dept. Agric. Indes Neerl. 18: 20. 1908.
Grammitis spongiosa (Maxon) Christenh., comb. nov. Grammitis subulatipinna (Alderw.) Christenh., comb.
Basionym: Enterosora spongiosa Maxon, Proc. Biol. nov. Basionym: Polypodium subulatipinnum Alderw.,
Soc. Washington 52: 113. 1939. Nova Guinea 14: 46. 1924.
Grammitis stella (Copel.) Christenh., comb. nov. Grammitis sunduei Christenh., nom. nov. Replaced
Basionym: Ctenopteris stella Copel., Philipp. J. Sci. synonym: Mycopteris longipilosa Sundue, Brittonia
84: 452. 1956. 66: 179. 2013, not: Grammitis longipilosa (C.Chr.)
Grammitis stellatosetosa (Copel.) Christenh., comb. Christenh. Etymology: Named for Michael Sundue,
nov. Basionym: Ctenopteris stellatosetosa Copel., who first described this species.
Univ. Calif. Publ. Bot. 18: 225. 1942. Grammitis taiwanensis (Parris & Ralf Knapp)
Grammitis stenobasis (Baker) Christenh., comb. nov. Christenh., comb. nov. Basionym: Radiogrammitis
Basionym: Polypodium stenobasis Baker, Ann. Bot. taiwanensis Parris & Ralf Knapp, Fl. China 2-3: 843.
(Oxford) 8: 130. 1894. 2013.
Grammitis steyermarkii (Labiak) Christenh., comb. Grammitis tamandarei (Rosenst.) Christenh., comb.
nov. Basionym: Terpsichore steyermarkii Labiak, nov. Basionym: Polypodium tamandarei Rosenst.,
Brittonia 52: 254. 2000. Hedwigia 56: 369. 1915.
Grammitis strangeana (Pic.Serm.) Christenh., comb. Grammitis taxodioides (Baker) Christenh., comb. nov.
nov. Basionym: Xiphopteris strangeana Pic.Serm., Basionym: Polypodium taxodioides Baker, J. Bot. 17:
Webbia 27: 453. 1972. 42. 1879.
Grammitis stuebelii (Hieron.) Christenh., comb. nov. Grammitis terpsichore Christenh., nom. nov. Replaced
Basionym: Polypodium stuebelii Hieron., Hedwigia synonym: Polypodium flexuosum Maxon, Contr. U.S.
48: 252. 1909. Natl. Herb. 17: 597. 1916, not: Grammitis flexuosa
Humb. & Bonpl. Etymology: Τερψιχόρη, the Muse of
Grammitis subcoriacea (Copel.) Christenh., comb. nov.
Dance in Ancient Greek mythology.
Basionym: Polypodium subcoriaceum Copel., Bull.
Grammitis thalia Christenh., nom. nov. Replaced

49
synonym: Polypodium emersonii (Hook. & Grev.) Univ. Calif. Publ. Bot. 18: 225. 1942.
C.Chr. var. samoense C.Chr., Bishop Mus. Bull. Bot. Grammitis vodonaivalui (Brownlie) Christenh., comb.
177: 111. 1943, not: Grammitis samoensis (Baker) nov. Basionym: Ctenopteris vodonaivalui Brownlie,
Ching. Etymology: Θάλεια, the Muse of Comedy in Nova Hedwigia Beih. 55: 357. 1977.
Ancient Greek mythology. Grammitis vomaensis (Brownlie) Christenh., comb.
Grammitis thelxinoe Christenh., nom. nov. Replaced nov. Basionym: Ctenopteris vomaensis Brownlie,
synonym: Prosaptia archboldii Copel., Univ. Calif. Nova Hedwigia Beih. 55: 354. 1977.
Publ. Bot. 18: 226. 1942, not: Grammitis archboldii Grammitis walleri (Maiden & Betche) Christenh.,
C.Chr. Etymology: Θελξινόη, the Muse of Happiness comb. nov. Basionym: Polypodium walleri Maiden &
in Ancient Greek mythology. Betche, Proc. Linn. Soc. New South Wales 35: 799.
Grammitis tintinnabula Christenh., nom. nov. Replaced 1910.
synonym: Prosaptia linearis Copel., Philipp. J. Sci., C Grammitis whartoniana (C.Chr.) Christenh., comb.
4: 115. 1909, not: Grammitis linearis Sw. Etymology: nov. Basionym: Polypodium whartonianum C.Chr.,
Named for a roman wind chime, tintinnabula, often Brittonia 2: 309. 1937.
of phallic shape.
Grammitis wooronooran (Bailey) Christenh., comb.
Grammitis toppingii (Copel.) Christenh., comb. nov. nov. Basionym: Vittaria wooronooran Bailey, not:
Basionym: Prosaptia toppingii Copel., Philipp. J. Sci. Grammitis wurunuran Parris
1(Suppl. 2): 158. 1906.
Grammitis wurdackii (L.E.Bishop) Christenh., comb.
Grammitis torulosa (Baker) Christenh., comb. nov. nov. Basionym: Cochlidium wurdackii L.E.Bishop,
Basionym: Polypodium torulosum Baker, J. Linn. Soc., Amer. Fern J. 68: 93. 1978.
Bot. 16: 204. 1877.
Grammitis yoderi (Copel.) Christenh., comb. nov.
Grammitis translucens (Parris & Jaman) Christenh., Basionym: Polypodium yoderi Copel., Philipp. J. Sci.
comb. nov. Basionym: Oreogrammitis translucens 1(Suppl. 2): 161. 1906.
Parris & Jaman, Sains Malaysiana 41: 299. 2012.
Grammitis zenkeri (Hieron.) Christenh., comb. nov.
Grammitis tristis (A.R.Sm. ) Christenh., comb. nov. Basionym: Polypodium zenkeri Hieron., Bot. Jahrb.
Basionym: Ceradenia tristis A.R.Sm., Novon 3: 184. Syst. 46: 385. 1911.
1993.
Lastreopsis poecilophlebia (Hook.) Christenh., comb.
Grammitis tryoniorum (B.León & A.R.Sm.) Christenh., nov. Basionym: Polypodium poecilophlebium Hook.,
comb. nov. Basionym: Ceradenia tryoniorum B.León Sp. Filic. 5: 14. 1863.
& A.R.Sm., Amer. Fern J. 93: 81. 2003.
Lomariopsis pteridiformis (Ces.) Christenh., comb.
Grammitis tsatchelae (Labiak) Christenh., comb. nov. nov. Basionym: Gymnogramma pteridiformis Ces.,
Basionym: Melpomene tsatchelae Labiak, Brittonia Rendiconto Accad. Sci. Soc. Borbon. Napoli 16: 27,
52: 251. 2000. 30. 1877.
Grammitis tunquiniensis (M.Kessler & A.R.Sm.) Selliguea amplexifolia (Christ) Christenh., comb. nov.
Christenh., comb. nov. Basionym: Ceradenia Basionym: Polypodium amplexifolium Christ, J. Bot.
tunquiniensis M.Kessler & A.R.Sm., Organisms (Morot) sér. 2, 1: 237, 269. 1908.
Diversity Evol. 8: 169. 2008.
Selliguea brachypodia (Copel.) Christenh., comb. nov.
Grammitis urania Christenh., nom. nov. Replaced Basionym: Polypodium brachypodium Copel., Phil.
synonym: Polypodium cucullatum Nees & Blume, Journ. Sci., C 12: 62. 1917.
Nova Acta Regiae Soc. Sci. Upsal., ser. 2, 11: 121. 3.
Selliguea cesatiana (Baker) Christenh., comb. nov.
1823, not: Grammitis cucullata Blume. Etymology:
Basionym: Polypodium cesatianum Baker, J. Bot. 42.
Οὐρανία, the Muse of Astronomy in Ancient Greek
1879.
mythology.
Selliguea cyrtomioides (S.G.Lu & C.D.Xu) Christenh.,
Grammitis urceolaris (Hayata) Christenh., comb. nov.
comb. nov. Basionym: Arthromeris cyrtomioides
Basionym: Polypodium urceolare Hayata, Icon. Pl.
S.G.Lu & C.D.Xu, Acta Phytotax. Sin. 45: 83. 2007.
Formosan. 5: 324. 1915.
Selliguea elegans (Ching) Christenh., comb. nov.
Grammitis vareschii (A.R.Sm. & Mostacero) Christenh.,
Basionym: Arthromeris elegans Ching, Sunyatsenia
comb. nov. Basionym: Galactodenia vareschii
6: 8. 1941.
A.R.Sm. & Mostacero, Syst. Bot. 37: 346. 2012.
Selliguea espinosae (Weath.) Christenh., comb. nov.
Grammitis vargasiana (A.Rojas) Christenh., comb. nov.
Basionym: Polypodium espinosae Weath., Contr.
Basionym: Lellingeria vargasiana A.Rojas, Brenesia
Gray Herb. 85: 15. 1929.
75-76: 8. 2011.
Selliguea feuillei (Bertero) Christenh., comb. nov.
Grammitis venulosoides (Copel.) Christenh., comb.
Basionym: Polypodium feuillei Bertero, Mercurio
nov. Basionym: Ctenopteris venulosoides Copel.,

Chileno: 745. 1829. Basionym: Polypodium tenuicaudum Hook., Sp. Filic.

Selliguea fraserjenkinsii (Kholia) Christenh., comb. 5: 90. 1864, as’ tenuicauda’.
nov. Basionym: Pichisermollodes fraserjenkinsii Selliguea tomentosa (W.M.Chu) Christenh., comb. nov.
Kholia, Ferns & Fern-Allies Sikkim 2: 137. 2014, as Basionym: Arthromeris tomentosa W.M.Chu, Acta
’fraser-jenkinsonii’. Bot. Yunnan., Suppl. 5: 53.1992.
Selliguea himalayensis (Hook.) Christenh., comb. nov. Selliguea translucens (Kunze) Christenh., comb. nov.
Basionym: Polypodium himalayense Hook., Sp. Filic. Basionym: Polypodium translucens Kunze, Anal.
5: 91. 1864. Pterid.: 16. 1837.
Selliguea himalovata (Fraser-Jenk. & Kandel) Selliguea wardii (C.B.Clarke) Christenh., comb. nov.
Christenh., comb. nov. Basionym: Arthromeris Basionym: Polypodium wardii C.B.Clarke, J. Linn.
himalovata Fraser-Jenkins & Kandel, Ferns & Fern- Soc., Bot. 25: 99. 1890.
Allies Nepal 1: 34. 2015.
Selliguea hovenkampiana Christenh., nom. nov. G4. Welwitschiaceae Reference: Leuenberger (2001).
Replaced synonym: Polypodium wallichianum Note: This taxon is geographically isolated with
Spreng., Syst. Veg. 4: 53. 1827, not: Seliguea numerous differences in male cone structure
wallichiana Hook. Etymology: Named for Peter compared to the other known species, Welwitschia
Hovenkamp, for his contribution to the taxonomy of mirabilis Hook., and is probably better treated as a
Asian Polypodiaceae. separate species. It is the second species known for
Selliguea lehmannii (Mett.) Christenh., comb. nov. this formerly monotypic family.
Basionym: Polypodium lehmannii Mett., Abh. Welwitschia namibiana (Leuenb.) Christenh. & Byng,
Senckenberg. Naturf. Gesell. 1: 117. 1856. comb. & stat. nov. Basionym: Welwitschia mirabilis
Selliguea leucophora (Baker) Christenh., comb. nov. Hook.f. subsp. namibiana Leuenb., Willdenowia 31:
Basionym: Polypodium leucophorum Baker, J. Linn. 375. 2001.
Soc., Bot. 22: 229. 1886.
Selliguea lui Christenh., nom. nov. Replaced synonym: 30. Alismataceae. Reference: Lehtonen & Myllys
Arthromeris nigropaleacea S.G.Lu, Indian Fern J. (2008).
14: 144. 1997, not: Selliguea nigropaleacea (Ching)
Note: Echinodorus berteroi (Spreng.) Fassett, type of
S.G.Lu, Hovenkamp & M.G.Gilbert. Etymology:
the genus Echinodorus, is found separate from the
Named for Shu Gang Lu, who first described this
remainder of Echinodorus, often with other genera
species.
like Albidella, Alisma or Baldellia interdigitated.
Selliguea lungtauensis (Ching) Christenh., comb. nov. A new genus for the latter is therefore hereby
Basionym: Arthromeris lungtauensis Ching, Contr. proposed.
Inst. Bot. Natl. Acad. Peiping 2: 98. 1933.
Aquarius Christenh. & Byng, gen. nov. Perennial
Selliguea mairei (Brause) Christenh., comb. nov. aquatic plants, submerged or emergent. Similar to
Basionym: Polypodium mairei Brause, Hedwigia 64: Echinodorus, but not annual and lacking winged ribs
208, 1914. on the achenes. Type: A. cordifolius (L.) Christenh. &
Selliguea phuluangensis (Tagawa & K.Iwats.) Byng. Etymology: Named for the star sign Aquarius,
Christenh., comb. nov. Basionym: Arthromeris as this is a plant of aquatic nature.
phuluangensis Tagawa & K.Iwats., Acta Phytotax. Aquarius bracteatus (Micheli) Christenh. & Byng, comb.
Geobot. 22: 100. 1967. nov. Basionym: Echinodorus bracteatus Micheli
Selliguea proavita (Copel.) Christenh., comb. nov. in A.L.P.de Candolle & A.C.P.de Candolle (eds.),
Basionym: Polypodium proavitum Copel., Philipp. J. Monogr. Phan. 3: 59. 1881.
Sci., C 8: 347. 1909. Aquarius cordifolius (L.) Christenh. & Byng, comb. nov.
Selliguea proteus (Copel.) Christenh., comb. nov. Basionym: Alisma cordifolium L., Sp. Pl.: 343. 1753.
Basionym: Polypodium proteus Copel., Philipp. J. Sci. Aquarius cylindricus (Rataj) Christenh. & Byng, comb.
1(Suppl. 2): 164, 1906. nov. Basionym: Echinodorus cylindricus Rataj, Stud.
Selliguea subebenipes (Ching) Christenh., comb. nov. CSA V 2: 66. 1975.
Basionym: Phymatopsis subebenipes Ching, Acta Aquarius decumbens (Kasselm.) Christenh. & Byng,
Phytotax. Sin. 9: 193. 1964. comb. nov. Basionym: Echinodorus decumbens
Selliguea tatsienensis (Franch & Bureau) Christenh., Kasselm., Aqua Pl. 25: 5. 2000.
comb. nov. Basionym: Polypodium tatsienense Aquarius densinervis (Somogyi) Christenh. & Byng,
Franch. & Bureau in Christ, Bull. Soc. Bot. France 52 comb. nov. Basionym: Echinodorus densinervis
(Mém. 1): 19. 1905.
Selliguea tenuicauda (Hook.) Christenh., comb. nov.

51
Figure 7: Angiosperms (Monocots). Oceana serrulata (R.Br.) Byng & Christenh., comb. nov.. Image used with
permission by Naturalis Biodiversity Center.

Somogyi, Phyton (Horn) 51: 151. 2011. Bull. 63: 548. 2009.
Aquarius emersus (Lehtonen) Christenh. & Byng, comb. Aquarius scaber (Rataj) Christenh. & Byng, comb. nov.
nov. Basionym: Echinodorus emersus Lehtonen, Kew Basionym: Echinodorus scaber Rataj, Folia Geobot.
Bull. 63: 538. 2009. Phytotax. 4: 438. 1969.
Aquarius floribundus (Seub.) Christenh. & Byng, comb. Aquarius subulatus (Mart. ex Schult.f.) Christenh.
nov. Basionym: Alisma floribundum Seub. in C.F.P. & Byng, comb. nov. Basionym: Alisma subalatum
von Martius (ed.), Fl. Bras. 3(1): 109. 1847. Mart. ex Schult.f. in J.J.Roemer & J.A.Schultes, Syst.
Aquarius glaucus (Rataj) Christenh. & Byng, comb. Veg., ed. 15 bis 7: 1609. 1830.
nov. Basionym: Echinodorus glaucus Rataj, Stud. CSA Aquarius trialatus (MicFassettheli) Christenh. & Byng,
V 2: 69. 1975. comb. nov. Basionym: Echinodorus trialatus Fassett,
Aquarius grandiflorus (Cham. & Schltdl.) Christenh. & Rhodora 57: 179. 1955.
Byng, comb. nov. Basionym: Alisma grandiflorum Aquarius tunicatus (Small) Christenh. & Byng, comb.
Cham. & Schltdl., Linnaea 2: 152. 1827. (Figure 6A) nov. Basionym: Echinodorus tunicatus Small, N.
Aquarius grisebachii (Small) Christenh. & Byng, comb. Amer. Fl. 17: 48. 1909.
nov. Basionym: Echinodorus grisebachii Small, N. Aquarius uruguayensis (Arechav.) Christenh. & Byng,
Amer. Fl. 17: 46. 1909. comb. nov. Basionym: Echinodorus uruguayensis
Aquarius horizontalis (Rataj) Christenh. & Byng, Arechav., Anales Mus. Nac. Montevideo 4: 66. 1902.
comb. nov. Basionym: Echinodorus horizontalis
Rataj, Folia Geobot. Phytotax. 4: 335. 1969. 32. Hydrocharitaceae. References: Les et al. (2006),
Aquarius inpai (Rataj) Christenh. & Byng, comb. nov. Chen et al. (2012).
Basionym: Echinodorus inpai Rataj, Akvar. Terar. 24: Note: Egeria Planch. is embedded in Elodea Michx. and
20. 1981. is hereby transferred. Limnobium Rich. is part of
Aquarius lanceolatus (Rataj) Christenh. & Byng, comb. Hydrocharis L. and the combination is here made.
nov. Basionym: Echinodorus lanceolatus Rataj, Bull. Elodea heterostemon (S.Koehler & C.P.Bove)
Jard. Bot. Natl. Belg. 38: 406. 1968. Byng & Christenh., comb. nov. Basionym:
Aquarius longipetalus (Micheli) Christenh. & Byng, Egeria heterostemon S.Koehler & C.P.Bove, Novon
comb. nov. Basionym: Echinodorus longipetalus 11: 63, 2001.
Micheli in A.L.P.de Candolle & A.C.P.de Candolle Hydrocharis laevigata (Humb. & Bonpl. ex Willd.)
(eds.), Monogr. Phan. 3: 60. 1881. Byng & Christenh., comb. nov. Basionym: Salvinia
Aquarius longiscapus (Arechav.) Christenh. & Byng, laevigata Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5:
comb. nov. Basionym: Echinodorus longiscapus 537. 1810.
Arechav., Anales Mus. Nac. Montevideo 4(1): 67.
1902.
41. Cymodoceaceae. Reference: Petersen et al. (2014).
Aquarius macrophyllus (Kunth) Christenh. & Byng,
Note: Molecular and morphological evidence warrants
comb. nov. Basionym: Alisma macrophyllum Kunth,
a separate genus status for Cymodocea serrulata
Enum. Pl. 3: 151. 1841.
(R.Br.) Asch. & Magnus, which is described below.
Aquarius major (Micheli) Christenh. & Byng, comb.
Oceana Byng & Christenh., gen. nov. Dioecious marine
nov. Basionym: Echinodorus martii var. major
plants. Rhizomes creeping, thick, smooth. Leaves
Micheli in A.L.P.de Candolle & A.C.P.de Candolle
linear with rounded apices, and serrulate margins,
(eds.), Monogr. Phan. 3: 49 (1881).
intrafoliar, leaf scars open, ligules narrowly truncate.
Aquarius palifolius (Nees & Mart.) Christenh. & Byng, Perianth lacking. Staminate flowers stalked, stamens
comb. nov. Basionym: Sagittaria palifolia Nees & two. Pistillate flowers sessile, with two free ovaries,
Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.- stigma subsessile, placentation apical. Fruit non-
Carol. Nat. Cur. 11: 21. 1823. fleshy. Type: Oceana serrulata (R.Br.) Byng &
Aquarius paniculatus (Micheli) Christenh. & Byng, Christenh. Distribution: Egypt to Mozambique and
comb. nov. Basionym: Echinodorus paniculatus Madagascar to the Western Pacific. Note: The genus
Micheli in A.L.P.de Candolle & A.C.P.de Candolle is similiar to Cymodocea K.D. Koenig but differs by
(eds.), Monogr. Phan. 3: 51. 1881. not having the leaf sheaths completely surrounding
Aquarius pubescens (Mart. ex Schult.f.) Christenh. the stem and broadly triangular, leaving open scars,
& Byng, comb. nov. Basionym: Alisma pubescens and the distinctly serrulate leaf margins. Etymology:
Mart. ex Schult.f. in J.J.Roemer & J.A.Schultes, Syst. Named for Oceanus, divine personification of the
Veg., ed. 15 bis 7: 1608. 1830. sea in Greek and Roman mythology.
Aquarius reptilis (Lehtonen) Christenh. & Byng, comb. Oceana serrulata (R.Br.) Byng & Christenh., comb. nov.
nov. Basionym: Echinodorus reptilis Lehtonen, Kew Basionym: Caulinia serrulata R.Br., Prodr. Fl. Nov.

53
A B
2 mm
C D
2 mm
1 mm
E F
1 mm
0.5 mm
Figure 6: Angiosperms (Monocots). A, Aquarius grandiflorus (Cham. & Schltdl.) Christenh. & Byng; B–F, Oxygyne
confusa Bidault, Merckx & Byng - B–C, flowering plants; D, open perianth; E, immature capsule with persistent
perianth; and F, mature open fruit showing seeds. Images: A, © Maarten Christenhusz, B–F, © Ehoarn Bidault.

Holland.: 339. 1810. Type: AUSTRALIA. R. Brown as a new species of Thismia Griff. and gave it the
5813 (holotype BM [BM000907122]!). (Figure 7) name “T. intermedia” because he thought the species
had intermediate characters between Thismia and
Bagnisia Becc., but the species was never described.
44. Burmanniaceae. References: Merckx et al. (2006,
Thassilo Franke studied several mycoheteotrophic
2009, 2013, 2017).
species in Cameroon (Franke 2004, Franke et al.
Note: While reviewing the taxonomy of Dioscoreales 2004, Sainge & Franke 2005, Sainge et al. 2005) and
(e.g. Merckx et al., 2013) an interesting specimen determined the specimen as possibly belonging to
from the Central African Republic was discovered in Oxygyne, but was uncertain about the number of
the genus Oxygyne Schltr. and is described here as stamens and so did not describe the species. As a
a new species. Additionally Gymnosiphon Blume is result we decided to give the species the epithet
expanded to include Cymbocarpa Miers. ‘confusa’ because of its long history of confusion by
Gymnosiphon saccatum (Sandwith) Merckx & several botanists.
Byng, comb. nov. Basionym: Cymbocarpa saccata
Sandwith, Bull. Misc. Inform. Kew 1931(1): 60. 1931.
46. Triuridaceae Reference: Mennes et al. (2013).
Gymnosiphon urbanii (K.I.Goebel & Sussenguth)
Note: The pantropical genus Sciaphila Blume is
Merckx & Byng, comb. nov. Basionym: Cymbocarpa
paraphyletic due to the inclusion of Seychellaria
urbanii K.I.Goebel & Sussenguth, Flora 117: 80.
Hemsley and Andruris Schltr. and to maintain these
1924, as ‘urbani’.
genera as distinct would require Sciaphila to be split
Oxygyne confusa Bidault, Merckx & Byng, sp. nov. into several genera. Morphologically Seychellaria
(Figures 6B–F). The species differs from the other and Andruris can be distinguished from Sciaphila
known African Oxygyne species, O. triandra mainly based on the presence of staminodes and
Schltr., in its smaller flower size and by having appendages on the connectives, respectively, in male
protruding stamens inserted on the perianth lobes, flowers. If Sciaphila were to be split it is highly likely
1 mm above their base. It is an achlorophyllous, any differences between any resulting genera would
mycoheterotrophic, glabrous herb. Roots are be distinguishable by similar minute floral characters
adventitious on a creeping stem, up to 2 mm long. or completely lacking any synapomorphies. We
Stems are simple, creeping for 1 cm and erect at apex therefore prefer to synonymise both Seychellaria
for 5 mm. Inflorescence 1 or 2-flowered; peduncle and Andruris with Sciaphila.
ca. 0.5–1.0 mm long, with 3–8 bracts at the base of
Sciaphila barbata (Nuraliev & Cheek) Merckx & Byng,
flowers, scale-like, 2–3 mm long, lanceolate, apex
comb. nov. Basionym: Seychellaria barbata Nuraliev
acute, green, caducous. Flowers are erect, green
& Cheek, Phytotaxa 268: 23. 2016.
at base, brown-yellowish, 7–8 × 2–3 mm. Perianth
segments are fused, 6-lobed, tube campanulate, Sciaphila mwanihanensis Merckx & Byng, nom. nov.
2.0–2.8 mm long, longitudinal veins on the outer Replaced synonym: Seychellaria africana Vollesen,
surface; perianth lobes are white, filiform, 3–6 mm Kew Bull. 36: 733. 1982, not: Sciaphila africana
long; perianth with an annular corona at throat, ca. A.Chev. Etymology: Named after the Mwanihana
0.6 mm long, apex lobed. Stamens 3, equal, attached Forest Reserve in Tanzania, the only known locality
to base of perianth lobes, protruding; filaments ca. of the species.
1 mm long, inflexed; anthers basifixed, bithecal, Sciaphila madagascariensis (C.H.Wright) Merckx
introrse, 0.5 mm long, white. Ovary ca. 1.0 × 0.7 mm & Byng, comb. nov. Basionym: Seychellaria
long, white; placentation parietal, ovules many (ca. madagascariensis C.H.Wright, Bull. Misc. Inform.
40), ca. 0.1 mm in diameter; style 1, ca. 1 mm long, Kew 1912: 196. 1912.
thick, ending in 3 short, triangular stigmatic branches, Sciaphila perrieri (Schltr.) Merckx & Byng, comb. nov.
ca. 0.25 × 0.2 mm. Fruits are capsular, ovoid, 1.5–2.0 Basionym: Seychellaria perrieri Schltr., Notizbl. Bot.
× 1.0–1.2 mm, opening apically, persistent perianth. Gart. Berlin-Dahlem 7: 318. 1923.
Seeds polygonal, 0.3–0.4 × 0.15–0.2 mm, slightly Sciaphila thomassetii (Hemsl.) Merckx & Byng, comb.
flattened, testa reticulate, radicle occasionally nov. Basionym: Seychellaria thomassetii Hemsl.,
emergent at the apex. Type: CENTRAL AFRICAN Ann. Bot. (Oxford) 21: 74. 1907.
REPUBLIC. Balimbwa, 40 km north of Bambari, 28 July
1928, C.Tisserant 2623 (holotype P [P00319729]!;
isotypes BM [BM000803697], P [P00319728]!). 66. Hypoxidaceae. Reference: Kocyan et al. (2011).
Note and etymology: The sole specimen for this Note: Curculigo Gaertn. is paraphyletic if genera like
species has had a long determination history. When Molineria Colla and Hypoxidia F.Friedmann are
Charles Tisserant collected his 2653 specimen, he accepted. We hereby treat it in the broad sense,
preliminary identified it as Hydnora Thunb. In 1983, because morphologically these species are relatively
Luis Diego Gómez Pignataro identified the specimen

55
similar. A.Gil & M.C.E.Amaral, Phytotaxa 164: 48. 2014.

Curculigo maheensis (F.Friedmann) Christenh. & Trimezia elegans (Ravenna) Christenh., comb. nov.
Byng, comb. nov. Basionym: Hypoxidia maheensis Basionym: Pseudotrimezia elegans Ravenna, Onira
F.Friedmann, Bull. Mus. Natl. Hist. Nat., B, Adansonia 1: 48. 1988.
sér. 4, 6: 459. 1985. Trimezia floscella (A.Gil & M.C.E.Amaral) Christenh.
Curculigo fakimensis (Odyuo, D.K.Roy & Khamdi) & Byng, comb. nov. Basionym: Neomarica floscella
Christenh. & Byng, comb. nov. Basionym: Molineria A.Gil & M.C.E.Amaral, Phytotaxa 164: 51. 2014.
fakimense Odyuo, D.K.Roy & Khamdi, Rheedea 26: Trimezia fulva (Ravenna) Christenh. & Byng, comb.
131. 2016. nov. Basionym: Pseudotrimezia fulva Ravenna,
Onira 5: 5. 2000.
70. Iridaceae References: Rodriguez & Sytsma (2006), Trimezia gracilis (Herb.) Christenh. & Byng, comb. nov.
Lovo et al. (2012). Basionym: Marica gracilis Herb., Bot. Mag. 66: t.
Note: Trimezia Salisb. ex Herb. is expanded to include 3713. 1839.
the entire tribe Trimezieae, as the traditional genera Trimezia imbricata (Hand.-Mazz.) Christenh. & Byng,
are not monophyletic and differences between comb. nov. Basionym: Marica imbricata Hand.-
genera are minute. Tigridia Juss. is thus expanded to Mazz., Denkschr. Kaiserl. Akad. Wiss., Wien. Math.-
prevent polyphyly. Naturwiss. Kl. 79: 215. 1908.
Tigridia citrina (Cruden) Christenh. & Byng, comb. nov. Trimezia involuta (A.Gil & M.C.E.Amaral) Christenh.
Basionym: Sessilanthera citrina Cruden, Brittonia 27: & Byng, comb. nov. Basionym: Neomarica involuta
108. 1975. A.Gil & M.C.E.Amaral, Phytotaxa 286: 93. 2016.
Tigridia heliantha (Ravenna) Christenh. & Byng, comb. Trimezia itacambirae (Ravenna) Christenh. & Byng,
nov. Basionym: Nemastylis latifolia subsp. heliantha comb. nov. Basionym: Pseudotrimezia itacambirae
Ravenna, Bonplandia (Corrientes) 2: 283. 1968. Ravenna, Revista Inst. Munic. Bot. 3: 26. 1969.
Trimezia barretoi (R.C.Foster) Christenh. & Byng, Trimezia laevis (Ravenna) Christenh. & Byng, comb.
comb. nov. Basionym: Pseudotrimezia barretoi nov. Basionym: Pseudotrimezia laevis Ravenna,
R.C.Foster, Contr. Gray Herb. 155: 8. 1945. Onira 5: 7. 2000.
Trimezia brevistaminea (Chukr) Christenh. & Byng, Trimezia longifolia (Link & Otto) Christenh. & Byng,
comb. nov. Basionym: Pseudotrimezia brevistaminea comb. nov. Basionym: Marica longifolia Link & Otto,
Chukr, Cad. Estud. Pesq., UNIP (São Paulo) 7: 3. 2001. Icon. Pl. Select. 10: 123. 1828. (Figure 8A)
Trimezia castaneomaculata (A.Gil & M.C.E.Amaral) Trimezia monticola (Klatt) Christenh. & Byng, comb.
Christenh. & Byng, comb. nov. Basionym: Neomarica nov. Basionym: Lansbergia monticola Klatt in Mart.,
castaneomaculata A.Gil & M.C.E.Amaral, Phytotaxa Fl. Bras. 3(1): 526. 1871.
286: 90. 2016. Trimezia nana (Lovo & Mello-Silva) Christenh. & Byng,
Trimezia cipoana (Ravenna) Christenh. & Byng, comb. comb. nov. Basionym: Pseudotrimezia nana Lovo &
nov. Basionym: Pseudotrimezia cipoana Ravenna, Mello-Silva, Phytotaxa 195: 146. 2015.
Revista Inst. Munic. Bot. 3: 25. 1969. Trimezia pumila (Ravenna) Christenh. & Byng, comb.
Trimezia concava (Ravenna) Christenh. & Byng, comb. nov. Basionym: Pseudotrimezia pumila Ravenna,
nov. Basionym: Pseudotrimezia concava Ravenna, Onira 5: 8. 2000.
Onira 1: 49. 1988. Trimezia recurvata (Ravenna) Christenh. & Byng,
Trimezia concolor Christenh. & Byng, nom. nov. comb. nov. Basionym: Pseudotrimezia recurvata
Replaced synonym: Pseudotrimezia gracilis Chukr, Ravenna, Revista Inst. Munic. Bot. 3: 27. 1969.
Bol. Bot. Univ. São Paulo 13: 104. 1992, not: Trimezia Trimezia rigida (Ravenna) Christenh. & Byng, comb.
gracilis (Herb.) Christenh. Etymology: From Latin, nov. Basionym: Trimezia caerulea subsp. rigida
of the same colour, because this species lacks any Ravenna, Onira 1: 13. 1988.
markings in the flower. Trimezia sergipensis (A.Gil & M.C.E.Amaral) Christenh.
Trimezia datensis (Ravenna) Christenh. & Byng, comb. & Byng, comb. nov. Basionym: Neomarica
nov. Basionym: Pseudotrimezia datensis Ravenna, sergipensis A.Gil & M.C.E.Amaral, Nordic J. Bot. 34:
Onira 11: 25. 2007. 559. 2016.
Trimezia diamantinensis (Ravenna) Christenh., comb. Trimezia setacea (Klatt) Christenh. & Byng, comb. nov.
nov. Basionym: Pseudotrimezia diamantinensis Basionym: Lansbergia setacea Klatt, Abh. Naturf.
Ravenna, Onira 5: 6. 2000. Ges. Halle 15: 364. 1882.
Trimezia eburnea (A.Gil & M.C.E.Amaral) Christenh. Trimezia speciosa (Chukr & A.Gil) Christenh. & Byng,
& Byng, comb. nov. Basionym: Neomarica eburnea comb. nov. Basionym: Pseudiris speciosa Chukr &

A B
Figure 8: Angiosperms (Monocots). A, Trimezia longifolia (Link & Otto) Christenh. & Byng; B, Gilliesia atropurpurea
(Phil.) M.F.Fay & Christenh.; C, Hippeastrum atamasco (L.) Christenh. & Byng; D, Hippeastrum zephyranthum
Christenh. & Byng; E, Dracaena dawei (Stapf) Byng & Christenh.; and F, Dracaena parva (N.E.Br.) Byng & Christenh..
All images © Maarten Christenhusz.
57
A.Gil, Proc. Calif. Acad. Sci., ser. 4, 59: 725. 2008. Habranthus albispiritus Ravenna, Onira 1: 54. 1988.
Trimezia striata (Lovo & Mello-Silva) Christenh. & Hippeastrum albolilacinum (Cárdenas) Christenh.
Byng, comb. nov. Basionym: Pseudotrimezia striata & Byng, comb. nov. Basionym: Zephyranthes
Lovo & Mello-Silva, Phytotaxa 195: 149. 2015. albolilacina Cárdenas, Pl. Life 29: 39. 1973.
Trimezia synandra (Ravenna) Christenh. & Byng, comb. Hippeastrum amambaicum (Ravenna)
nov. Basionym: Pseudotrimezia synandra Ravenna, Christenh. & Byng, comb. nov. Basionym:
Bol. Soc. Argent. Bot. 10: 320. 1965. Habranthus amambaicus Ravenna, Onira 9: 14.
Trimezia tenuissima (Ravenna) Christenh. & Byng, 2003.
comb. nov. Basionym: Pseudotrimezia tenuissima Hippeastrum americanum (Hoffmanns.) Christenh. &
Ravenna, Revista Inst. Munic. Bot. 3: 28. 1969. Byng, comb. nov. Basionym: Sternbergia americana
Trimezia warmingii (Klatt) Christenh. & Byng, comb. Hoffmanns., Verz. Pfl.-Kult.: 179. 1824.
nov. Basionym: Marica warmingii Klatt, Abh. Naturf. Hippeastrum amoenum (Phil.) Christenh. & Byng,
Ges. Halle 15: 375. 1882. comb. nov. Basionym: Placea amoena Phil., Anales
Trimezia xyridea (Ravenna) Christenh. & Byng, comb. Univ. Chile 93: 145. 1896.
nov. Basionym: Pseudotrimezia xyridea Ravenna, Hippeastrum andalgalense (Ravenna)
Onira 1: 50. 1988. Christenh. & Byng, comb. nov. Basionym:
Habranthus andalgalensis Ravenna, Sellowia 19: 30.
1967.
73. Amaryllidaceae. References: Meerow et al. (2000),
Fay et al. (2006). Hippeastrum araguaiense (Ravenna)
Christenh. & Byng, comb. nov. Basionym:
Note: Gethyum Phil. and Solaria Phil. are embedded in
Habranthus araguaiensis Ravenna, Onira 6: 41.
Gilliesia Lindl. and are hereby transferred. Similarly
2001.
Speea Loes. is placed in Miersia Lindl. Additionally,
Eucharis Planch. & Linden and Caliphruria Herb. are Hippeastrum arenicola (Brandegee) Christenh. &
placed in Urceolina Rchb., where they more naturally Byng, comb. nov. Basionym: Zephyranthes arenicola
reside and Hippeastrum Herb. is treated in the broad Brandegee, Proc. Calif. Acad. Sci., ser. 2, 2: 205. 1889.
sense, including the entire clade, encompassing Hippeastrum argyrium Christenh. & Byng, nom. nov.
Habranthus Herb., Rhodophiala C.Presl, Sprekelia Replaced synonym: Habranthus argentinus Traub,
Heist. and Zephyranthes Herb., among others, to Pl. Life 7: 42. 1951, not: Hippeastrum argentinum
make the genus monophyletic. (Pax) Hunz. Etymology: From Greek αργυρεια, silver.
Gilliesia atropurpurea (Phil.) M.F.Fay & Christenh., Hippeastrum arzae (Phil.) Christenh. & Byng, comb.
comb. nov. Basionym: Gethyum atropurpureum nov. Basionym: Placea arzae Phil., Anales Univ. Chile
Phil., Anales Univ. Chile 1873: 549. 1873. (Figure 8B) 43: 541. 1873.
Gilliesia attenuata (Ravenna) M.F.Fay & Christenh., Hippeastrum atamasco (L.) Christenh. & Byng, comb.
comb. nov. Basionym: Solaria attenuata Ravenna, nov. Basionym: Amaryllis atamasco L., Sp. Pl.: 292.
Bol. Soc. Argent. Bot. 11: 160. 1967. 1753. (Figure 8C)
Gilliesia brevicoalita (Ravenna) M.F.Fay & Christenh., Hippeastrum auratum (Ravenna) Christenh. & Byng,
comb. nov. Basionym: Solaria brevicoalita Ravenna, comb. nov. Basionym: Habranthus auratus Ravenna,
Onira 9: 64. 2005. Onira 9: 15. 2003.
Gilliesia curacavina (Ravenna) M.F.Fay & Christenh., Hippeastrum australe (Ravenna) Christenh. & Byng,
comb. nov. Basionym: Solaria curacavina Ravenna, comb. nov. Basionym: Phycella australis Ravenna,
Onira 9: 65. 2005. Pl. Life 37: 71. 1981.
Gilliesia cuspidata (Harv. ex Baker) M.F.Fay & Hippeastrum barrosianum (Hunz. & Di Fulvio)
Christenh., comb. nov. Basionym: Ancrumia Christenh. & Byng, comb. nov. Basionym:
cuspidata Harv. ex Baker, Hooker’s Icon. Pl. 13: t. Habranthus barrosianus Hunz. & Di Fulvio, Kurtziana
1227. 1877. 7: 255. 1973.
Gilliesia miersioides (Phil.) M.F.Fay & Christenh., comb. Hippeastrum bellum (T.M.Howard & S.Ogden)
nov. Basionym: Solaria miersioides Phil., Linnaea 29: Christenh. & Byng, comb. nov. Basionym:
72. 1858. Zephyranthes bella T.M.Howard & S.Ogden,
Hippeastrum albiellum (Traub) Christenh. & Byng, Herbertia 46: 105. 1991.
comb. nov. Basionym: Zephyranthes albiella Traub, Hippeastrum berteroi (Spreng.) Christenh. & Byng,
Pl. Life 6: 51. 1950. comb. nov. Basionym: Amaryllis berteroi Spreng.,
Hippeastrum albispiritum (Ravenna) Syst. Veg., ed. 16, 2: 49. 1825. Replaced synonym:
Christenh. & Byng, comb. nov. Basionym: Habranthus robustus Herb. in Sweet, Brit. Fl. Gard.
4: t. 14 (1829), not: Hippeastrum robustum (Otto &

A.Dietr.) A.Dietr. & Schult.f. in J.J.Roemer & J.A.Schultes, Syst.

Hippeastrum biflorum (Phil.) Christenh. & Byng, comb. Veg., ed. 15 bis 7: 799. 1830. Replaced synonym:
nov. Basionym: Rhodophiala biflora Phil., Linnaea Zephyranthes rosea Lindl., Bot. Reg. 10: t. 821. 1824,
29: 66. 1858. not: Hippeastrum roseum Baker
Hippeastrum bifolium (Aubl.) Christenh. & Byng, Hippeastrum catamarcense (Ravenna)
comb. nov. Basionym: Amaryllis bifolia Aubl., Hist. Christenh. & Byng, comb. nov. Basionym:
Pl. Guiane 1: 304. 1775. Habranthus catamarcensis Ravenna, Pl. Life 30: 43.
1974.
Hippeastrum botumirimense (R.S.Oliveira)
Christenh. & Byng, comb. nov. Basionym: Hippeastrum chacoense (Ravenna)
Habranthus botumirimensis R.S.Oliveira, Kew Bull. Christenh. & Byng, comb. nov. Basionym:
64: 538. 2009. Habranthus chacoensis Ravenna, Not. Mens. Mus.
Nac. Hist. Nat. 15(173): 4. 1970.
Hippeastrum brevipes (Standl.) Christenh. & Byng,
comb. nov. Basionym: Zephyranthes brevipes Hippeastrum chichimeca (T.M.Howard & S.Ogden)
Standl., Publ. Field Columb. Mus., Bot. Ser. 4: 199. Christenh. & Byng, comb. nov. Basionym:
1929. Zephyranthes chichimeca T.M.Howard & S.Ogden,
Herbertia 46: 105. 1991.
Hippeastrum breviscapum (Ravenna) Christenh.
& Byng, comb. nov. Basionym: Zephyranthes Hippeastrum chlorosolen (Herb.) Christenh. & Byng,
breviscapa Ravenna, Onira 3: 54. 1999. comb. nov. Basionym: Cooperia chlorosolen Herb.,
Edwards’s Bot. Reg. 22: t. 1835. 1836.
Hippeastrum brevitubum (Herb.) Christenh. & Byng,
comb. nov. Basionym: Phycella brevituba Herb., Hippeastrum chrysanthum (Greenm. & C.H.Thomps.)
Amaryllidaceae: 154. 1837. Christenh. & Byng, comb. nov. Basionym:
Zephyranthes chrysantha Greenm. & C.H.Thomps.,
Hippeastrum briquetii (J.F.Macbr.) Christenh. & Byng,
Ann. Missouri Bot. Gard. 1: 406. 1914.
comb. nov. Basionym: Zephyranthes briquetii
J.F.Macbr., Publ. Field Columb. Mus., Bot. Ser. 11: 9. Hippeastrum ciceroanum (M.M.Mejía & R.G.García)
1931. Christenh. & Byng, comb. nov. Basionym:
Zephyranthes ciceroana M.M.Mejía & R.G.García,
Hippeastrum caaguazuense (Ravenna)
Moscosoa 8: 23. 1994.
Habranthus caaguazuensis Ravenna, Onira 9: 15. Hippeastrum citrinum (Baker) Christenh. & Byng,
2003. comb. nov. Basionym: Zephyranthes citrina Baker,
Bot. Mag. 108: t. 6605. 1882.
Hippeastrum caeruleum (Griseb.) Christenh. & Byng,
comb. nov. Basionym: Amaryllis caerulea Griseb., Hippeastrum clintiae (Traub) Christenh. & Byng, comb.
Abh. Königl. Ges. Wiss. Göttingen 24: 320. 1879. nov. Basionym: Zephyranthes clintiae Traub, Pl. Life
8: 76. 1952.
Hippeastrum calderense (Ravenna)
Christenh. & Byng, comb. nov. Basionym: Hippeastrum colchicum Christenh. & Byng, nom. nov.
Habranthus calderensis Ravenna, Onira 9: 13. 2003. Replaced synonym: Zephyranthes fosteri Traub,
Herbertia 7: 65. 1941, not: Hippeastrum forsteri
Hippeastrum capivarinum (Ravenna) Christenh.
(Traub) H.E.Moore. Etymology: Named for its
& Byng, comb. nov. Basionym: Zephyranthes
resemblance to a member of the genus Colchicum
capivarina Ravenna, Pl. Life 30: 35. 1974.
(Colchicaceae).
Hippeastrum cardenasianum (Traub & I.S.Nelson)
Hippeastrum concinnum (Ravenna)
Habranthus cardenasianus Traub & I.S.Nelson, Pl.
Habranthus concinnus Ravenna, Onira 3: 63. 1999.
Life 13: 69. 1957.
Hippeastrum conzattii (Greenm.) Christenh. & Byng,
Hippeastrum cardinale (C.H.Wright) Christenh. &
comb. nov. Basionym: Zephyranthes conzattii
Byng, comb. nov. Basionym: Zephyranthes cardinalis
Greenm., Proc. Amer. Acad. Arts 33: 473. 1898.
C.H.Wright, Bot. Mag. 140: t. 8553. 1914.
Hippeastrum cordobense (Ravenna)
Hippeastrum carinatum (Herb.) Christenh. & Byng,
comb. nov. Basionym: Zephyranthes carinata Herb.,
Habranthus cordobensis Ravenna, Onira 6: 42. 2001.
Bot. Mag. 52: t. 2594. 1825.
Hippeastrum correntinum (Roitman, J.A.Castillo
Hippeastrum carmineum (Ravenna)
& M.R.Barrios) Christenh. & Byng, comb. nov.
Basionym: Habranthus correntinus Roitman,
Habranthus carmineus Ravenna, Not. Mens. Mus.
J.A.Castillo & M.R.Barrios, Bol. Soc. Argent. Bot. 43:
Nac. Hist. Nat. 15: 5. 1970.
153. 2008.
Hippeastrum carneum (Schult.) Christenh. & Byng,
Hippeastrum crassibulbum (Ravenna)
comb. nov. Basionym: Amaryllis carnea Schult.

59
Christenh. & Byng, comb. nov. Basionym: Not. Mens. Mus. Nac. Hist. Nat. 15(173): 4. 1970.
Habranthus crassibulbus Ravenna, Onira 3: 63. 1999. Hippeastrum erubescens (S.Watson) Christenh.
Hippeastrum crastinum Christenh. & Byng, nom. nov. & Byng, comb. nov. Basionym: Zephyranthes
Replaced synonym: Habranthus vittatus T.M.Howard, erubescens S.Watson, Proc. Amer. Acad. Arts 25:
Herbertia 46: 115. 1991, not: Hippeastrum vittatum 162. 1890.
(L.Hér.) Herb. Etymology: The common name of this Hippeastrum estense (Ravenna) Christenh. & Byng,
species is mañanita or tomorrow flower, hence the comb. nov. Basionym: Habranthus estensis Ravenna,
epithet meaning tomorrow in Latin. Pl. Life 30: 46. 1974.
Hippeastrum crinum Christenh. & Byng, nom. nov. Hippeastrum filifolium (Herb. ex Kraenzl.) Christenh.
Replaced synonym: Habranthus spectabilis Ravenna, & Byng, comb. nov. Basionym: Zephyranthes filifolia
Not. Mens. Mus. Nac. Hist. Nat. 15(173): 6. 1970, Herb. ex Kraenzl., Bull. Misc. Inform. Kew 1913: 190.
not: Hippeastrum spectabile M.Roem. Etymology: 1913.
Named for its resemblance to a member of the Hippeastrum flavissimum (Ravenna) Christenh. &
genus Crinum (Amaryllidaceae). Byng, comb. nov. Basionym: Zephyranthes flavissima
Hippeastrum crocus Christenh. & Byng, nom. nov. Ravenna, Sellowia 19: 28. 1967.
Replaced synonym: Zephyranthes crociflora Hippeastrum flavum (Phil.) Christenh. & Byng, comb.
T.M.Howard & S.Ogden, Herbertia 46: 106. 1991, nov. Basionym: Habranthus flavus Phil., Anales Univ.
not: Hippeastrum crociflorum Rusby. Etymology: Chile 27: 333. 1865.
Named for its resemblance to a Crocus (Iridaceae).
Hippeastrum fluviale (Ravenna) Christenh. & Byng,
Hippeastrum cubense (Urb.) Christenh. & Byng, comb. comb. nov. Basionym: Zephyranthes fluvialis
nov. Basionym: Zephyranthes cubensis Urb., Symb. Ravenna, Onira 6: 39. 2001.
Antill. 5: 292. 1907.
Hippeastrum fulgens (Hook.f.) Christenh. & Byng,
Hippeastrum cyrtanthoides (Sims) Christenh. & Byng, comb. nov. Basionym: Habranthus fulgens Hook.f.,
comb. nov. Basionym: Amaryllis cyrtanthoides Sims, Bot. Mag. 92: t. 5563. 1866.
Bot. Mag. 50: t. 2399. 1823.
Hippeastrum gameleirense (Ravenna)
Hippeastrum datense (Ravenna) Christenh. & Byng, Christenh. & Byng, comb. nov. Basionym:
comb. nov. Basionym: Habranthus datensis Ravenna, Habranthus gameleirensis Ravenna, Onira 3: 57.
Onira 3: 58. 1999. 1999.
Hippeastrum davidii (Ravenna) Christenh. & Byng, Hippeastrum germainii (Phil.) Christenh. & Byng,
comb. nov. Basionym: Placea davidii Ravenna, Pl. comb. nov. Basionym: Placea germainii Phil.,
Life 37: 73. 1981. Linnaea 29: 67. 1858.
Hippeastrum depauperatum (Herb.) Christenh. Hippeastrum gilliesianum (Herb.) Christenh. & Byng,
& Byng, comb. nov. Basionym: Zephyranthes comb. nov. Basionym: Habranthus bagnoldii var.
depauperata Herb., Amaryllidaceae: 171. 1837. gilliesianus Herb., Amaryllidaceae: 163. 1837.
Hippeastrum dichromanthum (T.M.Howard) Christenh. Hippeastrum glaucum (Lindl.) Christenh. & Byng,
& Byng, comb. nov. Basionym: Zephyranthes comb. nov. Basionym: Sprekelia glauca Lindl.,
dichromantha T.M.Howard, Herbertia 51: 38. 1996. Edwards’s Bot. Reg. 26(Misc.): 49. 1840.
Hippeastrum dilutum (Ravenna) Christenh. & Byng, Hippeastrum gracile (Herb.) Christenh. & Byng,
comb. nov. Basionym: Zephyranthes diluta Ravenna, comb. nov. Basionym: Zephyranthes gracilis Herb.,
Onira 3: 55. 1999. Amaryllidaceae: 172. 1837.
Hippeastrum drummondii (D.Don in R.Sweet) Hippeastrum gratissimum (Ravenna) Christenh.
Christenh. & Byng, comb. nov. Basionym: & Byng, comb. nov. Basionym: Zephyranthes
Zephyranthes drummondii D.Don in R.Sweet, Brit. Fl. gratissima Ravenna, Onira 6: 38. 2001.
Gard. 7: t. 328. 1836.
Hippeastrum guachipense (Ravenna)
Hippeastrum duarteanum (Ravenna) Christenh. & Byng, comb. nov. Basionym:
Christenh. & Byng, comb. nov. Basionym: Habranthus guachipensis Ravenna, Pl. Life 30: 43.
Habranthus duarteanus Ravenna, Pl. Life 30: 45. 1974.
1974.
Hippeastrum guaraniorum Christenh. & Byng, nom.
Hippeastrum dutilhianum (Büneker, R.E.Bastian nov. Replaced synonym: Zephyranthes paranaensis
& C.M.Costa) Christenh. & Byng, comb. nov. Ravenna, Pl. Life 30: 37. 1974, not: Hippeastrum
Basionym: Tocantinia dutilhiana Büneker, R.E.Bastian paranaense (Traub) Meerow. Etymology: Named for
& C.M.Costa, Balduinia 53: 6. 2016. the Guarani tribe of the Paraná region where this
Hippeastrum erectum (Ravenna) Christenh. & Byng, species occurs.
comb. nov. Basionym: Habranthus erectus Ravenna, Hippeastrum guatemalense (L.B.Spencer) Christenh.

& Byng, comb. nov. Basionym: Zephyranthes Byng, comb. nov. Basionym: Zephyranthes lagesiana
guatemalensis L.B.Spencer, Phytologia 59: 85. 1986. Ravenna, Onira 6: 40. 2001.
Hippeastrum habranthum Christenh. & Byng, nom. nov. Hippeastrum latissimifolium (L.B.Spencer) Christenh.
Replaced synonym:: Habranthus goianus Ravenna, & Byng, comb. nov. Basionym: Zephyranthes
Pl. Life 30: 45. 1974, not: Hippeastrum goianum latissimifolia L.B.Spencer, Phytologia 59: 86. 1986.
(Ravenna) Meerow. Etymology: Named for the Hippeastrum leonense (Ravenna)
genus Habranthus, now a synonym of Hippeastrum. Christenh. & Byng, comb. nov. Basionym:
Hippeastrum hermokrokos Christenh. & Byng, nom. Habranthus leonensis Ravenna, Onira 3: 56. 1999.
nov. Replaced synonym: Zephyranthes amoena Hippeastrum leptandrum (Ravenna)
Ravenna, Onira 3: 54. 1999, not: Hippeastrum Christenh. & Byng, comb. nov. Basionym:
amoenum (Phil.) Christenh. & Byng. Etymology: Habranthus leptandrus Ravenna, Pl. Life 34: 87.
Named for the mortal youth Κρόκος, companion of 1978.
Ἑρμῆς. Krokos was accidentally killed and in distress Hippeastrum leucanthum (T.M.Howard) Christenh.
Hermes turned his body into a flower in Greek & Byng, comb. nov. Basionym: Zephyranthes
mythology. leucantha T.M.Howard, Herbertia 49: 94. 1994.
Hippeastrum hondurense (Ravenna) Christenh. Hippeastrum lilium Christenh. & Byng, nom. nov.
& Byng, comb. nov. Basionym: Zephyranthes Replaced synonym: Zephyranthes elegans Ravenna,
hondurensis Ravenna, Onira 3: 54. 1999. Onira 3: 53. 1999, not: Hippeastrum elegans
Hippeastrum howardii (Traub) Christenh. & Byng, (Spreng.) H.E.Moore. Etymology: The species
comb. nov. Basionym: Zephyranthes howardii Traub, resembles a lily (Lilium, Liliaceae) hence the epithet.
Pl. Life 19: 49. 1963. Hippeastrum lindleyanum (Herb.) Christenh. & Byng,
Hippeastrum immaculatum (Traub & Clint) comb. nov. Basionym: Zephyranthes lindleyana
Christenh. & Byng, comb. nov. Basionym: Herb., Amaryllidaceae: 174. 1837.
Habranthus immaculatus Traub & Clint, Pl. Life 13: Hippeastrum longifolium (Hemsl.) Christenh. & Byng,
68. 1957. comb. nov. Basionym: Zephyranthes longifolia
Hippeastrum insularum (H.H.Hume ex Moldenke) Hemsl., Diagn. Pl. Nov. Mexic.: 55. 1880.
Christenh. & Byng, comb. nov. Basionym: Hippeastrum longipes (Baker) Christenh. & Byng,
Zephyranthes insularum H.H.Hume ex Moldenke, Pl. comb. nov. Basionym: Zephyranthes longipes Baker,
Life 8: 152. 1952. Bull. Misc. Inform. Kew 1898: 225. 1898.
Hippeastrum irwinianum (Ravenna) Hippeastrum longistylum (Pax) Christenh. & Byng,
Christenh. & Byng, comb. nov. Basionym: comb. nov. Basionym: Zephyranthes longistyla Pax,
Habranthus irwinianus Ravenna, Pl. Life 26: 97. Bot. Jahrb. Syst. 11: 320. 1889.
1970.
Hippeastrum longitubum (Flory ex Flagg & G.Lom.
Hippeastrum ischihualastum (Ravenna) Sm.) Christenh. & Byng, comb. nov. Basionym:
Christenh. & Byng, comb. nov. Basionym: Zephyranthes longituba Flory ex Flagg & G.Lom.Sm.,
Habranthus ischihualastus Ravenna, Onira 3: 65. Herbertia 64: 120. 2010.
1999.
Hippeastrum lucidum (R.S.Oliveira)
Hippeastrum itaobinum (Ravenna) Christenh. & Byng, comb. nov. Basionym:
Christenh. & Byng, comb. nov. Basionym: Habranthus lucidus R.S.Oliveira, Kew Bull. 64: 538.
Habranthus itaobinus Ravenna, Onira 3: 56. 1999. 2009.
Hippeastrum jonesii (Cory) Christenh. & Byng, comb. Hippeastrum luteum (Phil.) Christenh. & Byng, comb.
nov. Basionym: Cooperia jonesii Cory, Field & Lab. nov. Basionym: Placea lutea Phil., Linnaea 33: 259.
18: 45. 1950. 1865.
Hippeastrum jujuyense (E.Holmb.) Christenh. & Byng, Hippeastrum maasii (Ravenna) Christenh. & Byng,
comb. nov. Basionym: Zephyranthes jujuyensis comb. nov. Basionym: Habranthus maasii Ravenna,
E.Holmb., Anales Mus. Nac. Buenos Aires, ser. 3, 4: Pl. Life 34: 85. 1978.
523. 1905.
Hippeastrum macrosiphon (Baker) Christenh. & Byng,
Hippeastrum katheriniae (L.B.Spencer) Christenh. comb. nov. Basionym: Zephyranthes macrosiphon
& Byng, comb. nov. Basionym: Zephyranthes Baker, Gard. Chron., n.s., 16: 70. 1881.
katheriniae L.B.Spencer, Phytologia 59: 86. 1986.
Hippeastrum maculatum (L’Hér.) Christenh. & Byng,
Hippeastrum lacteum (S.Moore) Christenh. & Byng, comb. nov. Basionym: Amaryllis maculata L’Hér.,
comb. nov. Basionym: Zephyranthes lactea S.Moore, Sert. Angl.: 10. 1789.
Trans. Linn. Soc. London, Bot. 4: 495. 1895.
Hippeastrum magnoi (Ravenna) Christenh. & Byng,
Hippeastrum lagesianum (Ravenna) Christenh. & comb. nov. Basionym: Habranthus magnoi Ravenna,

61
Sellowia 19: 31. 1967. moctezumae T.M.Howard, Herbertia 51: 40. 1996.
Hippeastrum mannii (Roitman, J.A.Castillo & Hippeastrum modestum (Ravenna) Christenh. &
Maza) Christenh. & Byng, comb. nov. Basionym: Byng, comb. nov. Basionym: Zephyranthes modesta
Habranthus neumannii Roitman, J.A.Castillo & Maza, Ravenna, Onira 3: 62. 1999, not: Hippeastrum
Darwiniana 46: 67. 2008. modestrum (Phil.) Baker.
Hippeastrum martinezii (Ravenna) Hippeastrum morrisclintii (Traub & T.M.Howard)
Christenh. & Byng, comb. nov. Basionym: Christenh. & Byng, comb. nov. Basionym:
Habranthus martinezii Ravenna, Not. Mens. Mus. Zephyranthes morrisclintii Traub & T.M.Howard, Pl.
Nac. Hist. Nat. 15(173): 3. 1970. Life 26: 62. 1970.
Hippeastrum matacum (Ravenna) Christenh. & Byng, Hippeastrum nervosum (Herb.) Christenh. & Byng,
comb. nov. Basionym: Habranthus matacus Ravenna, comb. nov. Basionym: Zephyranthes nervosa Herb.,
Onira 3: 66. 1999. Amaryllidaceae: 172. 1837.
Hippeastrum medinae (L.O.Alvarado & García- Hippeastrum niveum (Ravenna) Christenh. & Byng,
Mend.) Christenh. & Byng, comb. nov. Basionym: comb. nov. Basionym: Habranthus niveus Ravenna,
Habranthus medinae L.O.Alvarado & García-Mend., Pl. Life 26: 95. 1970.
Novon 18: 283. 2008. Hippeastrum nymphaea (T.M.Howard & S.Ogden)
Hippeastrum mendocense (Baker) Christenh. & Byng, Christenh. & Byng, comb. nov. Basionym:
comb. nov. Basionym: Zephyranthes mendocensis Zephyranthes nymphaea T.M.Howard & S.Ogden,
Baker, Handb. Amaryll.: 36. 1888. Herbertia 46: 107. 1991.
Hippeastrum mesochloum (Herb. ex Lindl.) Christenh. Hippeastrum oaxacanum (T.M.Howard)
& Byng, comb. nov. Basionym: Zephyranthes Christenh. & Byng, comb. nov. Basionym:
mesochloa Herb. ex Lindl., Edwards’s Bot. Reg. 16: Habranthus oaxacanus T.M.Howard, Herbertia 51:
t. 1345. 1830. 43. 1996.
Hippeastrum mexicanum (T.M.Howard) Hippeastrum oltanum (Ravenna) Christenh. & Byng,
Christenh. & Byng, comb. nov. Basionym: comb. nov. Basionym: Habranthus oltanus Ravenna,
Habranthus mexicanus T.M.Howard, Herbertia 51: Onira 6: 41. 2001.
42. 1996. Hippeastrum oranense (Ravenna)
Hippeastrum microcarpum (Rusby) Christenh. & Byng, Christenh. & Byng, comb. nov. Basionym:
comb. nov. Basionym: Atamosco microcarpa Rusby, Habranthus oranensis Ravenna, Onira 3: 64. 1999.
Mem. New York Bot. Gard. 7: 213. 1927. Hippeastrum orellanae (Carnevali, Duno & J.L.Tapia)
Hippeastrum microstigmum (Ravenna) Christenh. Christenh. & Byng, comb. nov. Basionym:
& Byng, comb. nov. Basionym: Zephyranthes Zephyranthes orellanae Carnevali, Duno & J.L.Tapia,
microstigma Ravenna, Onira 10: 8. 2005. J. Torrey Bot. Soc. 137: 45. 2010.
Hippeastrum millarense (Ravenna) Hippeastrum ornatum (Miers) Christenh. & Byng,
Christenh. & Byng, comb. nov. Basionym: comb. nov. Basionym: Placea ornata Miers,
Habranthus millarensis Ravenna, Pl. Life 37: 66. Edwards’s Bot. Reg. 27: t. 50. 1841.
1981. Hippeastrum pantanalense (Ravenna)
Hippeastrum minimum (Herb.) Christenh. & Byng, Christenh. & Byng, comb. nov. Basionym:
comb. nov. Basionym: Zephyranthes minima Herb., Habranthus pantanalensis Ravenna, Onira 3: 59.
Amaryllidaceae: 172. 1837. 1999.
Hippeastrum minor (Ravenna) Christenh. & Byng, Hippeastrum philadelphicum (Ravenna)
comb. nov. Basionym: Habranthus minor Ravenna, Christenh. & Byng, comb. nov. Basionym:
Onira 9: 13. 2003. Habranthus philadelphicus Ravenna, Onira 9: 16.
Hippeastrum minutum (Kunth) Christenh. & Byng, 2003.
comb. nov. Basionym: Amaryllis minuta Kunth, Nov. Hippeastrum pictum (Ravenna) Christenh. & Byng,
Gen. Sp. 1: 278. 1816. comb. nov. Basionym: Habranthus pictus Ravenna,
Hippeastrum miradorense (Kraenzl.) Christenh. & Onira 3: 59. 1999.
Byng, comb. nov. Basionym: Cooperia miradorensis Hippeastrum plumieri (H.H.Hume ex Moldenke)
Kraenzl., Repert. Spec. Nov. Regni Veg. 21: 75. 1925. Christenh. & Byng, comb. nov. Basionym:
Hippeastrum mirum (Ravenna) Christenh. & Byng, Zephyranthes plumieri H.H.Hume ex Moldenke, Pl.
comb. nov. Basionym: Tocantinia mira Ravenna, Life 8: 152. 1952.
Onira 5: 10. 2000. Hippeastrum primulinum (T.M.Howard & S.Ogden)
Hippeastrum moctezumae (T.M.Howard) Christenh. Christenh. & Byng, comb. nov. Basionym:
& Byng, comb. nov. Basionym: Zephyranthes Zephyranthes primulina T.M.Howard & S.Ogden,

Herbertia 46: 110. 1991. Not. Mens. Mus. Nac. Hist. Nat. 15(173): 6. 1970.
Hippeastrum pseudocrocus (Solms) Christenh. & Byng, Hippeastrum sanavirone (Roitman, J.A.Castillo,
comb. nov. Basionym: Haylockia pseudocrocus G.M.Tourn & Uria) Christenh. & Byng, comb.
Solms, Bot. Zeitung (Berlin) 65: 135. 1907. Replaced nov. Basionym: Habranthus sanavirone Roitman,
synonym: Haylockia andina R.E.Fr., Nova Acta J.A.Castillo, G.M.Tourn & Uria, Novon 17: 393. 2007.
Regiae Soc. Sci. Upsal., ser. 4, 1: 160. 1905, not: Hippeastrum scarlatinum (Ravenna) Christenh. &
Hippeastrum andinum Phil. Byng, comb. nov. Basionym: Phycella scarlatina
Hippeastrum proctorii (Acev.-Rodr. & M.T.Strong) Ravenna, Pl. Life 37: 69. 1981.
Christenh. & Byng, comb. nov. Basionym: Hippeastrum schulzianum (Ravenna)
Zephyranthes proctorii Acev.-Rodr. & M.T.Strong, Christenh. & Byng, comb. nov. Basionym:
Contr. U.S. Natl. Herb. 52: 115. 2005. Habranthus schulzianus Ravenna, Not. Mens. Mus.
Hippeastrum puertoricense (Traub) Christenh. & Byng, Nac. Hist. Nat. 16: 8. 1972.
comb. nov. Basionym: Zephyranthes puertoricensis Hippeastrum sessile (Herb.) Christenh. & Byng,
Traub, Pl. Life 7: 37. 1951. comb. nov. Basionym: Zephyranthes sessilis Herb.,
Hippeastrum pulchellum (J.G.Sm.) Christenh. & Byng, Amaryllidaceae: 175. 1837.
comb. nov. Basionym: Zephyranthes pulchella Hippeastrum simpsonii (Chapm.) Christenh. & Byng,
J.G.Sm., Rep. (Annual) Missouri Bot. Gard. 6: 114. comb. nov. Basionym: Zephyranthes simpsonii
1895. Chapm., Fl. South. U.S., ed. 2(Suppl. 2): 696. 1892.
Hippeastrum purpurellum (Ravenna) Christenh. Hippeastrum smallii (Alexander) Christenh. & Byng,
& Byng, comb. nov. Basionym: Zephyranthes comb. nov. Basionym: Cooperia smallii Alexander,
purpurella Ravenna, Onira 8: 41. 2003. Addisonia 21: 7. 1939.
Hippeastrum quilmesianum (Ravenna) Hippeastrum smilakrokos Christenh. & Byng, nom.
Christenh. & Byng, comb. nov. Basionym: nov. Replaced synonym: Cooperia traubii W.Hayw.,
Habranthus quilmesianus Ravenna, Onira 1: 54. Herbertia 3: 64. 1936, not: Hippeastrum traubii
1988. (Moldenke) H.E.Moore. Etymology: Named for the
Hippeastrum refugiense (F.B.Jones) Christenh. & Byng, mortal youth Κρόκος, who was unhappy with his
comb. nov. Basionym: Zephyranthes refugiensis love affair with the nymph Σμίλαξ and both were
F.B.Jones, Rhodora 63: 214. 1961. turned into plants, one a saffron crocus, the other a
Hippeastrum riojanum (Ravenna) Christenh. & Byng, prickly bindweed in Greek mythology.
comb. nov. Basionym: Habranthus riojanus Ravenna, Hippeastrum sprekeliopsis Christenh. & Byng, nom. nov.
Pl. Life 26: 96. 1970. Replaced synonym: Sprekelia howardii Lehmiller,
Hippeastrum rosalense (Ravenna) Christenh. & Byng, Herbertia 54: 230. 2000., not: Hippeastrum howardii
comb. nov. Basionym: Zephyranthes rosalensis (Traub) Christenh. & Byng. Etymology: Named for
Ravenna, Pl. Life 30: 34. 1974. Sprekelia, a genus now included in Hippeastrum.
Hippeastrum royale Christenh. & Byng, nom. Hippeastrum stellare (Ravenna) Christenh. & Byng,
nov. Replaced synonym: Zephyranthes reginae comb. nov. Basionym: Zephyranthes stellaris
T.M.Howard & S.Ogden, Herbertia 46: 111. 1991, Ravenna, Sellowia 19: 28. 1967.
Not: Hippeastrum reginae (L.) Herb. Etymology: This Hippeastrum stellatoroseum (G.Lom.Sm., Spurrier,
regal species was renamed to remain royal. Flagg & Espejo) Christenh. & Byng, comb. nov.
Hippeastrum rubrum (Ravenna) Christenh. & Byng, Basionym: Zephyranthes stellatorosea G.Lom.Sm.,
comb. nov. Basionym: Habranthus ruber Ravenna, Spurrier, Flagg & Espejo, Novon 24: 289. 2015.
Pl. Life 26: 94. 1970. Hippeastrum steyermarkii (Ravenna)
Hippeastrum ruizlealii (Ravenna) Christenh. & Byng, Christenh. & Byng, comb. nov. Basionym:
comb. nov. Basionym: Habranthus ruizlealii Ravenna, Habranthus steyermarkii Ravenna, Pl. Life 34: 85.
Pl. Life 30: 44. 1974. 1978.
Hippeastrum saipinense (Ravenna) Hippeastrum stigmovittatum (Büneker, R.E.Bastian
Christenh. & Byng, comb. nov. Basionym: & C.M.Costa) Christenh. & Byng, comb. nov.
Habranthus saipinensis Ravenna, Onira 10: 10. 2005. Basionym: Tocantinia stigmovittata Büneker,
R.E.Bastian & C.M.Costa, Balduinia 53: 8. 2016.
Hippeastrum salinarum (Ravenna)
Christenh. & Byng, comb. nov. Basionym: Hippeastrum subflavum (L.B.Spencer) Christenh. &
Habranthus salinarum Ravenna, Not. Mens. Mus. Byng, comb. nov. Basionym: Zephyranthes subflava
Nac. Hist. Nat. 15(173): 3. 1970. L.B.Spencer, Phytologia 59: 87. 1986.
Hippeastrum saltense (Ravenna) Christenh. & Byng, Hippeastrum susatanum (Fern.Alonso & Groenend.)
comb. nov. Basionym: Habranthus saltensis Ravenna, Christenh. & Byng, comb. nov. Basionym:

63
Zephyranthes susatana Fern.Alonso & Groenend., Linden ex Planch., J. Gén. Hort. 12: 1216. 1857.
Revista Acad. Colomb. Ci. Exact. 28: 179. 2004. Urceolina caucana (Meerow) Christenh. & Byng,
Hippeastrum tepicense (Greenm. ex Flagg & G.Lom. comb. nov. Basionym: Eucharis caucana Meerow,
Sm.) Christenh. & Byng, comb. nov. Basionym: Ann. Missouri Bot. Gard. 76: 191. 1989.
Habranthus tepicensis Greenm. ex Flagg & G.Lom. Urceolina cyaneosperma (Meerow) Christenh. &
Sm., Herbertia 63: 175. 2009. Byng, comb. nov. Basionym: Eucharis cyaneosperma
Hippeastrum tiltilense (Traub & Moldenke) Christenh. Meerow, Sida 12: 39. 1987.
& Byng, comb. nov. Basionym: Amaryllis tiltilensis Urceolina formosa (Meerow) Christenh. & Byng,
Traub & Moldenke, Amaryllis Manual: 88. 1949. comb. nov. Basionym: Eucharis formosa Meerow,
Replaced synonym: Hippeastrum laetum Phil., Sida 12: 42. 1987.
Anales Univ. Chile 93: 157. 1890, not: Hippeastrum Urceolina moorei (Baker) Christenh. & Byng, comb.
laetum Phil. nov. Basionym: Eucharis grandiflora var. moorei
Hippeastrum treatiae (S.Watson) Christenh. & Byng, Baker, Gard. Chron., ser. 3, 4: 628. 1888.
comb. nov. Basionym: Zephyranthes treatiae Urceolina plicata (Meerow) Christenh. & Byng, comb.
S.Watson, Proc. Amer. Acad. Arts 14: 300. 1879. nov. Basionym: Eucharis plicata Meerow, Brittonia
Hippeastrum tucumanense (Hunz.) Christenh. & Byng, 36: 18. 1984.
comb. nov. Basionym: Zephyranthes tucumanensis Urceolina ruthiana (L.Jost, Oleas & Meerow)
Hunz., Kurtziana 5: 362. 1969. Christenh. & Byng, comb. nov. Basionym: Eucharis
Hippeastrum tulipanum Christenh. & Byng, nom. ruthiana L.Jost, Oleas & Meerow, PhytoKeys 48: 6.
nov. Replaced synonym: Zephyranthes nelsonii 2015.
Greenm., Proc. Amer. Acad. Arts 33: 473. 1898, not:
Hippeastrum nelsonii (Cárdenas) Van Scheepen.
74. Asparagaceae. References: Bos (1998), Rudall et
Etymology: Named for its superficial resemblance to
al. (2000), Jankalski (2008), Lu & Morden (2014),
a species of Tulipa (Liliaceae).
Knirsch et al. (2016), Mabberley (2017), WCSP
Hippeastrum uruguaianicum (Ravenna) Christenh. (2018).
& Byng, comb. nov. Basionym: Zephyranthes
Note: As traditionally circumscribed Dracaena L. is
uruguaianica Ravenna, Onira 10: 9. 2005.
paraphyletic with Sansevieria Thunb. and part of
Hippeastrum vandenbergianum Christenh. & Byng, Pleomele Salisb. embedded in it. As these genera
nom. nov. Replaced synonym: Habranthus bahiensis are morphologically variable, but share substantial
Ravenna, Onira 1: 53. 1988, not: Hippeastrum similarities, it is best treated as a single genus.
bahiense M.Roem. Etymology: Named for Cassio van Several species were already transferred into
den Berg, for his contributions to Brazilian botany. Dracaena and the genus was officially synonymised
Hippeastrum venturiana (Ravenna) by Mabberley (2017), but not all combinations were
Christenh. & Byng, comb. nov. Basionym: made. We hereby transfer the remaining taxa to
Habranthus venturianus Ravenna, Onira 3: 65. 1999. standardise the taxonomy.
Hippeastrum wrightii (Baker) Christenh. & Byng, Dracaena aethiopica (Thunb.) Byng & Christenh.,
comb. nov. Basionym: Zephyranthes wrightii Baker, comb. nov. Basionym: Sansevieria aethiopica
Handb. Amaryll.: 32. 1888. Thunb., Prodr. Pl. Cap.: 65. 1794.
Hippeastrum yaviense (Ravenna) Christenh. & Byng, Dracaena angolensis (Welw. ex Carrière) Byng &
comb. nov. Basionym: Zephyranthes yaviensis Christenh., comb. nov. Basionym: Sansevieria
Ravenna, Onira 1: 55. 1988. angolensis Welw. ex Carrière, Rev. Hort. (Paris) 1861:
Hippeastrum zephyranthum Christenh. & Byng, nom. 447. 1861. Replaced synonym: Sansevieria cylindrica
nov. Replaced synonym: Amaryllis candida Lindl., Bojer ex Hook., Bot. Mag. 85: t. 5093. 1859, not:
Bot. Reg. 9: t. 724. 1823, not: Hippeastrum candidum Dracaena cylindrica Hook.f.
Stapf. Etymology: Named for Zephyranthes, now a Dracaena arborescens (Cornu ex Gérôme & Labroy)
synonym of Hippeastrum. (Figure 8D) Byng & Christenh., comb. nov. Basionym:
Miersia humilis (Phil.) M.F.Fay & Christenh., comb. Sansevieria arborescens Cornu ex Gérôme & Labroy,
nov. Basionym: Geanthus humilis Phil., Descr. Nuev. Bull. Mus. Hist. Nat. (Paris) 9: 173. 1903.
Pl., reimpr. 1884: 8. 1884. Dracaena ascendens (L.E.Newton) Byng & Christenh.,
Miersia triloba (Ravenna) M.F.Fay & Christenh., comb. comb. nov. Basionym: Sansevieria ascendens
nov. Basionym: Speea triloba Ravenna, Pl. Life 34: L.E.Newton, Bradleya 28: 19. 2010.
148. 1978. Dracaena aubrytiana (Carrière) Byng & Christenh.,
Urceolina amazonica (Linden ex Planch.) Christenh. comb. nov. Basionym: Sansevieria aubrytiana
& Byng, comb. nov. Basionym: Eucharis amazonica

Carrière, Rev. Hort. (Paris) 1861: 449. 1861. Mattei, Boll. Stud. Inform. Reale Giardino Colon. 4:
Dracaena bacularis (Pfennig ex A.Butler & Jankalski) 170. 1918.
Byng & Christenh., comb. nov. Basionym: Dracaena fasciata (Cornu ex Gérôme & Labroy) Byng
Sansevieria bacularis Pfennig ex A.Butler & Jankalski, & Christenh., comb. nov. Basionym: Sansevieria
Sansevieria 22: 3. 2010. fasciata Cornu ex Gérôme & Labroy, Bull. Mus. Hist.
Dracaena bagamoyensis (N.E.Br.) Byng & Christenh., Nat. (Paris) 9: 173. 1903.
comb. nov. Basionym: Sansevieria bagamoyensis Dracaena forskaoliana (Schult. & Schult.f.) Byng
N.E.Br., Bull. Misc. Inform. Kew 1913: 306. 1913. & Christenh., comb. nov. Basionym: Smilacina
Dracaena ballyi (L.E.Newton) Byng & Christenh., forskaoliana Schult. & Schult.f., Syst. Veg., ed. 15 bis,
comb. nov. Basionym: Sansevieria ballyi L.E.Newton, 7: 304. 1829, as ‘forskaliana’.
Brit. Cactus Succ. J. 22: 11. 2004. Dracaena francisii (Chahin.) Byng & Christenh.,
Dracaena bella (L.E.Newton) Byng & Christenh., comb. comb. nov. Basionym: Sansevieria francisii Chahin.,
nov. Basionym: Sansevieria bella L.E.Newton, Cact. Sansevieria J. 4: 12. 1995.
Succ. J. (Los Angeles) 72: 224. 2000. Dracaena frequens (Chahin.) Byng & Christenh., comb.
Dracaena bugandana Byng & Christenh., nom. nov. nov. Basionym: Sansevieria frequens Chahin., Cact.
Replaced synonym: Sansevieria lineata T.G.Forrest, Succ. J. (Los Angeles) 72: 130. 2000.
Sansevieria 30: 11. 2013, not: Dracaena lineata hort. Dracaena gracillima (Chahin.) Byng & Christenh.,
ex Baker. Etymology: This species is found in Uganda, comb. nov. Basionym: Sansevieria gracillima
which takes its name from the Buganda kingdom. Chahin., Sansevieria 12: 3. 2005.
Dracaena burdettii (Chahin.) Byng & Christenh., comb. Dracaena hallii (Chahin.) Byng & Christenh., comb. nov.
nov. Basionym: Sansevieria burdettii Chahin., Brit. Basionym: Sansevieria hallii Chahin., Sansevieria J. 5:
Cactus Succ. J. 18: 132. 2000. 7. 1996.
Dracaena burmanica (N.E.Br.) Byng & Christenh., Dracaena hargeisana (Chahin.) Byng & Christenh.,
comb. nov. Basionym: Sansevieria burmanica comb. nov. Basionym: Sansevieria hargeisana
N.E.Br., Bull. Misc. Inform. Kew 1915: 228. 1915. Chahin., Sansevieria J. 3: 53. 1994.
Dracaena canaliculata (Carrière) Byng & Christenh., Dracaena humiflora (D.J.Richards) Byng & Christenh.,
comb. nov. Basionym: Sansevieria canaliculata comb. nov. Basionym: Sansevieria humiflora
Carrière, Rev. Hort. (Paris) 1861: 449. 1861. D.J.Richards, Sansevieria 10: 3. 2004.
Dracaena caulescens (N.E.Br.) Byng & Christenh., Dracaena ×itumea (Mbugua) Byng & Christenh., comb.
comb. nov. Basionym: Sansevieria caulescens nov. Basionym: Sansevieria aethiopica subsp. itumea
N.E.Br., Bull. Misc. Inform. Kew 1915: 200. 1915. Mbugua, Fl. Trop. E. Afr., Dracaenac. : 17. 2007. (= D.
Dracaena conspicua N.E.Br., Byng & Christenh., comb. serpenta × D. raffillii)
nov. Basionym: Sansevieria conspicua N.E.Br., Bull. Dracaena liberica (Gérôme & Labroy) Byng &
Misc. Inform. Kew 1913: 306. 1913. Christenh., comb. nov. Basionym: Sansevieria
Dracaena dawei (Stapf) Byng & Christenh., comb. nov. liberica Gérôme & Labroy, Bull. Mus. Hist. Nat.
Basionym: Sansevieria dawei Stapf, J. Linn. Soc., Bot. (Paris) 9: 173. 1903.
37: 529. 1906. (Figure 8E) Dracaena longiflora (Sims) Byng & Christenh., comb.
Dracaena dooneri (N.E.Br.) Byng & Christenh., comb. nov. Basionym: Sansevieria longiflora Sims, Bot.
nov. Basionym: Sansevieria dooneri N.E.Br., Bull. Mag. 53: t. 2634. 1826.
Misc. Inform. Kew 1915: 231. 1915. Dracaena longiflora (Sims) Byng & Christenh. var.
Dracaena downsii (Chahin.) Byng & Christenh., comb. fernandopoensis (N.E.Br.) Byng & Christenh.,
nov. Basionym: Sansevieria downsii Chahin., Brit. comb. nov. Basionym: Sansevieria longiflora var.
Cactus Succ. J. 18: 133. 2000. fernandopoensis N.E.Br., Bull. Misc. Inform. Kew
1915: 257. 1915.
Dracaena dumetescens (L.E.Newton) Byng &
Christenh., comb. nov. Basionym: Sansevieria Dracaena longistyla (la Croix) Byng & Christenh.,
dumetescens L.E.Newton, Bradleya 27: 157. 2009. comb. nov. Basionym: Sansevieria longistyla la Croix,
Kew Bull. 59: 617. 2004.
Dracaena ebracteata (Cav.) Byng & Christenh., comb.
nov. Basionym: Salmia ebracteata Cav., Icon. 3: 24. Dracaena lunatifolia (L.E.Newton) Byng & Christenh.,
1795. comb. nov. Basionym: Sansevieria lunatifolia
L.E.Newton, Cact. Succ. J. (Los Angeles) 87: 118.
Dracaena eilensis (Chahin.) Byng & Christenh.,
2015.
comb. nov. Basionym: Sansevieria eilensis Chahin.,
Sansevieria J. 4: 9. 1995. Dracaena malawiana Byng & Christenh., nom. nov.
Replaced synonym: Sansevieria formosa Chahin.,
Dracaena erythraeae (Mattei) Byng & Christenh.,
Sansevieria 26: 2. 2012, not: Dracaena formosa
comb. nov. Basionym: Sansevieria erythraeae

65
hort. ex Baker. Etymology: This species is only found & stat. nov. Basionym: Sansevieria pinguicula var.
in Malawi, hence the name. nana Chahin., Sansevieria 30: 15. 2013.
Dracaena masoniana (Chahin.) Byng & Christenh., Dracaena powellii (N.E.Br.) Byng & Christenh., comb.
comb. nov. Basionym: Sansevieria masoniana nov. Basionym: Sansevieria powellii N.E.Br., Bull.
Chahin., Cact. Succ. J. (Los Angeles) 72: 31. 2000. Misc. Inform. Kew 1915: 198. 1915.
Dracaena newtoniana (T.G.Forrest) Byng & Christenh., Dracaena powysii (L.E.Newton) Byng & Christenh.,
comb. nov. Basionym: Sansevieria newtoniana comb. nov. Basionym: Sansevieria powysii
T.G.Forrest, Cact. Succ. J. (Los Angeles) 86: 76. 2014. L.E.Newton, Bradleya 28: 24. 2010.
Dracaena nilotica (Baker) Byng & Christenh., comb. Dracaena raffillii (N.E.Br.) Byng & Christenh., comb.
nov. Basionym: Sansevieria nilotica Baker, J. Linn. nov. Basionym: Sansevieria raffillii N.E.Br., Bull. Misc.
Soc., Bot. 14: 548. 1875. Inform. Kew 1915: 252. 1915.
Dracaena nitida (Chahin.) Byng & Christenh., comb. Dracaena roxburghiana (Schult. & Schult.f.) Byng
nov. Basionym: Sansevieria nitida Chahin., Cact. & Christenh., comb. nov. Basionym: Sansevieria
Succ. J. (Los Angeles) 73: 120. 2001. roxburghiana Schult. & Schult.f. in J.J.Roemer &
Dracaena parva (N.E.Br.) Byng & Christenh., comb. J.A.Schultes, Syst. Veg., ed. 15 bis 7: 357. 1829.
nov. Basionym: Sansevieria parva N.E.Br., Bull. Misc. Dracaena sambiranensis (H.Perrier) Byng & Christenh.,
Inform. Kew 1915: 233. 1915. (Figure 8F) comb. nov. Basionym: Sansevieria sambiranensis
Dracaena patens (N.E.Br.) Byng & Christenh., comb. H.Perrier, Notul. Syst. (Paris) 5: 154. 1936.
nov. Basionym: Sansevieria patens N.E.Br., Bull. Dracaena scimitariformis (D.J.Richards) Byng &
Misc. Inform. Kew 1915: 210. 1915. Christenh., comb. nov. Basionym: Sansevieria
Dracaena pearsonii (N.E.Br.) Byng & Christenh., comb. scimitariformis D.J.Richards, Sansevieria 5: 8. 2002.
nov. Basionym: Sansevieria pearsonii N.E.Br., Bull. Dracaena senegambica (Baker) Byng & Christenh.,
Misc. Inform. Kew 1911: 97. 1911. comb. nov. Basionym: Sansevieria senegambica
Dracaena pedicellata (la Croix) Byng & Christenh., Baker, J. Linn. Soc., Bot. 14: 548.1875.
comb. nov. Basionym: Sansevieria pedicellata la Dracaena serpenta Byng & Christenh., nom. nov.
Croix, Kew Bull. 59: 620. 2004. Replaced synonym: Sansevieria gracilis N.E.Br., Bull.
Dracaena perrotii (Warb.) Byng & Christenh., Misc. Inform. Kew 1911: 96. 1911, not: Dracaena
comb. nov. Basionym: Sansevieria perrotii Warb., gracilis Salisb. Etymology: This is often called ‘snake
Tropenpflanzer 5: 191. 1901. plant’, hence the epithet chosen for this species.
Dracaena pethera Byng & Christenh., nom. nov. Dracaena serpenta Byng & Christenh. var. humbertiana
Replaced synonym: Sansevieria kirkii Baker, Bull. (Guillaumin) Byng & Christenh., comb. nov.
Misc. Inform. Kew 1887: 3. 1887, not: Dracaena kirkii Basionym: Sansevieria humbertiana Guillaumin,
Baker. Etymology: From Greek πεθερά, mother-in- Bull. Mus. Natl. Hist. Nat., sér. 2, 12: 353. 1940.
law. Dracaena singularis (N.E.Br.) Byng & Christenh.,
Dracaena pethera Byng & Christenh. var. pulchra comb. nov. Basionym: Sansevieria singularis N.E.Br.,
(N.E.Br.) Byng & Christenh., comb. nov. Basionym: Bull. Misc. Inform. Kew 1911: 97. 1911. Replaced
Sansevieria kirkii var. pulchra N.E.Br., Bull. Misc. synonym: Boophone fischeri Baker, Fl. Trop. Afr. 7:
Inform. Kew 1915: 256. 1915. 577. 1898, not: Dracaena fischeri Baker.
Dracaena pfisteri (D.J.Richards) Byng & Christenh., Dracaena sinussimiorum (Chahin.) Byng & Christenh.,
comb. nov. Basionym: Sansevieria pfisteri comb. nov. Basionym: Sansevieria sinussimiorum
D.J.Richards, Sansevieria 19: 4. 2009. Chahin., Sansevieria 3: 24. 2002.
Dracaena phillipsiae (N.E.Br.) Byng & Christenh., Dracaena sordida (N.E.Br.) Byng & Christenh., comb.
comb. nov. Basionym: Sansevieria phillipsiae N.E.Br., nov. Basionym: Sansevieria sordida N.E.Br., Bull.
Hooker’s Icon. Pl. 30: t. 3000. 1913. Misc. Inform. Kew 1915: 214. 1915.
Dracaena pinguicula (P.R.O.Bally) Byng & Christenh., Dracaena spathulata Byng & Christenh., nom. nov.
comb. nov. Basionym: Sansevieria pinguicula Replaced synonym: Sansevieria concinna N.E.Br.,
P.R.O.Bally, Candollea 19: 145. 1964. Bull. Misc. Inform. Kew 1915: 233. 1915, not:
Dracaena concinna Kunth. Etymology: This species
Dracaena pinguicula (P.R.O.Bally) Byng & Christenh.
has spoon-like leaves, hence the name.
subsp. disticha (Pfennig ex A.Butler) Byng &
Christenh., comb. nov. Basionym: Sansevieria Dracaena stuckyi (God.-Leb.) Byng & Christenh.,
pinguicula subsp. disticha Pfennig ex A.Butler, comb. nov. Basionym: Sansevieria stuckyi God.-Leb.,
Sansevieria 28: 17. 2012. Sansev. Gigant. Afr. Or.: 13. 1903.
Dracaena pinguicula (P.R.O.Bally) Byng & Christenh. Dracaena subspicata (Baker) Byng & Christenh., comb.
subsp. nana (Chahin.) Byng & Christenh., comb. nov. Basionym: Sansevieria subspicata Baker, Gard.

Chron., ser. 3, 6: 436. 1889. Rhodora 77: 253. 1975.

Dracaena subtilis (N.E.Br.) Byng & Christenh., comb. Callisia disgrega (Kunth) Christenh. & Byng, comb.
nov. Basionym: Sansevieria subtilis N.E.Br., Bull. nov. Basionym: Tradescantia disgrega Kunth, Enum.
Misc. Inform. Kew 1915: 237. 1915. Pl. 4: 97. 1843.
Dracaena suffruticosa (N.E.Br.) Byng & Christenh., Callisia diuretica (Mart.) Christenh. & Byng, comb.
comb. nov. Basionym: Sansevieria suffruticosa nov. Basionym: Tradescantia diuretica Mart., Reise
N.E.Br., Bull. Misc. Inform. Kew 1915: 202. 1915. Bras. 1: 281. 1823.
Dracaena suffruticosa (N.E.Br.) Byng & Christenh. var. Callisia elata (D.R.Hunt) Christenh. & Byng, comb. nov.
longituba (Pfennig) Byng & Christenh., comb. nov. Basionym: Tripogandra elata D.R.Hunt, Kew Bull. 33:
Basionym: Sansevieria suffruticosa var. longituba 405. 1979.
Pfennig, Bot. Jahrb. Syst. 102: 178. 1981. Callisia encolea (Diels) Christenh. & Byng, comb. nov.
Dracaena testudinea Byng & Christenh., nom. nov. Basionym: Tradescantia encolea Diels, Bot. Jahrb.
Replaced synonym: Sansevieria braunii Engl. Syst. 37: 385. 1906.
& Krause, Bot. Jahrb. Syst. 45: 153. 1910, not: Callisia glandulosa (Seub.) Christenh. & Byng, comb.
Dracaena braunii Engl. Etymology: The white- nov. Basionym: Tradescantia glandulosa Seub. in
mottled purplish-green leaves resemble a tortoise Martius (ed.), Fl. Bras. 3: 253. 1855.
shell, hence the name. Callisia ionantha (Diels) Christenh. & Byng, comb. nov.
Dracaena varians (N.E.Br.) Byng & Christenh., comb. Basionym: Tradescantia ionantha Diels, Bot. Jahrb.
nov. Basionym: Sansevieria varians N.E.Br., Bull. Syst. 37: 382. 1906.
Misc. Inform. Kew 1915: 209. 1915. Callisia kruseana (Matuda) Christenh. & Byng, comb.
Dracaena volkensii (Gürke) Byng & Christenh., comb. nov. Basionym: Tripogandra kruseana Matuda,
nov. Basionym: Sansevieria volkensii Gürke in Anales Inst. Biol. Univ. Nac. México 37: 77. 1967.
H.G.A.Engler, Pflanzenw. Ost-Afrikas, C: 144. 1895. Callisia mexicana Christenh. & Byng, nom. nov.
Dracaena zebra Byng & Christenh., nom. nov. Replaced Replaced synonym: Tripogandra guerrerensis
synonym: Sansevieria metallica Gérôme & Labroy, Matuda, Anales Inst. Biol. Univ. Nac. México 36: 113.
Bull. Mus. Hist. Nat. (Paris) 9: 173. 1903, not: 1966, not: Callisia guerrerensis Matuda. Etymology:
Dracaena metallica W.Bull. Etymology: This species This species is found in Mexico.
has silver-striped leaves, like the stripes of a zebra. Callisia montana (Handlos) Christenh. & Byng, comb.
Dracaena zebra Byng & Christenh. var. nyasica Byng nov. Basionym: Tripogandra montana Handlos,
& Christenh., comb. nov. Basionym: Sansevieria Rhodora 77: 276. 1975.
metallica var. nyasica N.E.Br., Bull. Misc. Inform. Kew Callisia neglecta (Handlos) Christenh. & Byng, comb.
1915: 247. 1915. nov. Basionym: Tripogandra neglecta Handlos,
Drimia jackyi (Knirsch, Mart.-Azorin & Wetschnig) Rhodora 77: 287. 1975.
Christenh. & Byng, comb. nov. Basionym: Callisia palmeri (Rose) Christenh. & Byng, comb. nov.
Rhodocodon jackyi Knirsch, Mart.-Azorin & Basionym: Tradescantia palmeri Rose, Contr. U.S.
Wetschnig, Phytotaxa 253: 202. 2016. Natl. Herb. 1: 113. 1891.
Callisia purpurascens (S.Schauer) Christenh. & Byng,
78. Commelinaceae References: Evans et al. (2003), comb. nov. Basionym: Tradescantia purpurascens
Hertweck et al. (2014). S.Schauer, Linnaea 19: 700. 1847.
Note: Callisia Loefl. is expanded to include Tripogandra Callisia saxicola (Greenm.) Christenh. & Byng, comb.
Raf. nov. Basionym: Tradescantia saxicola Greenm.,
Callisia amplexans (Handlos) Christenh. & Byng, comb. Proc. Amer. Acad. Arts 39: 70. 1903.
nov. Basionym: Tripogandra amplexans Handlos, Callisia serrulata (Vahl) Christenh. & Byng, comb. nov.
Rhodora 77: 242. 1975. Basionym: Commelina serrulata Vahl, Eclog. Amer. 2:
Callisia amplexicaulis (Klotzsch ex C.B.Clarke) 4. 1798.
Christenh. & Byng, comb. nov. Basionym: Callisia silvatica (Handlos) Christenh. & Byng, comb.
Tradescantia amplexicaulis Klotzsch ex C.B.Clarke in nov. Basionym: Tripogandra silvatica Handlos,
A.L.P.P. de Candolle & A.C.P. de Candolle, Monogr. Rhodora 77: 310. 1975.
Phan. 3: 304. 1881. Callisia tripogandra Christenh. & Byng, nom. nov.
Callisia angustifolia (B.L.Rob.) Christenh. & Byng, Replaced synonym: Tradescantia multiflora Sw.,
comb. nov. Basionym: Tradescantia angustifolia Prodr. Veg. Ind. Occ.: 57. 1788, not: Callisia multiflora
B.L.Rob., Proc. Amer. Acad. Arts 27: 185. 1893. (M.Martens & Galeotti) Standl. Etymology: This
Callisia brasiliensis (Handlos) Christenh. & Byng, comb. species is the type of the genus Tripogandra, now
nov. Basionym: Tripogandra brasiliensis Handlos,

67
included in Callisia. Manzan. & Gouda, Phytotaxa 3: 2. 2010.

Callisia warmingiana (Seub.) Christenh. & Byng, comb. Tillandsia euryelytra (J.R.Grant) Christenh. & Byng,
nov. Basionym: Tradescantia warmingiana Seub., comb. nov. Basionym: Racinaea euryelytra J.R.Grant,
Vidensk. Meddel. Naturhist. Foren. Kjøbenhavn Phytologia 76: 284. 1994.
1872: 126. 1872. Tillandsia gentryana (Manzan. & W.Till) Christenh. &
Cyanotis kewoides Christenh. & Byng, nom. nov. Byng, comb. nov. Basionym: Racinaea gentryana
Replaced synonym: Belosynapsis kewensis Hassk., Manzan. & W.Till, J. Bromeliad Soc. 60: 159. 2011.
Flora 54: 259. 1871, not: Cyanotis kewensis Tillandsia geotropica Christenh. & Byng, nom. nov.
C.B.Clarke. Etymology: This species has a superficial Replaced synonym: Racinaea penduliflora Gouda
resemblance to Kewa (Kewaceae). & Manzan., J. Bromeliad Soc. 58: 156. 2008, not:
Tillandsia penduliflora Griseb. Etymology: The
91. Bromeliaceae. References: Barfuss et al. (2005), inflorescences of this species are downward secund,
Givnish et al. (2011), Evans et al. (2015). pointing to the earth, hence the name.
Note: To prevent unecessary splitting of traditional Tillandsia goudai (Manzan. & W.Till) Christenh. &
genera in Bromeliaceae, Cipuropsis Ule, Navia Byng, comb. nov. Basionym: Racinaea goudai
Schult.f., Tillandsia L. and Werauhia J.R.Grant are Manzan. & W.Till, J. Bromeliad Soc. 60: 210. 2011,
expanded, rather than split into several genera. as ‘goudae’.
Cipuropsis asplundii (L.B.Sm.) Christenh. & Byng, Tillandsia grantii (Manzan. & W.Till) Christenh. & Byng,
comb. nov. Basionym: Tillandsia asplundii L.B.Sm., comb. nov. Basionym: Racinaea grantii Manzan. &
Phytologia 6: 441. 1959. W.Till, J. Bromeliad Soc. 60: 166. 2011.
Cipuropsis bicolor (L.B.Sm.) Christenh. & Byng, comb. Tillandsia guacamayosensis (Manzan. & Gouda)
nov. Basionym: Mezobromelia bicolor L.B.Sm., Proc. Christenh. & Byng, comb. nov. Basionym: Racinaea
Amer. Acad. Arts 70: 151. 1935. guacamayosensis Manzan. & Gouda, Phytotaxa 3: 5.
2010.
Cipuropsis capituligera (Griseb.) Christenh. & Byng,
comb. nov. Basionym: Tillandsia capituligera Tillandsia hoepfelii Christenh. & Byng, nom. nov.
Griseb., Cat. Pl. Cub.: 254. 1866. Replaced synonym: Racinaea wuelfinghoffii Höpfel
& Scharf, Bromelie 2008: 136. 2008, not: Tillandsia
Cipuropsis delicatula (L.B.Sm.) Christenh. & Byng,
wuelfinghoffii Ehlers. Etymology: This species was
comb. nov. Basionym: Tillandsia delicatula L.B.Sm.,
named for Frank Höpfel, who first described it.
Phytologia 5: 36. 1954.
Tillandsia kalliantha (J.R.Grant) Christenh. & Byng,
Cipuropsis hospitalis (L.B.Sm.) Christenh. & Byng,
comb. nov. Basionym: Racinaea kalliantha J.R.Grant,
comb. nov. Basionym: Tillandsia hospitalis L.B.Sm.,
Caldasia 5: 13. 1948.
Tillandsia lymansmithiana (J.R.Grant) Christenh.
Cipuropsis magdalenae (L.B.Sm.) Christenh. & Byng,
& Byng, comb. nov. Basionym: Racinaea
comb. nov. Basionym: Vriesea magdalenae L.B.Sm.,
lymansmithiana J.R.Grant, Phytologia 76: 286. 1994.
Tillandsia macrantha (H.Luther) Christenh. & Byng,
Cipuropsis pinnata (Mez & Sodiro) Christenh. & Byng,
comb. nov. Basionym: Racinaea macrantha
comb. nov. Basionym: Tillandsia pinnata Mez &
H.Luther, Selbyana 28: 10. 2007.
Sodiro, Bull. Herb. Boissier, sér. 2, 4: 1130. 1904.
Tillandsia manzanaresii Christenh. & Byng, nom. nov.
Cipuropsis singularis (Mez & Wercklé) Christenh. &
Replaced synonym: Racinaea lutheri Manzan. &
Byng, comb. nov. Basionym: Tillandsia singularis
W.Till, J. Bromeliad Soc. 62: 264. 2013, not: Tillandsia
Mez & Wercklé, Bull. Herb. Boissier, sér. 2, 5: 103.
lutheri (Manzan. & W.Till) J.R.Grant. Etymology: This
1905.
species is named for José Manuel Manzanares, who
Cipuropsis truncata (L.B.Sm.) Christenh. & Byng, comb. first described it.
nov. Basionym: Tillandsia truncata L.B.Sm., Contr.
Tillandsia marioportillae (Höpfel & Scharf) Christenh.
U.S. Natl. Herb. 29: 530. 1954.
& Byng, comb. nov. Basionym: Racinaea
Navia duidensis (L.B.Sm.) Christenh. & Byng, comb. marioportillae Höpfel & Scharf, Bromelie 2008: 97.
nov. Basionym: Brewcaria duidensis L.B.Sm., 2008.
Steyerm. & H.Rob., Acta Bot. Venez. 14: 10. 1984.
Tillandsia pattersoniae (Manzan. & W.Till) Christenh. &
Navia marahuacae (L.B.Sm., Steyerm. & H.Rob.) Byng, comb. nov. Basionym: Racinaea pattersoniae
Christenh. & Byng, comb. nov. Basionym: Brewcaria Manzan. & W.Till, J. Bromeliad Soc. 57: 201. 2007.
marahuacae L.B.Sm., Steyerm. & H.Rob., Ann.
Tillandsia racinaea Christenh. & Byng, nom. nov.
Missouri Bot. Gard. 73: 714. 1987.
Replaced synonym: Racinaea kessleri H.Luther,
Tillandsia condorensis (Manzan. & Gouda) Christenh. Selbyana 19: 86. 1998, not: Tillandsia kessleri
& Byng, comb. nov. Basionym: Racinaea condorensis

H.Luther. Etymology: Named for the genus Racinaea, beyrichianum Sporl. ex Körn., Linnaea 27: 567. 1856.
now a synonym of Tillandsia. Paepalanthus cubensis (Ruhland) Christenh. & Byng,
Tillandsia scharfii Christenh. & Byng, nom. nov. comb. nov. Basionym: Lachnocaulon cubense
Replaced synonym: Racinaea hasei Höpfel & Scharf, Ruhland, Repert. Spec. Nov. Regni Veg. 22: 34. 1925.
Bromelie 2011: 133. 2011, not: Tillandsia hasei Paepalanthus deflexus (F.N.Costa) Christenh. & Byng,
Ehlers & L.Hrom. Etymology: This species is named comb. nov. Basionym: Actinocephalus deflexus
for Uwe Scharf, who first described it. F.N.Costa, Novon 16: 214. 2006.
Tillandsia strobeliorum (Manzan. & W.Till) Christenh. & Paepalanthus delicatus (Sano) Christenh. & Byng,
Byng, comb. nov. Basionym: Racinaea strobeliorum comb. nov. Basionym: Actinocephalus delicatus
Manzan. & W.Till, J. Bromeliad Soc. 59: 109. 2009. Sano, Phytotaxa 27: 27. 2011.
Tillandsia terrestris (Manzan. & Gouda) Christenh. Paepalanthus digynus (Körn.) Christenh. & Byng,
& Byng, comb. nov. Basionym: Racinaea terrestris comb. nov. Basionym: Lachnocaulon digynum Körn.,
Manzan. & Gouda, Phytotaxa 3: 9. 2010. Linnaea 27: 570. 1856.
Tillandsia zingleri (Höpfel & Scharf) Christenh. & Byng, Paepalanthus ekmanii (Ruhland) Christenh. & Byng,
comb. nov. Basionym: Racinaea zingleri Höpfel & comb. nov. Basionym: Lachnocaulon ekmanii
Scharf, Bromelie 2011: 128. 2011. Ruhland, Repert. Spec. Nov. Regni Veg. 22: 34. 1925.
Tillandsia zonai Christenh. & Byng, nom. nov. Paepalanthus engleri (Ruhland) Christenh. & Byng,
Replaced synonym: Racinaea tillii Manzan. & comb. nov. Basionym: Lachnocaulon engleri Ruhland
Gouda, Phytotaxa 3: 13. 2010, not: Tillandsia tillii in H.G.A.Engler (ed.), Pflanzenr. IV, 30: 241. 1903.
Ehlers. Etymology: The species is named for Scott Paepalanthus fluviatilis (Aubl.) Christenh. & Byng,
Zona, for his contribution to the understanding of comb. nov. Basionym: Tonina fluviatilis Aubl., Hist.
littertrapping in plants, particularly in Bromeliaceae. Pl. Guiane 2: 857. 1775.
Werauhia bibeatricis (Morillo) Christenh. & Byng, Paepalanthus giuliettiae (Sano) Christenh. & Byng,
comb. nov. Basionym: Vriesea bibeatricis Morillo, comb. nov. Basionym: Actinocephalus giuliettiae
Ernstia 39: 2. 1986, as ‘bi-beatricis’. Sano, Phytotaxa 27: 31. 2011.
Werauhia glutinosa (Lindl.) Christenh. & Byng, comb. Paepalanthus glabrescens (Silveira) Christenh. & Byng,
nov. Basionym: Vriesea glutinosa Lindl., Gard. Chron. comb. nov. Basionym: Paepalanthus ciliatus var.
1856: 388. 1856. glabrescens Silveira, Floral. Mont. 1: 213. 1928.
Werauhia monstrum (Mez) Christenh. & Byng, comb. Paepalanthus graminifolius (F.N.Costa) Christenh.
nov. Basionym: Tillandsia monstrum Mez, Repert. & Byng, comb. nov. Basionym: Actinocephalus
Spec. Nov. Regni Veg. 16: 78. 1919. graminifolius F.N.Costa, Novon 16: 215. 2006.
Werauhia soderstromii (L.B.Sm.) Christenh. & Byng, Paepalanthus koernickeanus (Trovó & F.N.Costa)
comb. nov. Basionym: Vriesea soderstromii L.B.Sm., Christenh. & Byng, comb. nov. Basionym:
Bull. Bromeliad Soc. 13: 53. 1963. Actinocephalus koernickeanus Trovó & F.N.Costa,
Werauhia splendens (Brongn.) Christenh. & Byng, Novon 19: 256. 2009.
comb. nov. Basionym: Tillandsia splendens Brongn., Paepalanthus minus (Chapm.) Christenh. & Byng,
Hort. Universel 6: 199. 1845. comb. nov. Basionym: Lachnocaulon michauxii var.
minus Chapm., Fl. South. U.S., ed. 3: 531. 1897.
94. Eriocaulaceae. Reference: Andrade et al. (2010). Paepalanthus verae (Sano & Trovó) Christenh. & Byng,
Note: Paepalanthus Mart., nom. cons. is paraphyletic comb. nov. Basionym: Actinocephalus verae Sano &
with Actinocephalus (Kórn.) Sano, Lachnocaulon Trovó, Brittonia 62: 35. 2010.
Kunth and Tonina Aubl. embedded. The genus was
officially expanded by Mabberley (2017), but not all 106. Papaveraceae. References: Carolan et al. (2008),
combinations were transferred, which are hereby Kadereit et al. (2011), Grey-Wilson (2012).
provided.
Note: The circumscription of Papaver L. has been
Paepalanthus aggregatus (F.N.Costa) Christenh. contentious, and molecular studies have made
& Byng, comb. nov. Basionym: Actinocephalus the situation worse with Meconopsis Vig. being
aggregatus F.N.Costa, Novon 16: 212. 2006. embedded and polyphyletic, with the type M.
Paepalanthus anceps (Walter) Christenh. & Byng, cambrica Vig. deeply embedded in Papaver and
comb. nov. Basionym: Eriocaulon anceps Walter, Fl. another clade of Meconopsis being embedded in a
Carol.: 83. 1788. seprarate clade of Papaver. The genera Stylomecon
Paepalanthus beyrichianus (Sporl. ex Körn.) Christenh. G.Taylor and Roemeria Medik. are also embedded
& Byng, comb. nov. Basionym: Lachnocaulon in Papaver. As a result the type of Meconopsis was
changed, but in doing so the genus Meconopsis is

69
still not monophyletic, resulting in the acceptance 1891.

of Cathcarthia Hook.f.. As these genera have few Papaver comptum (Prain) Christenh. & Byng, comb.
separating characters (mainly presence or absence nov. Basionym: Meconopsis compta Prain, Bull.
of style, which is not consistent within the genera) Misc. Inform. Kew 1918: 212. 1918.
and all are known as poppies, it is better to Papaver concinnum (Prain) Christenh. & Byng, comb.
expand the genus Papaver to include Cathcarthia, nov. Basionym: Meconopsis concinna Prain, Bull.
Meconopsis and Roemeria. Misc. Inform. Kew 1915: 163. 1915.
Papaver argemonanthum (Prain) Christenh. & Byng, Papaver ×cookei (G.Taylor) Christenh. & Byng, comb.
comb. nov. Basionym: Meconopsis argemonantha nov. Basionym: Meconopsis ×cookei G.Taylor, J.
Prain, Bull. Misc. Inform. Kew 1915: 161. 1915. Roy. Hort. Soc., London 76: 231. 1951. (= Papaver
Papaver autumnale (P.A.Egan) Christenh. & Byng, puniceum × Papaver quintuplinervium).
comb. nov. Basionym: Meconopsis autumnalis Papaver delavayi (Franch.) Christenh. & Byng, comb.
P.A.Egan, Phytotaxa 20: 48. 2011. nov. Basionym: Cathcartia delavayi Franch., Bull.
Papaver baileyi (Prain) Christenh. & Byng, comb. nov. Soc. Bot. France 33: 390. 1886.
Basionym: Meconopsis baileyi Prain, Bull. Misc. Papaver dhwojii (G.Taylor ex Hay) Christenh. & Byng,
Inform. Kew 1915: 161. 1915. comb. nov. Basionym: Meconopsis dhwojii G.Taylor
Papaver balangense (Tosh.Yoshida, H.Sun & Boufford) ex Hay, New Fl. & Silva 4: 225. 1932.
Christenh. & Byng, comb. nov. Basionym: Papaver discigerum (Prain) Christenh. & Byng, comb.
Meconopsis balangensis Tosh.Yoshida, H.Sun & nov. Basionym: Meconopsis discigera Prain, Ann.
Boufford, Acta Bot. Yunnan. 33: 409. 2011. Bot. (Oxford) 20: 356. 1906.
Papaver bellum (Prain) Christenh. & Byng, comb. nov. Papaver exile (Tosh.Yoshida, H.Sun & Grey-Wilson)
Basionym: Meconopsis bella Prain, J. Asiat. Soc. Christenh. & Byng, comb. nov. Basionym:
Bengal, Pt. 2, Nat. Hist. 63: 82. 1894. Meconopsis exilis Tosh.Yoshida, H.Sun & Grey-
Papaver betonicifolium (Franch.) Christenh. & Byng, Wilson, Curtis’s Bot. Mag. 29: 204. 2012.
comb. nov. Basionym: Meconopsis betonicifolia Papaver florindae (Kingdon-Ward) Christenh. &
Franch., Pl. Delavay. 1: 42. 1889. (Figure 9A) Byng, comb. nov. Basionym: Meconopsis florindae
Papaver bhutanicum (Tosh.Yoshida & Grey-Wilson) Kingdon-Ward, Gard. Chron., ser. 3, 79: 232. 1926.
Christenh. & Byng, comb. nov. Basionym: Papaver forrestii (Prain) Christenh. & Byng, comb. nov.
Meconopsis bhutanica Tosh.Yoshida & Grey-Wilson, Basionym: Meconopsis forrestii Prain, Bull. Misc.
Plantsman n.s. 11: 98. 2012. Inform. Kew 1907: 316. 1907.
Papaver bijiangense (H.Ohba, Tosh.Yoshida & Papaver ganeshense (Grey-Wilson) Christenh. & Byng,
H.Sun) Christenh. & Byng, comb. nov. Basionym: comb. nov. Basionym: Meconopsis ganeshensis
Meconopsis bijiangensis H.Ohba, Tosh.Yoshida & Grey-Wilson, Curtis’s Bot. Mag. 23: 188. 2006.
H.Sun, J. Jap. Bot. 84: 294. 2009.
Papaver georgei (G.Taylor) Christenh. & Byng, comb.
Papaver bulbiliferum (Tosh.Yoshida, H.Sun & Grey- nov. Basionym: Meconopsis georgei G.Taylor,
Wilson) Christenh. & Byng, comb. nov. Basionym: Account Gen. Meconopsis: 86. 1934.
Meconopsis bulbilifera Tosh.Yoshida, H.Sun & Grey-
Papaver gracilipes (G.Taylor) Christenh. & Byng, comb.
Wilson, Curtis’s Bot. Mag. 29: 200. 2012.
nov. Basionym: Meconopsis gracilipes G.Taylor,
Papaver calciphilum (Kingdon-Ward) Christenh. & Account Gen. Meconopsis: 38. 1934.
Byng, comb. nov. Basionym: Meconopsis calciphila
Papaver grande (Prain) Christenh. & Byng, comb. nov.
Kingdon-Ward, Gard. Chron., ser. 3, 82: 506. 1927.
Basionym: Meconopsis grandis Prain, J. Asiat. Soc.
Papaver cathcartia Christenh. & Byng, nom. nov. Bengal, Pt. 2, Nat. Hist. 64: 320. 1896.
Replaced synonym: Cathcartia villosa Hook.f.,
Papaver guilelmi-waldemarii (Klotzsch & Garcke)
Bot. Mag. 77: t. 4596. 1851, not: Papaver villosum
K.Koch. Etymology: Named for the genus Cathcartia,
Meconopsis guilelmi-waldemarii Klotzsch & Garcke,
now included in Papaver.
Bot. Ergebn. Reise Prinz Waldem.: 129. 1862.
Papaver chankheliense (Grey-Wilson) Christenh. Replaced synonym: Meconopsis aculeata Royle, Ill.
& Byng, comb. nov. Basionym: Meconopsis Bot. Himal. Mts.1: 67 1834, not: Papaver aculeatum
chankheliensis Grey-Wilson, Curtis’s Bot. Mag. 23: Thunb.
203. 2006.
Papaver ×harleyanum (G.Taylor) Christenh. & Byng,
Papaver chelidonifolium (Bureau & Franch.) Christenh. comb. nov. Basionym: Meconopsis ×harleyana
& Byng, comb. nov. Basionym: Meconopsis G.Taylor, New Fl. & Silva 6: 48. 1933. (= Papaver
chelidonifolia Bureau & Franch, J. Bot. (Morot) 5: 19. simplicifolium × Papaver integrifolium)
Papaver henrici (Bureau & Franch.) Christenh. & Byng,

comb. nov. Basionym: Meconopsis henrici Bureau & Boufford, Acta Bot. Yunnan. 34(2): 145. 2012.
Franch., J. Bot. (Morot) 5(2): 19. 1891. Papaver napaulense (DC.) Christenh. & Byng, comb.
Papaver heterandrum (Tosh.Yoshida, H.Sun & nov. Basionym: Meconopsis napaulensis DC., Prodr.
Boufford) Christenh. & Byng, comb. nov. Basionym: 1: 121. 1824.
Meconopsis heterandra Tosh.Yoshida, H.Sun & Papaver nyingchiense (L.H.Zhou) Christenh. & Byng,
Boufford, Acta Bot. Yunnan. 32(6): 505. 2010. comb. nov. Basionym: Meconopsis nyingchiensis
Papaver horridulum (Hook.f. & Thomson) Christenh. & L.H.Zhou, Bull. Bot. Lab. N.E. Forest. Inst., Harbin
Byng, comb. nov. Basionym: Meconopsis horridula 1980: 98. 1980.
Hook.f. & Thomson, Fl. Ind. 1: 252. 1855. Papaver oliverianum (Franch. ex Prain) Christenh. &
Papaver impeditum (Prain) Christenh. & Byng, comb. Byng, comb. nov. Basionym: Meconopsis oliveriana
nov. Basionym: Meconopsis impedita Prain, Bull. Franch. ex Prain, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist.
Misc. Inform. Kew 1915: 162. 1915. 64: 312. 1896.
Papaver kafiristanicum Christenh. & Byng, nom. nov. Papaver ouvrardianum (Hand.-Mazz.) Christenh.
Replaced synonym: Meconopsis neglecta G.Taylor, & Byng, comb. nov. Basionym: Meconopsis
Account Gen. Meconopsis: 102. 1934, not: Papaver ouvrardianum Hand.-Mazz., Anz. Akad. Wiss. Wien,
neglectum Fedde. Etymology: Named for Kāfiristān, Math.-Naturwiss. Kl. 59: 247. 1922. Replaced
a historical region around the province of Nuristan in synonym: Meconopsis speciosa Prain, Trans. & Proc.
Afghanistan, where this species is found. Bot. Soc. Edinburgh 23: 258. 1907, not: Papaver
Papaver ×kongboense (G.Taylor) Christenh. & Byng, speciosum Tourlet
comb. nov. Basionym: Meconopsis ×kongboensis Papaver pinnatifolium (C.Y.Wu & X.Zhuang) Christenh.
Grey-Wilson non Hort. ex R.L.Harley in Grey Wilson, & Byng, comb. nov. Basionym: Meconopsis
Gen. Meconopsis: 203. 2014. (= Papaver sulphureum pinnatifolia C.Y.Wu & X.Zhuang, Acta Phytotax. Sin.
× Papaver simplicifolium) 17: 114. 1979.
Papaver lamjungense (Tosh.Yoshida, H.Sun & Grey- Papaver polygonoides (Prain) Christenh. & Byng,
Wilson) Christenh. & Byng, comb. nov. Basionym: comb. nov. Basionym: Cathcartia polygonoides
Meconopsis lamjungensis Tosh.Yoshida, H.Sun & Prain, J. Asiat. Soc. Bengal. Pt. 2. Nat. Hist. 64: 326.
Grey-Wilson, Curtis’s Bot. Mag. 29: 207. 2012. 1896.
Papaver lancifolium (Franch.) Christenh. & Byng, Papaver prainianum (Kingdon-Ward) Christenh. &
comb. nov. Basionym: Cathcartia lancifolia Franch., Byng, comb. nov. Basionym: Meconopsis prainiana
Bull. Soc. Bot. France 33: 391. 1889. Kingdon-Ward, Gard. Chron., ser. 3, 79: 308. 1926.
Papaver latifolium (Prain) Christenh. & Byng, comb. Papaver prattii (Prain) Christenh. & Byng, comb. &
nov. Basionym: Meconopsis sinuata Prain var. stat. nov. Basionym: Meconopsis sinuata Prain var.
latifolia Prain, Curtis’s Bot. Mag. 134: f. 8223. 1908. prattii Prain, J. Asiat. Soc. Bengal. Pt. 2. Nat. Hist. 64:
Papaver lhasaense (Grey-Wilson) Christenh. & Byng, 314. 1896.
comb. nov. Basionym: Meconopsis lhasaensis Grey- Papaver primulinum (Prain) Christenh. & Byng, comb.
Wilson, Gen. Meconopsis: 248. 2014. nov. Basionym: Meconopsis primulina Prain, J. Asiat.
Papaver lijiangense (Grey-Wilson) Christenh. & Soc. Bengal, Pt. 2, Nat. Hist. 64: 319. 1896.
Byng, comb. & stat. nov. Basionym: Meconopsis Papaver przewalskii Christenh. & Byng, nom. nov.
integrifolia (Maxim.) Franch. susbp. lijiangensis Replaced synonym: Cathcartia integrifolia Maxim.,
Grey-Wilson, Plantsman n.s. 3: 33. 1996. Bull. Acad. Imp. Sci. Saint-Pétersbourg, sér. 3,
Papaver ludlowii (Grey-Wilson) Christenh. & Byng, 23: 310. 1877, not: Papaver integrifolium Vig.
comb. nov. Basionym: Meconopsis ludlowii Grey- Etymology: The type specimen of this species was
Wilson, Gen. Meconopsis : 350. 2014. collected by Nikolai Przewalski, and the species is
named in honour of him. (Figure 9B)
Papaver lyratum (H.A.Cummins & Prain) Christenh.
& Byng, comb. nov. Basionym: Cathcartia lyrata Papaver pseudointegrifolium (Prain) Christenh.
H.A.Cummins & Prain ex Prain, J. Asiat. Soc. Bengal, & Byng, comb. nov. Basionym: Meconopsis
Pt. 2, Nat. Hist. 64: 325. 1896. pseudointegrifolia Prain, Ann. Bot. (Oxford) 20: 353.
1906.
Papaver manasluense (P.A.Egan) Christenh. & Byng,
comb. nov. Basionym: Meconopsis manasluensis Papaver pseudovenustum (G.Taylor) Christenh. & Byng,
P.A.Egan, Phytotaxa 20: 50. 2011. comb. nov. Basionym: Meconopsis pseudovenusta
G.Taylor, Account Gen. Meconopsis: 85. 1934.
Papaver muscicola (Tosh.Yoshida, H.Sun & Boufford)
Christenh. & Byng, comb. nov. Basionym: Papaver psilonomma (Farrer) Christenh. & Byng,
Meconopsis muscicola Tosh.Yoshida, H.Sun & comb. nov. Basionym: Meconopsis psilonomma

71
Farrer, Gard. Chron., ser. 3, 57: 110. 1915. Wilson, Gen. Meconopsis: 188. 2014.
Papaver pulchellum (Tosh.Yoshida, H.Sun & Boufford) Papaver superbum (King ex Prain) Christenh. & Byng,
Christenh. & Byng, comb. nov. Basionym: comb. nov. Basionym: Meconopsis superba King ex
Meconopsis pulchella Tosh.Yoshida, H.Sun & Prain, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 64: 317.
Boufford, Acta Bot. Yunnan. 32: 503. 2010. 1896.
Papaver puniceum (Maxim.) Christenh. & Byng, comb. Papaver taylorii (L.H.J.Williams) Christenh. &
nov. Basionym: Meconopsis punicea Maxim., Fl. Byng, comb. nov. Basionym: Meconopsis taylorii
Tangut. 1: 34. 1889. (Figure 9C) L.H.J.Williams, Trans. Bot. Soc. Edinburgh 41: 348.
Papaver quintuplinervium (Regel) Christenh. & Byng, 1972.
comb. nov. Basionym: Meconopsis quintuplinervia Papaver tibeticum (Grey-Wilson) Christenh. & Byng,
Regel, Gartenflora 25: 291. 1876. comb. nov. Basionym: Meconopsis tibetica Grey-
Papaver racemosum (Maxim.) Christenh. & Byng, Wilson, Bull. Alpine Gard. Soc. Gr. Brit. 74: 222. 2006.
comb. nov. Basionym: Meconopsis racemosa Papaver torquatum (Prain) Christenh. & Byng, comb.
Maxim., Bull. Acad. Imp. Sci. St.-Pétersbourg 28: 310. nov. Basionym: Meconopsis torquata Prain, Ann.
1877. Bot. (Oxford) 20: 355. 1906.
Papaver regium (G.Taylor) Christenh. & Byng, comb. Papaver venustum (Prain) Christenh. & Byng, comb.
nov. Basionym: Meconopsis regia G.Taylor, J. Bot. 67: nov. Basionym: Meconopsis venusta Prain, Bull.
259. 1929 Misc. Inform. Kew 1915: 164. 1915.
Papaver robustum (Hook.f. & Thomson) Christenh. & Papaver violaceum (Kingdon-Ward) Christenh. &
Byng, comb. nov. Basionym: Meconopsis robusta Byng, comb. nov. Basionym: Meconopsis violacea
Hook.f. & Thomson, Fl. Ind. 1: 253. 1855. Kingdon-Ward, Garden 91: 450. 1927.
Papaver roemeria Christenh. & Byng, nom. nov. Papaver wallichii (Hook.) Christenh. & Byng, comb.
Replaced synonym: Chelidonium hybridum L., Sp. nov. Basionym: Meconopsis wallichii Hook., Curtis’s
Pl. 506. 1753, not: Papaver hybridum L. Etymology: Bot. Mag. 78: t. 4668. 1852.
This species is named for Roemeria, a genus now a Papaver wilsonii (Grey-Wilson) Christenh. & Byng,
synonym of Papaver. comb. nov. Basionym: Meconopsis wilsonii Grey-
Papaver rude (Prain) Christenh. & Byng, comb. nov. Wilson, Curtis’s Bot. Mag. 23: 195. 2006.
Basionym: Meconopsis horridula Hook.f. & Thomson Papaver wumungense (K.M.Feng ex C.Y.Wu &
var. rudis Prain, J. Asiat. Soc. Bengal. Pt. 2. Nat. Hist. H.Zhuang) Christenh. & Byng, comb. nov. Basionym:
64: 314. 1896. Meconopsis wumungensis K.M.Feng ex C.Y.Wu &
Papaver sherriffii (G.Taylor) Christenh. & Byng, comb. H.Zhuang, Fl. Yunnan. 2: 33. 1979.
nov. Basionym: Meconopsis sherriffii G.Taylor, New Papaver xiangchengense (R.Li & Z.L.Dao) Christenh.
Fl. & Silva 9: 155. 1937. & Byng, comb. nov. Basionym: Meconopsis
Papaver simikotense (Grey-Wilson) Christenh. & Byng, xiangchengensis R.Li & Z.L.Dao, Novon 22: 180.
comb. nov. Basionym: Meconopsis simikotensis 2012.
Grey-Wilson, Bull. Alpine Gard. Soc. Gr. Brit. 74: 220. Papaver yaoshanense (Tosh.Yoshida, H.Sun &
2006. Boufford) Christenh. & Byng, comb. nov. Basionym:
Papaver sinomaculatum (Grey-Wilson) Christenh. Meconopsis yaoshanensis Tosh.Yoshida, H.Sun &
& Byng, comb. nov. Basionym: Meconopsis Boufford, Acta Bot Yunnan. 34: 148. 2012.
sinomaculata Grey-Wilson, Plantsman n.s., 1: 227. Papaver zangnanense (L.H.Zhou) Christenh. & Byng,
2002. comb. nov. Basionym: Meconopsis zangnanensis
Papaver sinuatum (Prain) Christenh. & Byng, comb. L.H.Zhou, Acta Phytotax. Sin. 17: 112. 1979.
nov. Basionym: Meconopsis sinuata Prain, J. Asiat. Papaver zhongdianense (Grey-Wilson) Christenh.
Soc. Bengal, Pt. 2, Nat. Hist. 64: 314. 1896. & Byng, comb. nov. Basionym: Meconopsis
Papaver smithianum (Hand.-Mazz.) Christenh. & Byng, zhongdianensis Grey-Wilson, Gen. Meconopsis : 258.
comb. nov. Basionym: Cathcartia smithiana Hand.- 2014.
Mazz., Anz. Akad. Wiss. Wien, Math.-Naturwiss. Kl.
60: 182. 1924. 110. Berberidaceae. References: Liu et al. (2002), Wang
Papaver staintonii (Grey-Wilson) Christenh. & Byng, et al. (2007).
comb. nov. Basionym: Meconopsis staintonii Grey- Note: Diphylleia Michx. is embedded in Podophyllum
Wilson, Curtis’s Bot. Mag. 23: 190. 2006. L., which resulted in the acceptance of Dysosma
Papaver sulphureum (Grey-Wilson) Christenh. & Byng, Woodson and Sinopodophyllum T.S.Ying. As this
comb. nov. Basionym: Meconopsis sulphurea Grey- destabalises the traditional concept of Podophyllum
and defies the fact that Diphylleia is morphologically

similar, it is better to merge the three species of demissa Hook.f. & Thomson, Fl. Ind. 1: 23. 1855.
Diphylleia with Podophyllum, keeping the genus in Anemonidium flaccidum (F.Schmidt) Christenh. &
the traditional sense. Byng, comb. nov. Basionym: Anemone flaccida
Podophyllum cymosum (Michx.) Christenh. & Byng, F.Schmidt, Reis. Amur-Land., Bot.: 103. 1868.
comb. nov. Basionym: Diphylleia cymosa Michx., Fl. Anemonidium keiskeanum (T.Ito ex Maxim.) Christenh.
Bor.-Amer. 1: 203. 1803. & Byng, comb. nov. Basionym: Anemone keiskeana
Podophyllum grayi (F.Schmidt) Christenh. & Byng, T. Itô ex Maxim., Bull. Acad. Imp. Sci. St.-Petersbourg,
comb. nov. Basionym: Diphylleia grayi F.Schmidt, sér. 3, 32: 478. 1888.
Reis. Amur-Land., Bot.: 109. 1868. Anemonidium narcissiflorum (L.) Christenh. & Byng,
Podophyllum sinense (H.L.Li) Christenh. & Byng, comb. comb. nov. Basionym: Anemone narcissiflora L., Sp.
nov. Basionym: Diphylleia sinensis H.L.Li, J. Arnold Pl. 1: 542. 1753.
Arbor. 28: 443. 1947. Anemonidium obtusilobum (D.Don) Christenh. &
Podophyllum tsayuense (T.S.Ying) Christenh. & Byng, Byng, comb. nov. Basionym: Anemone obtusiloba
comb. nov. Basionym: Dysosma tsayuensis T.S.Ying, D.Don, Prodr. Fl. Nepal.: 194. 1825. (Figure 9D)
Acta Phytotax. Sin. 17: 20. 1979. Anemonidium rupestre (Wall. ex Hook.f. & Thomson)
Christenh. & Byng, comb. nov. Basionym: Anemone
111. Ranunculaceae. References: Hoot et al. (1994), rupestris Wall. ex Hook.f. & Thomson, Fl. Ind. 1: 21.
Hörandl et al. (2005), Jabbour & Renner (2011, 1855.
2012), Wang et al. (2013), Lehtonen et al. (2016), Anemonidium tenuicaule (Cheeseman) Christenh. &
Jiang et al. (2017), Xiang et al. (2017). Byng, comb. nov. Basionym: Ranunculus tenuicaulis
Note: Anemone L. in the sense of Hoot et al. (1994) forms Cheesman, Trans. & Proc. New Zealand Inst. 17: 235.
a paraphyletic assembly leading up to a monophyletic 1885.
Clematis L. (Lehtonen et al., 2016). The original idea Anemonidium tetrasepalum (Royle) Christenh. &
of treating it as a single large genus resulted from Byng, comb. nov. Basionym: Anemone tetrasepala
using Clematis as the outgroup by Hoot et al. (1994), Royle, Ill. Bot. Himal. Mts. 1: 53. 1834.
when in fact it is part of the ingroup. As merging the Anemonidium trullifolium (Hook.f. & Thomson)
two genera would destabilise the nomenclature too Christenh. & Byng, comb. nov. Anemone trullifolia
much and is unlikely to be followed by the general Hook.f. & Thomson, Fl. Ind. 1: 22. 1855.
public, the genus Anemone is to be segregated Delphinium arenarium (Carlström) Jabbour, comb.
again into Anemone, Anemonidium (Spach) Holub, nov. Basionym: Consolida arenaria Carlström,
Eriocapitella Nakai, Knowltonia Salisb. (here Willdenowia 14: 16. 1984.
expanded to include most Southern Hemisphere
Delphinium coelesyriacum (Mouterde) Jabbour, comb.
Anemone), Hepatica Mill. and Pulsatilla Mill.; the
nov. Basionym: Consolida coelesyriaca Mouterde,
latter two treated in their historical concepts. Jiang
Nouv. Fl. Liban & Syrie 2: 23. 1970.
et al. (2017) produced another phylogenetic study
on Anemoneae, but failed to include Anemone s.s. Delphinium kandaharicum (Iranshahr) Jabbour, comb.
(e.g. Anemone nemorosa L. and related species), nov. Basionym: Consolida kandaharica Iranshahr, Pl.
hence they found that there are only two clades in Syst. Evol. 155: 56. 1987.
Anemone, albeit with weak support. On the basis Delphinium lineolatum (Hub.-Mor. & C.Simon)
of the study by Jiang et al. (2017) no nomenclatural Jabbour, comb. nov. Basionym: Consolida lineolata
changes should be made, but we follow the more Hub.-Mor & C.Simon, Bauhinia 6: 285. 1978.
complete study of Lehtonen et al. (2016) instead. Delphinium lorestanicum (Iranshahr) Jabbour, comb.
Jabbour & Renner (2011) first found Consolida Gray nov. Basionym: Consolida lorestanica Iranshahr, Pl.
and Aconitella Spach embedded inside Delphinium Syst. Evol. 155: 55. 1987.
L. and subsequent studies confirmed the result Delphinium samium (P.H.Davis) Jabbour, comb. nov.
(Jabbour & Renner, 2012; Wang et al., 2013; Xiang Basionym: Consolida samia P.H.Davis, Notes Roy.
et al., 2017). Similarly, Ranunculus L. is expanded to Bot. Gard. Edinburgh 26: 172. 1965.
include Ficaria Haller and Myosurus L., to provide
Delphinium staminosum (P.H.Davis & Sorger) Jabbour,
a stable taxonomy for that genus (Hörandl et al.,
comb. nov. Basionym: Consolida staminosa
2005).
P.H.Davis & Sorger, Notes Roy. Bot. Gard. Edinburgh
Anemonidium coelestinum (Franch.) Christenh. & 40: 89. 1982.
Byng, comb. nov. Basionym: Anemone coelestina
Delphinium stapfianum (P.H.Davis & Sorger) Jabbour,
Franch., Bull. Soc. Bot. France 32: 4. 1885.
comb. nov. Basionym: Consolida stapfiana P.H.Davis
Anemonidium demissum (Hook.f. & Thomson) & Sorger, Notes Roy. Bot. Gard. Edinburgh 40: 89.
Christenh. & Byng, comb. nov. Basionym: Anemone

73
A B
C D
E F
Figure 9: Angiosperms (Eudicots). A, Papaver betonicifolium (Franch.) Christenh. & Byng; B, Papaver przewalskii
Christenh. & Byng; C, Papaver puniceum (Maxim.) Christenh. & Byng; D, Anemonidium obtusilobum (D.Don)
Christenh. & Byng; E, Eriocapitella hupehensis (Lemoine) Christenh. & Byng; and F, Eriocapitella rivularis (Buch.Ham.
ex DC.) Christenh. & Byng. All images © Maarten Christenhusz.

1982. 1: 217. 1817.

Eriocapitella hupehensis (Lemoine) Christenh. & Byng, Knowltonia major (Phil.) Christenh. & Byng, comb.
comb. nov. Basionym: Anemone japonica (Thunb.) nov. Basionym: Barneoudia major Phil., Linnaea 28:
Siebold & Zucc. var. hupehensis Lemoine, Catalogue 609. 1857.
170: 42. 1908. (Figure 9E) Knowltonia mexicana (Kunth) Christenh. & Byng,
Eriocapitella ×hybrida (L.H.Bailey) Christenh. & Byng, comb. nov. Basionym: Anemone mexicana Kunth,
comb. nov. Basionym: Anemone japonica (Thunb.) Nov. Gen. Sp. (quarto ed.) 5: 41. 1817.
Siebold & Zucc. var. hybrida L.H.Bailey, Stand. Knowltonia moorei (Espinosa) Christenh. & Byng,
Cycl. Hort. 1: 286. 1914. (= Eriocapitella vitifolia × comb. nov. Basionym: Anemone moorei Espinosa,
Eriocapitella japonica) Bol. Mus. Nac., Santiago de Chile 18: 26. 1940.
Eriocapitella rivularis (Buch.Ham. ex DC.) Christenh. Knowltonia peruviana (Britton) Christenh. & Byng,
& Byng, comb. nov. Basionym: Anemone rivularis comb. nov. Basionym: Anemone peruviana Britton,
Buch.-Ham. ex DC., Syst. Nat. 1: 211. 1818. (Figure Ann. New York Acad. Sci. 4: 225. 1891.
9F) Knowltonia pulsatilloides Christenh. & Byng, nom.
Eriocapitella rupicola (Cambess.) Christenh. & Byng, nov. Replaced synonym: Atragene capensis L., Sp.
comb. nov. Basionym: Anemone rupicola Cambess., Pl. 1: 543. 1753, not: Knowltonia capensis Huth.
Voy. Inde [Jacquemont] 4(Bot.): 5. 1844. Etymology: This species somewhat resembles a
Eriocapitella tomentosa (Maxim.) Christenh. & Byng, Pulsatilla of the Cape, hence its name.
comb. nov. Basionym: Anemone japonica (Thunb.) Knowltonia sellowii (Pritz.) Christenh. & Byng, comb.
Siebold & Zucc. var. tomentosa Maxim., Fl. Tangut.: nov. Basionym: Anemone sellowii Pritz., Linnaea 15:
7. 1889. 667. 1842.
Knowltonia assisbrasiliana (Kuhlm. & Porto) Ranunculus adamastus Christenh. & Byng, nom. nov.
Christenh. & Byng, comb. nov. Basionym: Anemone Replaced synonym: Myosurus capensis Tamura, Acta
assisbrasiliana Kuhlm. & Porto, Arch. Jard. Bot. Rio Phytotax. Geobot. 44: 27. 1993, not: Ranunculus
de Janeiro 6: 114. 1933. capensis Thunb. Etymology: From Greek αδάμαστος,
Knowltonia balliana (Britton) Christenh. & Byng, indomitable or untamed.
comb. nov. Basionym: Barneoudia balliana Britton, Ranunculus alopecuroides (Greene) Christenh. & Byng,
Ann. New York Acad. Sci. 6: 236. 1891. comb. nov. Basionym: Myosurus alopecuroides
Knowltonia caffra (Harv.) Christenh. & Byng, comb. Greene, Bull. Calif. Acad. Sci. 1: 278. 1885.
nov. Basionym: Anemone caffra Harv., Gen. S. Afr. Ranunculus aotearoanus Christenh. & Byng, nom.
Pl.: 9. 1838. nov. Replaced synonym: Myosurus novae-zelandiae
Knowltonia chilensis (Gay) Christenh. & Byng, comb. W.R.B.Oliv., Trans. & Proc. New Zealand Inst. 56:
nov. Basionym: Barneoudia chilensis Gay, Fl. Chil. 1: 4. 1926, not: Ranunculus novae-zelandiae Petrie.
29. 1845. Etymology: Aotearoa is the Maori name for New
Knowltonia crassifolia (Hook.) Christenh. & Byng, Zealand, where this species is found.
comb. nov. Basionym: Anemone crassifolia Hook., Ranunculus aristatus (Benth.) Christenh. & Byng,
Icon. Pl. 3: t. 257. 1840. comb. nov. Basionym: Myosurus aristatus Benth.,
Knowltonia fanninii (Harv. & Hook.f.) Christenh. & London J. Bot. 6: 459. 1847. Replaced synonym:
Byng, comb. nov. Basionym: Anemone fanninii Harv. Myosurus apetalus Gay, Fl. Chil. 1: 31. 1845, not:
& Hook.f., Bot. Mag. 113: t. 6958. 1887. Ranunculus apetalus Farr
Knowltonia helleborifolia (DC.) Christenh. & Byng, Ranunculus breviscapus (Huth) Christenh. & Byng,
comb. nov. Basionym: Anemone helleborifolia DC., comb. nov. Basionym: Myosurus breviscapus Huth,
Syst. Nat. 1: 211. 1817. Bot. Jahrb. Syst. 16: 285. 1893.
Knowltonia hepaticifolia (Hook.f.) Christenh. & Byng, Ranunculus cauda-muris (Neck.) Christenh. & Byng,
comb. nov. Basionym: Anemone hepaticifolia comb. nov. Basionym: Myosurus cauda-muris Neck.,
Hook.f., Icon. Pl. 1: t. 1. 1837. Delic. Gallo-Belg. 1: 161. 1786.
Knowltonia hootae Christenh. & Byng, nom. nov. Ranunculus cupulatus (S.Wats.) Christenh. & Byng,
Replaced synonym: Anemone rigida Gay, Fl. Chil. 1: comb. nov. Basionym: Myosurus cupulatus S.Watson,
25. 1845, not: Knowltonia rigida Salisb. Etymology: Proc. Amer. Acad. Arts & Sci. 17: 362. 1882.
Named for Sara Hoot, for her contributions to the Ranunculus egglestonii Christenh. & Byng, comb. nov.
understanding of the evolution of Ranunculales. Basionym: Myosurus egglestonii Wooton & Standley,
Knowltonia integrifolia (DC.) Christenh. & Byng, comb. Contr. U.S. Natl. Herb. 16: 123. 1913. Replaced
nov. Basionym: Hepatica integrifolia DC., Syst. Nat. synonym: Myosurus nitidus Eastw., Bull. Torrey Bot.

75
Club 32: 194. 1905, not: Ranunculus nitidus Walter comb. nov. Basionym: Grevillea acuaria F.Muell. ex
Ranunculus heldreichii (H.Lév.) Christenh. & Byng, Benth., Fl. Austral. 5: 421. 1870.
comb. nov. Basionym: Myosurus heldreichii H.Lév., Hakea adenotricha (McGill.) Christenh. & Byng, comb.
Bull. Acad. Int. Géogr. Bot. 11: 296. 1902. nov. Basionym: Grevillea adenotricha McGill., New
Ranunculus musculus Christenh. & Byng, nom. nov. Names Grevillea: 1. 1986.
Replaced synonym: Myosurus tenellus Greene, Hakea adpressa (Olde & Marriott) Christenh. & Byng,
Pittonia 3: 258. 1898, not: Ranunuculus tenellus comb. nov. Basionym: Grevillea adpressa Olde &
Viv., nor: Ranunculus tenellus Nutt. Etymology: From Marriott, Nuytsia 9: 250. 1993.
Latin mus, a mouse, in deminutive, as the plant Hakea agrifolia (A.Cunn. ex R.Br.) Christenh. & Byng,
resembles little mouse tails. comb. nov. Basionym: Grevillea agrifolia A.Cunn. ex
Ranunculus ondine Christenh. & Byng, nom. nov. R.Br., Prodr. Fl. Nov. Holl., Suppl. 1: 24. 1830.
Replaced synonym: Myosurus major Greene, Pittonia Hakea albiflora (C.T.White) Christenh. & Byng, comb.
3: 257. 1898, not: Ranunculus major Dumort. nov. Basionym: Grevillea albiflora C.T.White, Proc.
Etymology: Undine (or Ondine) is a water nymph Roy. Soc. Queensland 55: 79. 1944.
from the alchemical writings of Paracelsus. As this Hakea alpina (Lindl.) Christenh. & Byng, comb. nov.
species is aquatic, the name seemed appropriate. Basionym: Grevillea alpina Lindl., Three Exped.
Ranunculus oz Christenh. & Byng, nom. nov. Replaced Australia [Mitchell] 2: 178, in adnot. 1838.
synonym: Myosurus australis F.Muell. Trans. Philos. Hakea alpivaga (Gand.) Christenh. & Byng, comb. nov.
Soc. Victoria 1: 6. 1854, not: Ranunculus australis Basionym: Grevillea alpivaga Gand., Bull. Soc. Bot.
Eastw. Etymology: As this species is Australian, ‘oz’ France 66: 231, in clavi. 1919.
is a colloquial abbreviation commonly used for the
Hakea althoferorum (Olde & Marriott) Christenh. &
country where this species grows.
Byng, comb. nov. Basionym: Grevillea althoferorum
Ranunculus sessilis (S.Wats.) Christenh. & Byng, comb. Olde & Marriott, Nuytsia 9: 295. 1993.
nov. Basionym: Myosurus sessilis S.Wats., Proc.
Hakea amplexans (F.Muell. ex Benth.) Christenh. &
Amer. Acad. Arts & Sci. 17: 362. 1882.
Byng, comb. nov. Basionym: Grevillea amplexans
Ranunculus tehuelchicus Christenh. & Byng, nom. nov. F.Muell. ex Benth., Fl. Austral. 5: 488. 1870.
Replaced synonym: Myosurus patagonicus Speg.,
Hakea anethifolia (R.Br.) Christenh. & Byng, comb.
Rev. Fac. Agron. Univ. Nac. La Plata 3: 488. 1897, not:
nov. Basionym: Grevillea anethifolia R.Br., Prodr. Fl.
Ranunculus patagonicus Poepp. Etymology: Named
Nov. Holl., Suppl. 1: 21. 1830.
for the Tehuelche people that inhabit Patagonia,
where this species is found. Hakea aneura (McGill.) Christenh. & Byng, comb. nov.
Basionym: Grevillea aneura McGill., New Names
Grevillea: 1. 1986.
115. Proteaceae. Reference: Mast et al. (2015).
Hakea angulata (R.Br.) Christenh. & Byng, comb. nov.
Note: Hakea s.s. and Finschia make Grevillea Basionym: Grevillea angulata R.Br., Prodr. Fl. Nov.
paraphyletic and the two genera are hereby merged. Holl., Suppl. 1: 24. 1830.
Hakea has nomenclatural priority. A full discussion is
Hakea angustiloba (F.Muell.) Christenh. & Byng,
provided in the introduction.
comb. & stat. nov. Basionym: Grevillea ilicifolia var.
Hakea abbreviata (McGill.) Christenh. & Byng, comb. angustiloba F.Muell., Fragm. Phytogr. Austral. 6:
& stat. nov. Basionym: Grevillea nana C.Gardner 212. 1868.
subsp. abbreviata McGill., New Names Grevillea: 10.
Hakea annulifera (F.Muell.) Christenh. & Byng, comb.
1986.
nov. Basionym: Grevillea annulifera F.Muell., Fragm.
Hakea acanthifolia (A.Cunn.) Christenh. & Byng, comb. Phytogr. Austral. 4: 85. 1864. (Figure 10A)
nov. Basionym: Grevillea acanthifolia A.Cunn.,
Hakea anthira Christenh. & M.W.Chase, nom. nov.
Geogr. Mem. New South Wales (Field): 328. 1825.
Replaced synonym: Grevillea uncinulata subsp.
Hakea acerata (McGill.) Christenh. & Byng, comb. nov. florida McGill., New Names Grevillea: 15. 1986, not:
Basionym: Grevillea acerata McGill., New Names Hakea florida R.Br. Etymology: From ανθηρος Greek
Grevillea: 1. 1986. for florid.
Hakea acrobotrya (Meisn.) Christenh. & Byng, comb. Hakea apiciloba (F.Muell.) Christenh. & Byng, comb.
nov. Basionym: Grevillea acrobotrya Meisn., nov. Basionym: Grevillea apiciloba F.Muell., Fragm.
Hooker’s J. Bot. Kew Gard. Misc. 7: 74. 1855. Phytogr. Austral. 10: 45. 1876.
Hakea acropogon (Makinson) Christenh. & Byng, Hakea apiciloba (F.Muell.) Christenh. & Byng, comb.
comb. nov. Basionym: Grevillea acropogon nov. Basionym: Grevillea apiciloba F.Muell., Fragm.
Makinson, Fl. Australia 17A: 496. 2000. Phytogr. Austral. 10: 45. 1876.
Hakea acuaria (F.Muell. ex Benth.) Christenh. & Byng, Hakea aquifolium (Lindl.) Christenh. & Byng, comb.

nov. Basionym: Grevillea aquifolium Lindl., Three F.Muell. ex Benth., Fl. Austral. 5: 436. 1870.
Exped. Australia [Mitchell] 2: 178, in adnot. 1838. Hakea barrettiorum Christenh. & M.W.Chase, nom.
(Figure 10B) nov. Basionym: Grevillea lemanniana Meisn.,
Hakea arachnoidea Christenh. & M.W.Chase, nom. Prodr. (A.P. de Candolle) 14: 366. 1856, not:
nov. Replaced synonym: Embothrium sericeum Hakea lehmanniana Meisn., orth. var. Etymology:
Sm., Spec. Bot. N. Holl. 25. 1793, not: Hakea sericea This species is named for the brothers Matthew
Schrad. & J.C.Wendl. Etymology: This species has and Russell Barrett, for their contribution to the
pink spidery flowers, hence the epithet. (Figure 10C) knowledge of the Australian flora.
Hakea arenaria (R.Br.) Christenh. & Byng, comb. nov. Hakea bathurstensis Christenh. & Byng, nom. nov.
Basionym: Grevillea arenaria R.Br., Trans. Linn. Soc. Replaced synonym: Grevillea divaricata R.Br.,
London, Bot. 10: 172. 1810. Prodr. Fl. Nov. Holl., Suppl. 1: 20. 1830, not: Hakea
Hakea argyrophylla (Meisn.) Christenh. & Byng, comb. divaricata L.A.S.Johnson. Etymology: This species
nov. Basionym: Grevillea argyrophylla Meisn., has only been found only once near Bathurst, New
Hooker’s J. Bot. Kew Gard. Misc. 7: 75. 1855. South Wales.
Hakea aridocapensis Christenh. & M.W.Chase, nom. Hakea batrachioides (F.Muell. ex McGill.) Christenh. &
nov. Replaced synonym: Grevillea baxteri R.Br., Byng, comb. nov. Basionym: Grevillea batrachioides
Prodr. Fl. Nov. Holl., Suppl. 1: 22. 1830, not: Hakea F.Muell. ex McGill., New Names Grevillea: 2. 1986.
baxteri R.Br. Etymology: This species is endemic to Hakea baueri (R.Br.) Christenh. & Byng, comb. nov.
Cape Aride, hence the name. Basionym: Grevillea baueri R.Br., Trans. Linn. Soc.
Hakea armigera (Meisn.) Christenh. & Byng, comb. London, Bot. 10: 173. 1810.
nov. Basionym: Grevillea armigera Meisn., Prodr. [A. Hakea beadleana (McGill.) Christenh. & Byng, comb.
P. de Candolle] 14: 373. 1856. nov. Basionym: Grevillea beadleana McGill., New
Hakea asparagoides (Meisn.) Christenh. & Byng, Names Grevillea: 2. 1986.
comb. nov. Basionym: Grevillea asparagoides Hakea beardiana (McGill.) Christenh. & Byng, comb.
Meisn., Prodr. [A. P. de Candolle] 14: 373. 1856. nov. Basionym: Grevillea beardiana McGill., New
(Figure 10D) Names Grevillea: 2. 1986.
Hakea aspera (R.Br.) Christenh. & Byng, comb. nov. Hakea bedggoodiana (J.H.Willis ex McGill.)
Basionym: Grevillea aspera R.Br., Trans. Linn. Soc. Christenh. & Byng, comb. nov. Basionym: Grevillea
London, Bot. 10: 172. 1810. bedggoodiana J.H.Willis ex McGill., New Names
Hakea asperula (McGill.) Christenh. & Byng, comb. & Grevillea: 3. 1986.
stat. nov. Basionym: Grevillea baueri subsp. asperula Hakea behrii (Schltdl.) Christenh. & Byng, comb. nov.
McGill., New Names Grevillea: 2. 1986. Basionym: Grevillea behrii Schltdl., Linnaea 20: 585.
Hakea aspleniifolia (Knight) Christenh. & Byng, comb. 1847. Replaced synonym: Anadenia ilicifolia R.Br.,
nov. Basionym: Grevillea aspleniifolia Knight, Cult. Trans. Linn. Soc. London, Bot. 10: 167. 1810, not:
Prot.: 120. 1809. Hakea ilicifolia R.Br.
Hakea asteriscosa (Diels ex Diels & E.Pritz.) Christenh. Hakea bemboka (Stajsic & Molyneux) Christenh. &
& Byng, comb. nov. Basionym: Grevillea asteriscosa Byng, comb. nov. Basionym: Grevillea bemboka
Diels ex Diels & E.Pritz., Bot. Jahrb. Syst. 35: 151. Stajsic & Molyneux, Muelleria 22: 28. 2005.
1904. (Figure 10E) Hakea benthamiana (McGill.) Christenh. & Byng,
Hakea aurea (Olde & Mariott) Christenh. & Byng, comb. nov. Basionym: Grevillea benthamiana
comb. nov. Basionym: Grevillea aurea Olde & McGill., New Names Grevillea: 3. 1986.
Marriott, Telopea 5: 407. 1993. Hakea berryana (Ewart & Jean White) Christenh. &
Hakea australis (R.Br.) Christenh. & Byng, comb. nov. Byng, comb. nov. Basionym: Grevillea berryana
Basionym: Grevillea australis R.Br., Trans. Linn. Soc. Ewart & Jean White, Proc. Roy. Soc. Victoria n.s. 22:
London, Bot. 10: 171. 1810. 14. 1909.
Hakea banksii (R.Br.) Christenh. & Byng, comb. nov. Hakea biformis (Meisn.) Christenh. & Byng, comb.
Basionym: Grevillea banksii R.Br., Trans. Linn. Soc. nov. Basionym: Grevillea biformis Meisn., Pl. Preiss.
London, Bot. 10: 176. 1810. (Lehmann) 2: 258. 1848.
Hakea banyabba (Olde & Marriott) Christenh. & Byng, Hakea biternata (Meisn.) Christenh. & Byng, comb.
comb. nov. Basionym: Grevillea banyabba Olde & nov. Basionym: Grevillea biternata Meisn., Pl. Preiss.
Marriott, Telopea 5: 719. 1994. (Lehmann) 1: 549. 1845.
Hakea barklyana (F.Muell. ex Benth.) Christenh. & Hakea brachystachya (Meisn.) Christenh. & Byng,
Byng, comb. nov. Basionym: Grevillea barklyana comb. nov. Basionym: Grevillea brachystachya

77
Meisn., Pl. Preiss. (Lehmann) 2: 254. 1848. large evergreen leaves, like a Camellia (Theaceae).
Hakea brachystylis (Meisn.) Christenh. & Byng, comb. Hakea candelabroides (C.A.Gardner) Christenh.
nov. Basionym: Grevillea brachystylis Meisn., Pl. & Byng, comb. nov. Basionym: Grevillea
Preiss. (Lehmann) 1: 538. 1845. candelabroides C.A.Gardner, J. Roy. Soc. Western
Hakea bracteosa (Meisn.) Christenh. & Byng, comb. Australia 47: 56. 1964.
nov. Basionym: Grevillea bracteosa Meisn., Pl. Hakea candicans (C.A.Gardner) Christenh. &
Preiss. (Lehmann) 2: 254. 1848. Byng, comb. nov. Basionym: Grevillea candicans
Hakea brevifolia (F.Muell. ex Benth.) Christenh. & C.A.Gardner, J. Roy. Soc. Western Australia 27: 170.
Byng, comb. nov. Basionym: Grevillea brevifolia 1943.
F.Muell. ex Benth., Fl. Austral. 5: 467. 1870. Hakea capitellata (Meisn.) Christenh. & Byng, comb.
Hakea brevis (Olde & Marriott) Christenh. & Byng, nov. Basionym: Grevillea capitellata Meisn., Prodr.
comb. nov. Basionym: Grevillea brevis Olde & (A.P. de Candolle) 14: 356. 1856.
Marriott, Telopea 5: 410. 1993. Hakea carduifolia (A.Cunn. ex Benth.) Christenh. &
Hakea brindabella (Stajsic) Christenh. & Byng, comb. Byng, comb. nov. Basionym: Grevilea carduifolia
& stat. nov. Basionym: Grevillea victoriae subsp. A.Cunn. ex Benth., Fl. Austral. 5: 456. 1870.
brindabella Stajsic, Muelleria 28: 19. 2010. Replaced synonym: Grevillea cunninghamii R.Br.,
Prodr. Fl. Nov. Holl., Suppl. 1: 23, 1830. not: Hakea
Hakea bronwenae (Keighery) Christenh. & Byng, comb.
cunninghamii R.Br.
nov. Basionym: Grevillea bronwenae Keighery,
Nuytsia 7: 128. 1990. Hakea celata (Molyneux) Christenh. & Byng, comb.
nov. Basionym: Grevillea celata Molyneux, Muelleria
Hakea bundera (Keighery) Christenh. & Byng, comb.
8: 311. 1995.
& stat. nov. Basionym: Grevillea variifolia subsp.
bundera Keighery, Nuytsia 12: 294. 1998. Hakea centristigma (McGill.) Christenh. & Byng,
comb. nov. Basionym: Grevillea drummondii subsp.
Hakea burrowa (Molyneux & Forrester) Christenh.
centristigma McGill., New Names Grevillea: 5. 1986.
& Byng, comb. nov. Basionym: Grevillea burrowa
Molyneux & Forrester, Muelleria 34: 51. 2015. Hakea ceratocarpa (Diels ex Diels & E.Pritz.) Christenh.
& Byng, comb. nov. Basionym: Grevillea ceratocarpa
Hakea buxifolia (Sm.) Christenh. & Byng, comb. nov.
Diels ex Diels & E.Pritz., Bot. Jahrb. Syst. 35: 157.
Basionym: Embothrium buxifolium Sm., Spec. Bot. N.
1904.
Holl. 29. 1793.
Hakea cheilocarpa (Makinson) Christenh. & Byng,
Hakea byrnesii (McGill.) Christenh. & Byng, comb. nov.
comb. nov. Basionym: Grevillea cheilocarpa
Basionym: Grevillea byrnesii McGill., New Names
Makinson, Fl. Australia 17A: 505. 2000.
Grevillea: 3. 1986.
Hakea chloroxantha (Diels) Christenh. & Byng, comb.
Hakea cagiana (McGill.) Christenh. & Byng, comb. nov.
nov. Basionym: Finschia chloroxantha Diels, Bot.
Basionym: Grevillea cagiana McGill., New Names
Jahrb. Syst. 54: 204. 1916.
Grevillea: 3. 1986.
Hakea christineae (McGill.) Christenh. & Byng, comb.
Hakea calcicola (A.S.George) Christenh. & Byng, comb.
nov. Basionym: Grevillea christineae McGill., New
nov. Basionym: Grevillea calcicola A.S.George, J. Roy.
Names Grevillea: 4. 1986.
Soc. Western Australia 50: 97. 1968.
Hakea chrysophaea (F.Muell. ex Meisn.) Christenh. &
Hakea caleyana (Olde) Christenh. & Byng, comb.
Byng, comb. nov. Basionym: Grevillea chrysophaea
& stat. nov. Basionym: Grevillea laurifolia subsp.
F.Muell. ex Meisn., Linnaea 26: 357. 1854.
caleyana Olde, Telopea 18: 104. 2015.
Hakea cincta (Molyneux & Stajsic) Christenh. & Byng,
Hakea caleyi (R.Br.) Christenh. & Byng, comb. nov.
comb. & stat. nov. Basionym: Grevillea miqueliana
Basionym: Grevillea caleyi R.Br., Prodr. Fl. Nov. Holl.,
subsp. cincta Molyneux & Stajsic. Muelleria 22: 50.
Suppl. 1: 22. 1830.
2005.
Hakea calliantha (Makinson & Olde) Christenh. &
Hakea cirsiifolia (Meisn.) Christenh. & Byng, comb.
Byng, comb. nov. Basionym: Grevillea calliantha
nov. Basionym: Grevillea cirsiifolia Meisn., Pl. Preiss.
Makinson & Olde, Telopea 4: 351. 1991.
(Lehmann) 2: 253. 1848.
Hakea callichlaena (Molyneux & Stajsic) Christenh. &
Hakea coccinea (Meisn.) Christenh. & Byng, comb.
Byng, comb. nov. Basionym: Grevillea callichlaena
nov. Basionym: Grevillea coccinea Meisn., Hooker’s
Molyneux & Stajsic, Muelleria 22: 38. 2005.
J. Bot. Kew Gard. Misc. 7: 76. 1855.
Hakea camellia Christenh. & Byng, nom. nov. Replaced
Hakea concinna (R.Br.) Christenh. & Byng, comb. nov.
synonym: Grevillea latifolia C.Gardner, Bull. Woods
Basionym: Grevillea concinna R.Br., Trans. Linn. Soc.
Forests Dept., W. Australia 32: 43. 1923, not: Hakea
London, Bot. 10: 172. 1810.
latifolia hort. ex Colla. Etymology: This species has
Hakea confertifolia (F.Muell.) Christenh. & Byng,

comb. nov. Basionym: Grevillea confertifolia delta McGill., New Names Grevillea: 15. 1986.
F.Muell., Trans. Phil. Soc. Victoria 1: 22. 1854. Hakea depauperata (R.Br.) Christenh. & Byng, comb.
Hakea corrugata (Olde & Marriott) Christenh. & Byng, nov. Basionym: Grevillea depauperata R.Br., Prodr.
comb. nov. Basionym: Grevillea corrugata Olde & Fl. Nov. Holl., Suppl. 1: 21. 1930.
Marriott, Nuytsia 9: 247. 1993. Hakea deurrens (Ewart) Christenh. & Byng, comb. nov.
Hakea cravenii (Makinson) Christenh. & Byng, comb. Basionym: Grevillea decurrens Ewart in Ewart &
nov. Basionym: Grevillea cravenii Makinson, Fl. Davies, Fl. Northern Territory: 83. 1917.
Australia 17A: 494. 2000. Hakea didymobotrya (Meisn.) Christenh. & Byng,
Hakea crithmifolia (R.Br.) Christenh. & Byng, comb. comb. nov. Basionym: Grevillea didymobotrya
nov. Basionym: Grevillea crithmifolia R.Br., Prodr. Fl. Meisn., Prodr. [A. P. de Candolle] 14: 386. 1856.
Nov. Holl., Suppl. 1: 23. 1830. Hakea didymochiton (Makinson) Christenh. & Byng,
Hakea crowleyae (Olde & Marriott) Christenh. & Byng, comb. & stat. nov. Basionym: Grevillea irrasa subsp.
comb. nov. Basionym: Grevillea crowleyae Olde & didymochiton Makinson, Fl. Australia 17A: 503.
Marriott, Nuytsia 9: 271. 1993. 2000.
Hakea ctenophylla (Meisn.) Christenh. & Byng, comb. Hakea dielsiana (C.A.Gardner) Christenh. &
nov. Basionym: Grevillea ctenophylla Meisn., Prodr. Byng, comb. nov. Basionym: Grevillea dielsiana
[A. P. de Candolle] 14: 372. 1856. Replaced synonym C.A.Gardner, J. Roy. Soc. Western Australia 27: 169.
Grevillea pectinata R.Br., Prodr. Fl. Nov. Holl., Suppl. 1943.
1: 23. 1830, not: Hakea pectinata Dum.Cours. Hakea diffusa (Sieber ex Spreng.) Christenh. & Byng,
Hakea cullenii (Olde & Marriott) Christenh. & Byng, comb. nov. Basionym: Grevillea diffusa Sieber ex
comb. & stat. nov. Basionym: Grevillea obliquistigma Spreng, Cur. Post. Syst. Veg. 4(2): 46. 1827.
subsp. cullenii Olde & Marriott, Grevillea Book 1: 1. Hakea digitata (F.Muell.) Christenh. & Byng, comb. &
1994. stat. nov. Basionym: Grevillea apiciloba var. digitata
Hakea curviloba (McGill.) Christenh. & Byng, comb. F.Muell., Fragm. Phytogr. Austral. 10: 46. 1876.
nov. Basionym: Grevillea curviloba McGill., New Hakea dilatata (R.Br.) Christenh. & Byng, comb. nov.
Names Grevillea: 4. 1986. Basionym: Grevillea ilicifolia var. dilatata R.Br., Prodr.
Hakea cygnopotamica Christenh. & M.W.Chase, nom. Fl. Nov. Holl., Suppl. 1: 21. 1830.
nov. Replaced synonym: Anadenia tenuiflora Lindl., Hakea dimidiata (F.Muell.) Christenh. & Byng, comb.
Sketch Veg. Swan R. 31. 1839, not: Hakea tenuiflora nov. Basionym: Grevillea dimidiata F.Muell., Fragm.
Dum.Cours. Etymology: This species is named for Phytogr. Austral. 3: 146. 1863.
the Swan River in Perth, from where this species was Hakea diminuta (L.A.S.Johnson) Christenh. &
described. Byng, comb. nov. Basionym: Grevillea diminuta
Hakea cymbiformis (Olde & Marriott) Christenh. L.A.S.Johnson, Contr. New South Wales Natl. Herb.
& Byng, comb. & stat. nov. Basionym: Grevillea 3: 95. 1962.
biformis subsp. cymbiformis Olde & Marriott. Hakea dimorpha (F.Muell.) Christenh. & Byng, comb.
Grevillea Book 1: 176. 1994. nov. Basionym: Grevillea dimorpha F.Muell., Trans.
Hakea cyranostigma (McGill.) Christenh. & Byng, Phil. Soc. Victoria 1: 21. 1854.
comb. nov. Basionym: Grevillea cyranostigma Hakea disjuncta (F.Muell.) Christenh. & Byng, comb.
McGill., Telopea 1: 20. 1975. nov. Basionym: Grevillea disjuncta F.Muell., Fragm.
Hakea dasycarpa (McGill.) Christenh. & Byng, comb. Phytogr. Austral. 6: 206. 1868.
& stat. nov. Basionym: Grevillea dryandri subsp. Hakea diversifolia (Meisn.) Christenh. & Byng, comb.
dasycarpa McGill., New Names Grevillea: 5. 1986. nov. Basionym: Grevillea diversifolia Meisn., Pl.
Hakea decipiens (McGill.) Christenh. & Byng, comb. Preiss. (Lehmann) 1: 547. 1845.
nov. Basionym: Grevillea decipiens McGill., New Hakea dixoniana Christenh. & M.W.Chase, nom. nov.
Names Grevillea: 4. 1986. Replaced synonym: Grevillea preissii Meisn., Pl.
Hakea decora (Domin) Christenh. & Byng, comb. nov. Preiss. (Lehmann) 1: 543. 1845, not: Hakea preissii
Basionym: Grevillea decora Domin, Biblioth. Bot. Meisn. Etymology: This species is named in honour of
22(89): 589. 1921. Kingsley Dixon, for his contributions to restauration
Hakea deflexa (F.Muell.) Christenh. & Byng, comb. nov. ecology and conservation of the Australian flora.
Basionym: Grevillea deflexa F.Muell., Australasian Hakea dolichopoda (McGill.) Christenh. & Byng, comb.
Chemist and Druggist 5: 72. 1883. & stat. nov. Basionym: Grevillea disjuncta subsp.
Hakea delta (McGill.) Christenh. & Byng, comb. & stat. dolichopoda McGill., New Names Grevillea: 5. 1986.
nov. Basionym: Grevillea thelemanniana subsp. Hakea donaldiana (Kenneally) Christenh. & Byng,
comb. nov. Basionym: Grevillea donaldiana, Western

79
A B
C D
E F
Figure 10: Angiosperms (Eudicots). A, Hakea annulifera (F.Muell.) Christenh. & Byng; B, Hakea aquifolium (Lindl.)
Christenh. & Byng; C, Hakea arachnoidea Christenh. & M.W.Chase; D, Hakea asparagoides (Meisn.) Christenh. &
Byng; E, Hakea asteriscosa (Diels ex Diels & E.Pritz.) Christenh. & Byng; and F, Hakea dryandroides (C.A.Gardner)
Christenh. & Byng. All images © Maarten Christenhusz.

Australian Naturalist 17: 113. 1988. Nov. Holl.: 20. 1830, not: Hakea lanigera Ten.
Hakea dryandri (R.Br.) Christenh. & Byng, comb. nov. Hakea eriobotrya (F.Muell.) Christenh. & Byng, comb.
Basionym: Grevillea dryandri R.Br., Trans. Linn. Soc. nov. Basionym: Grevillea eriobotrya F.Muell., Fragm.
London, Bot. 10: 175. 1910. Phytogr. Austral. 10: 44. 1876.
Hakea dryandroides (C.A.Gardner) Christenh. & Hakea eriostachya (Lindl.) Christenh. & Byng, comb.
Byng, comb. nov. Basionym: Grevillea dryandroides nov. Basionym: Grevillea eriostachya Lindl., Sketch
C.A.Gardner, J. Roy. Soc. Western Australia 19: 81. Veg. Swan R.: 36. 1840.
1934. (Figure 10F) Hakea eryngioides (Benth.) Christenh. & Byng, comb.
Hakea dryophylla (N.A.Wakef.) Christenh. & Byng, nov. Basionym: Grevillea eryngioides Benth., Fl.
comb. nov. Basionym: Grevillea dryophylla Austral. 5: 476. 1870.
N.A.Wakef., Victorian Naturalist 73: 74. 1956. Hakea erythroclada (W.Fitzg.) Christenh. & Byng,
Hakea dubia (Domin) Christenh. & Byng, comb. & comb. nov. Basionym: Grevillea erythroclada
stat. nov. Basionym: Grevillea fasciculata var. dubia W.Fitzg., J. Roy. Soc. Western Australia 3: 132. 1918.
Domin, Mem. Soc. Sci. Bohème (Věstn. Král. České Hakea evanescens (Olde & Marriott) Christenh. &
Společn. Nauk. Tř. Mat.-Přír.) 2: 10. 1923. Replaced Byng, comb. nov. Basionym: Grevillea evanescens
synonym: Grevillea crassifolia Domin, Mem. Soc. Sci. Olde & Marriott. Grevillea Book 1: 181. 1994.
Bohème (Věstn. Král. České Společn. Nauk. Tř. Mat.- Hakea evansiana (MacKee) Christenh. & Byng, comb.
Přír.) 2: 10. 1923, not: Hakea crassifolia Meisn. nov. Basionym: Grevillea evansiana MacKee, Proc.
Hakea dumetorum (Meisn.) Christenh. & Byng, comb. Linn. Soc. New South Wales 78: 49. 1953.
nov. Basionym: Grevillea dumetorum Meisn., Prodr. Hakea excelsior (Diels ex Diels & E.Pritz.) Christenh. &
[A. P. de Candolle] 14: 378. 1856. Byng, comb. nov. Basionym: Grevillea excelsior Diels
Hakea dunlopii (Makinson) Christenh. & Byng, comb. ex E.Pritz., Bot. Jahrb. Syst. 35: 151. 1904.
nov. Basionym: Grevillea dunlopii Makinson, Fl. Hakea exposita (Olde & Marriott) Christenh. & Byng,
Australia 17A: 494. 2000. comb. nov. Basionym: Grevillea exposita Olde &
Hakea ecorniculata (Olde & Marriott) Christenh. Marriot, Grevillea Book 1: 181. 1994.
& Byng, comb. & stat. nov. Basionym: Grevillea Hakea extorris (S.Moore) Christenh. & Byng, comb.
buxifolia subsp. ecorniculata Olde & Marriott, nov. Basionym: Grevillea extorris S.Moore, J. Linn.
Telopea 5: 709. 1994. Soc., Bot. 34: 221. 1899.
Hakea elbertii (Sleumer) Christenh. & Byng, comb. Hakea exul (Lindl.) Christenh. & Byng, comb. nov.
nov. Basionym: Grevillea elbertii Sleumer, Blumea Basionym: Grevillea exul Lindl., J. Hort. Soc. London
8: 2. 1955. 6: 273. 1851.
Hakea elliotii (Olde & Marriott) Christenh. & Byng, Hakea fasciculata (R.Br.) Christenh. & Byng, comb.
comb. & stat. nov. Basionym: Grevillea insignis nov. Basionym: Grevillea fasciculata R.Br., Prodr. Fl.
subsp. elliotii Olde & Marriott, Nuytsia 9: 285. 1993. Nov. Holl., Suppl. 1: 20. 1830.
Hakea elongata (Olde & Marriott) Christenh. & Byng, Hakea fastigiata (Olde & Marriott) Christenh. & Byng,
comb. nov. Basionym: Grevillea elongata Olde & comb. nov. Basionym: Grevillea fastigiata Olde &
Marriott, Grevillea Book 1: 175. 1994. Marriott, Grevillea Book 1: 186. 1994.
Hakea endlicheriana (Meisn.) Christenh. & Byng, Hakea fecunda (Makinson) Christenh. & Byng, comb.
comb. nov. Basionym: Grevillea endlicheriana & stat. nov. Basionym: Grevillea obtusiflora subsp.
Meisn., Pl. Preiss. (Lehmann) 1: 546. 1845. fecunda Makinson, Telopea 7: 144. 1997.
Hakea epicroca (Stajsic & Molyneux) Christenh. & Hakea ferricola (Keighery) Christenh. & Byng, comb. &
Byng, comb. nov. Basionym: Grevillea epicroca stat. nov. Basionym: Grevillea manglesioides subsp.
Stajsic & Molyneux, Fl. Australia 17A: 502. 2000. ferricola Keighery, Fl. Australia 17A: 497. 2000.
Hakea erectiloba (F.Muell.) Christenh. & Byng, comb. Hakea ferruginiflora (C.T.White) Christenh. & Byng,
nov. Basionym: Grevillea erectiloba F.Muell., Fragm. comb. nov. Basionym: Finschia ferruginiflora
Phytogr. Austral. 1: 44. 1858. C.T.White, Pacific Sci. 3: 190. 1949.
Hakea eremophila (Diels ex Fiels & E.Pritz.) Christenh. Hakea fililoba (McGill.) Christenh. & Byng, comb. &
& Byng, comb. & stat. nov. Basionym: Grevillea stat. nov. Basionym: Grevillea thelemanniana subsp.
integrifolia var. eremophila Diels ex Diels & E.Pritz., fililoba McGill., New Names Grevillea: 15. 1986.
Bot. Jahrb. Syst. 35: 156. 1904.
Hakea filipendula (McGill.) Christenh. & Byng, comb.
Hakea ericifolia (R.Br.) Christenh. & Byng, comb. nov. Basionym: Grevillea diffusa subsp. filipendula
nov. Basionym: Grevillea ericifolia R.Br., Prodr. Fl. McGill., New Names Grevillea: 4. 1986.
Nov. Holl., Suppl. 1: 20. 1830. Replaced synonym:
Hakea filix Christenh. & Byng, nom. nov. Replaced
Grevillea lanigera A.Cunn. ex R.Br., Suppl. Prodr. Fl.

81
synonym: Grevillea longifolia R.Br., Suppl. Prodr. nov. Replaced synonym: Grevillea commutata
Fl. Nov. Holl.: 22. 1830, not: Hakea longifolia Dum. F.Muell., Fragm. Phytogr. Austral. 6: 207. 1868, not:
Cours. Etymology: This species is called ‘fern-leaf Hakea commutata F.Muell. Etymology: This species
spider flower’, hence filix, meaning fern. is found near Geraldton, Western Australia, hence
Hakea fistulosa (A.S.George) Christenh. & Byng, comb. the name.
nov. Basionym: Grevillea fistulosa A.S.George, Hakea gillivrayi (Hook.) Christenh. & Byng, comb. nov.
Nuytsia 1: 371. 1974. Basionym: Grevillea gillivrayi Hook., Hooker’s J. Bot.
Hakea fitzgeraldii Christenh. & M.W.Chase, nom. nov. Kew Gard. Misc. 6: 358. 1854. (Figure 11A)
Replaced synonym: Grevillea stenophylla W.Fitzg., J. Hakea glabrescens (Olde & Marriott) Christenh. &
W. Austral. Nat. Hist. Soc. 2: 30. 1905, not: Hakea Byng, comb. nov. Basionym: Grevillea glabrescens
stenophylla A.Cunn. ex R.Br. Etymology: This species Olde & Marriott, Telopea 5: 406. 1993.
is named for William Fitzgerald, who first described Hakea globosa (C.A.Gardner) Christenh. & Byng, comb.
it. nov. Basionym: Grevillea globosa C.A.Gardner, J.
Hakea flexuosa (Lindl.) Christenh. & Byng, comb. nov. Roy. Soc. Western Australia 47: 55. 1964.
Basionym: Anadenia flexuosa Lindl., Sketch Veg. Hakea glossadenia (McGill.) Christenh. & Byng, comb.
Swan R.: 31. 1839. nov. Basionym: Grevillea glossadenia McGill.,
Hakea floribunda (R.Br.) Christenh. & Byng, comb. nov. Telopea 1: 21. 1975.
Basionym: Grevillea floribunda R.Br., Prodr. Fl. Nov. Hakea goodii (R.Br.) Christenh. & Byng, comb. nov.
Holl., Suppl. 1: 19. 1830. Basionym: Grevillea goodii R.Br., Trans. Linn. Soc.
Hakea floripendula (R.V.Sm.) Christenh. & Byng, comb. London, Bot. 10: 174. 1910.
nov. Basionym: Grevillea floripendula R.V.Sm., Hakea gordoniana (C.A.Gardner) Christenh. &
Muelleria 4: 423. 1981. Byng, comb. nov. Basionym: Grevillea gordoniana
Hakea fluviatilis (Makinson) Christenh. & Byng, comb. C.A.Gardner, J. Roy. Soc. Western Australia 47: 56.
& stat. nov. Basionym: Grevillea neurophylla subsp. 1964.
fluviatilis Makinson, Fl. Australia 17A: 501. 2000. Hakea grandis (Keighery) Christenh. & Byng, comb. &
Hakea formosa (McGill.) Christenh. & Byng, comb. stat. nov. Basionym: Grevillea brachystylis subsp.
nov. Basionym: Grevillea formosa McGill., New grandis Keighery, Western Australian Naturalist 27:
Names Grevillea: 6. 1986. 12. 2009.
Hakea fuchsioides Christenh. & M.W.Chase, nom. Hakea granitica Christenh. & M.W.Chase, nom. nov.
nov. Replaced synonym: Grevillea bipinnatifida Replaced synonym: Grevillea neurophylla Gand.,
R.Br., Prodr. Fl. Nov. Holl., Suppl. 1: 23. 1830, not: Bull. Soc. Bot. France 66: 231. 1919, not: Hakea
Hakea bipinnatifida R.Br. Etymology: This species neurophylla Meisn. Etymology: This is commonly
resembles the genus Fuchsia (Onagraceae). called ‘granite grevillea’.
Hakea fulgens (C.A.Gardner) Christenh. & Byng, comb. Hakea granulifera (McGill.) Christenh. & Byng, comb.
nov. Basionym: Grevillea fulgens C.A.Gardner, J. Roy. & stat. nov. Basionym: Grevillea obtusiflora subsp.
Soc. Western Australia 47: 55. 1964. granulifera McGill., New Names Grevillea: 11. 1986.
Hakea funicularis (Olde & Marriott) Christenh. & Byng, Hakea granulosa (McGill.) Christenh. & Byng, comb.
comb. & stat. nov. Basionym: Grevillea Grevillea nov. Basionym: Grevillea granulosa McGill., New
obliquistigma subsp. funicularis Olde & Marriott, Names Grevillea: 7. 1986.
Grevillea Book 1: 180. 1994. Hakea grevilleoides Christenh. & M.W.Chase, nom.
Hakea fuscolutea (Keighery) Christenh. & Byng, comb. nov. Replaced synonym: Grevillea hookeriana
nov. Basionym: Grevillea fuscolutea Keighery, Meisn., Pl. Preiss. (Lehmann) 1: 546. 1845, not:
Nuytsia 8: 228. 1992. Hakea hookeriana Meisn., nor: Hakea pritzelii Diels.
Hakea gariwerdensis (Makinson) Christenh. & Byng, Etymology: This species is a ‘typical’ toothbrush
comb. nov. Basionym: Grevillea gariwerdensis grevillea, hence the name.
Makinson, Fl. Australia 17A: 501. 2000. Hakea guthrieana (Olde & Marriott) Christenh. &
Hakea ×gaudichaudii (R.Br. ex Gaudich.) Christenh. & Byng, comb. nov. Basionym: Grevillea guthrieana
Byng, comb. nov. Basionym: Grevillea gaudichaudii Olde & Marriott, Telopea 5: 731. 1994.
R.Br. ex Gaudich., Voy. Uranie, Bot.: 443. 1829 (= H. Hakea hakeoides (Meisn.) Christenh. & Byng, comb.
acanthifolia × H. laurifolia). nov. Basionym: Grevillea hakeoides Meisn., Pl.
Hakea georgeana (McGill.) Christenh. & Byng, comb. Preiss. (Lehmann) 2: 252. 1848.
nov. Basionym: Grevillea georgeana McGill., New Hakea halmaturina (Tate) Christenh. & Byng, comb.
Names Grevillea: 6. 1986. nov. Basionym: Grevillea halmaturina Tate, Handb.
Hakea geraldtonica Christenh. & M.W.Chase, nom.

A B
C D
Figure 11: Angiosperms (Eudicots). A, Hakea gillivrayi (Hook.) Christenh. & Byng; B, Hakea lavandulacea (Schltdl.)
Christenh. & Byng; C, Hakea plurijuga (F.Muell.) Christenh. & Byng; D, Hakea robusta (A.Cunn. ex R.Br.) Christenh. &
M.W.Chase; E, Hakea rosea Christenh. & M.W.Chase; and F, Hakea saccata (Benth.) Christenh. & Byng. All images ©
Maarten Christenhusz.

83
Fl. Extratrop. South Australia: 83, 228. 1890. nov. Basionym: Grevillea humilis Makinson, Fl.
Hakea haplantha (F.Muell.) Christenh. & Byng, comb. Australia 17A: 499. 2000.
nov. Basionym: Grevillea haplantha F.Muell. ex Hakea hypargyrea (F.Muell.) Christenh. & Byng, comb.
Benth., Fl. Austral. 5: 451. 1870. & stat. nov. Basionym: Grevillea ramosissima var.
Hakea heliosperma (R.Br.) Christenh. & Byng, comb. hypargyrea F.Muell., Fragm. Phytogr. Austral. 8: 150.
nov. Basionym: Grevillea heliosperma R.Br., Trans. 1874.
Linn. Soc. London, Bot. 10: 176. 1810. Hakea iaspicula (McGill.) Christenh. & Byng, comb.
Hakea helmsiae (F.M.Bailey) Christenh. & Byng, comb. nov. Basionym: Grevillea iaspicula McGill., New
nov. Basionym: Grevillea helmsiae F.M.Bailey, Names Grevillea: 7. 1986.
Queensland Agric. J. 4: 195. 1899. Hakea ilex Christenh. & M.W.Chase, nom. nov. Replaced
Hakea hilliana (F.Muell.) Christenh. & Byng, comb. synonym: Grevillea longicuspis McGill., New Names
nov. Basionym: Grevillea hilliana F.Muell., Trans. Grevillea: 9. 1986, not: Hakea longicuspis Hook.
Phil. Inst. Victoria 2: 72. 1857. Etymology: This species has a holly-like leaf, hence
the name.
Hakea hirsuta (Olde & Marriott) Christenh. & Byng,
comb. & stat. nov. Basionym: Grevillea dryandroides Hakea imberbis (Makinson) Christenh. & Byng, comb.
subsp. hirsuta Olde & Marriott, Nuytsia 9: 270. 1993. nov. Basionym: Grevillea imberbis Makinson, Fl.
Australia 17A: 501. 2000.
Hakea hirtella (Benth.) Christenh. & Byng, comb. &
stat. nov. Basionym: Grevillea pinaster var. hirtella Hakea inconspicua (Diels ex Diels & E.Pritz.) Christenh.
Benth., Fl. Austral. 5: 427. 1870. & Byng, comb. nov. Basionym: Grevillea inconspicua
Diels ex Diels & E.Pritz., Bot. Jahrb. Syst. 35: 153.
Hakea hislopii (Olde & Marriott) Christenh. & Byng,
1904.
comb. nov. Basionym: Grevillea hislopii Olde &
Marriott, Nuytsia 18: 224. 2008. Hakea incurva (Diels) Christenh. & Byng, comb. & stat.
nov. Basionym: Grevillea integrifolia var. incurva
Hakea hockingsii (Molyneux & Olde) Christenh. &
Diels, Bot. Jahrb. Syst. 35: 157. 1904.
Byng, comb. nov. Basionym: Grevillea hockingsii
Molyneux & Olde, Telopea 5: 784. 1994. Hakea infecunda (McGill.) Christenh. & Byng, comb.
nov. Basionym: Grevillea infecunda McGill., New
Hakea hodgei (Olde & Marriott) Christenh. & Byng,
Names Grevillea: 7. 1986.
comb. nov. Basionym: Grevillea hodgei Olde &
Marriott, Grevillea Book 1: 185. 1994. Hakea infundibularis (A.S.George) Christenh. & Byng,
comb. nov. Basionym: Grevillea infundibularis
Hakea holleae Christenh. & Byng, nom. nov. Replaced
A.S.George, Nuytsia 1: 371. 1974.
synonym: Grevillea pinifolia Meisn., Prodr. [A. P. de
Candolle] 14: 350. 1856, not: Hakea pinifolia Knight. Hakea insignis (Kippist ex Meisn.) Christenh. & Byng,
Etymology: This species has needle-like leaves like comb. nov. Basionym: Grevillea insignis Kippist ex
the spindle that fell into the well of Mother Holle Meisn., Hooker’s J. Bot. Kew Gard. Misc. 7: 76. 1855.
(or Frau Hulda), from an old German fairytale, Hakea integrifolia (Endl.) Christenh. & Byng, comb.
popuralised by the brothers Grimm in the 19th nov. Basionym: Anadenia integrifolia Endl., Ann.
Century. Wien. Mus. II, 209: 21. 1839.
Hakea hopperiana Christenh. & M.W.Chase, nom. Hakea involucrata (A.S.George) Christenh. & Byng,
nov. Replaced synonym: Grevillea drummondii comb. nov. Basionym: Grevillea involucrata
Meisn., Pl. Preiss. (Lehmann) 1: 536, 1845, not: A.S.George, Nuytsia 1: 372. 1974.
Hakea drummondii G.Don ex Meisn. Etymology: Hakea involuta (McGill.) Christenh. & Byng, comb. &
This species is named in honour of Stephen Hopper, stat. nov. Basionym: Grevillea didymobotrya subsp.
for his contributions on conservation and flora of involuta McGill., New Names Grevillea: 4. 1986.
Western Australia. Hakea irrasa (Makinson) Christenh. & Byng, comb.
Hakea howatharra (Olde & Marriott) Christenh. nov. Basionym: Grevillea irrasa Makinson, Fl.
& Byng, comb. & stat. nov. Basionym: Grevillea Australia 17A: 503. 2000.
bracteosa subsp. howatharra Olde & Marriott, Hakea jephcottii (J.H.Willis) Christenh. & Byng, comb.
Nuytsia 18: 232. 2008. nov. Basionym: Grevillea jephcottii J.H.Willis,
Hakea huegelii (Meisn.) Christenh. & Byng, comb. Muelleria 1: 117. 1967.
nov. Basionym: Grevillea huegelii Meisn., Pl. Preiss. Hakea johnsonii (McGill.) Christenh. & Byng, comb.
(Lehmann) 1: 543. 1845. nov. Basionym: Grevillea johnsonii McGill., Telopea
Hakea humifusa (Olde & Marriott) Christenh. & Byng, 1: 22. 1975.
comb. nov. Basionym: Grevillea humifusa Olde & Hakea juncifolia (Hook.) Christenh. & Byng, comb.
Marriott, Grevillea Book 1: 182 1994. nov. Basionym: Grevillea juncifolia Hook., J. Exp.
Hakea humilis (Makinson) Christenh. & Byng, comb.

Inter. Trop. Australia (Mitchell): 341. 1848. Grevillea Book 1: 175. 1994.
Hakea juniperina (R.Br.) Christenh. & Byng, comb. nov. Hakea linearifolia (Cav.) Christenh. & Byng, comb.
Basionym: Grevillea juniperina R.Br., Trans. Linn. Soc. nov. Basionym: Embothrium linearifolium Cav., Icon.
London, Bot. 10: 171. 1810. Descr. Pl. 4: 59. 1797.
Hakea kedumbensis (McGill.) Christenh. & Byng, Hakea linsmithii (McGill.) Christenh. & Byng, comb.
comb. & stat. nov. Basionym: Grevillea obtusiflora nov. Basionym: Grevillea linsmithii McGill., New
subsp. kedumbensis McGill., New Names Grevillea: Names Grevillea: 8. 1986.
11. 1986. Hakea lissopleura (McGill.) Christenh. & Byng, comb.
Hakea kenneallyi (McGill.) Christenh. & Byng, comb. nov. Basionym: Grevillea lissopleura McGill., New
nov. Basionym: Grevillea kenneallyi McGill., New Names Grevillea: 9. 1986.
Names Grevillea: 8. 1986. Hakea lobata (F.Muell.) Christenh. & Byng, comb. nov.
Hakea kennedyana (F.Muell.) Christenh. & Byng, comb. Basionym: Grevillea lobata F.Muell., Trans. Phil. Soc.
nov. Basionym: Grevillea kennedyana F.Muell., Victoria 1: 22. 1854.
Australasian Chemist and Druggist 2: 4. 1887. Hakea loboana (Domin) Christenh. & Byng, comb. nov.
Hakea kimberleyensis Christenh. & Byng, nom. nov. Basionym: Grevillea loboana Domin, Mem. Soc. Sci.
Basionym: Grevillea microcarpa Olde & Marriott, Bohème (Věstn. Král. České Společn. Nauk. Tř. Mat.-
Telopea 5: 415. 1993, not: Hakea microcarpa R.Br. Přír.) 2: 11. 1923. Syn.: Grevillea diversifolia subsp.
Etymology: This is a species native to the Kimberley subtersericata McGill., New Names Grevillea: 5.
Range. 1986.
Hakea kirkalocka (Olde & Marriott) Christenh. & Byng, Hakea longistyla (Hook.) Christenh. & Byng, comb.
comb. nov. Basionym: Grevillea kirkalocka Olde & nov. Basionym: Grevillea longistyla Hook., J. Exp. Int.
Marriott, Nuytsia 15: 93. 2002. Trop. Australia (Mitchell): 343. 1848.
Hakea lanata (Olde & Mariott) Christenh. & Byng, Hakea lucens (Makinson) Christenh. & Byng, comb. &
comb. & stat. nov. Basionym: Grevillea coccinea stat. nov. Basionym: Grevillea humilis subsp. lucens
subsp. lanata Olde & Marriott, Nuytsia 9: 277. 1993. Makinson, Fl. Australia 17A: 500. 2000.
Hakea laurifolia (Sieber ex Spreng.) Christenh. & Byng, Hakea lullfitzii (McGill.) Christenh. & Byng, comb. nov.
comb. nov. Basionym: Grevillea laurifolia Sieber ex Basionym: Grevillea lullfitzii McGill., New Names
Spreng., Cur. Post. Syst, Veg. 4: 46. 1827. Grevillea: 9. 1986.
Hakea lavandulacea (Schltdl.) Christenh. & Byng, Hakea maccutcheonii (Keighery & Cranfield)
comb. nov. Basionym: Grevillea lavandulacea Christenh. & Byng, comb. nov. Basionym: Grevillea
Schltdl., Linnaea 20: 586. 1847. (Figure 11B) maccutcheonii Keighery & Cranfield, Nuytsia 11: 33.
Hakea ledifolia Christenh. & Byng, nom. nov. Replaced 1996.
synonym: Grevillea costata C.Gardner ex A.S.George, Hakea macleayana (McGill.) Christenh. & Byng, comb.
Nuytsia 1: 370. 1974, not: Hakea costata Meisn. & stat. nov. Basionym: Grevillea barklyana subsp.
Etymology: This species has leaves that resemble macleayana McGill., New Names Grevillea: 2. 1986.
those of Rhododendron subsect. Ledum. Hakea macleayana (McGill.) Christenh. & Byng, comb.
Hakea leiophylla (F.Muell. ex Benth.) Christenh. & & stat. nov. Basionym: Grevillea barklyana subsp.
Byng, comb. nov. Basionym: Grevillea leiophylla macleayana McGill., New Names Grevillea: 2. 1986.
F.Muell. ex Benth., Fl. Austral. 5: 471. 1870. Hakea macrostylis (F.Muell.) Christenh. & Byng, comb.
Hakea leptobotrys (Meisn.) Christenh. & Byng, comb. nov. Basionym: Grevillea macrostylis F.Muell.,
nov. Basionym: Grevillea leptobotrys Meisn., Pl. Fragm. Phytogr. Austral. 1: 137. 1859.
Preiss. (Lehmann) 2: 256. 1848. Hakea magnifica (McGill.) Christenh. & Byng, comb. &
Hakea leptopoda (McGill.) Christenh. & Byng, comb. stat. nov. Basionym: Grevillea petrophiloides subsp.
nov. Basionym: Grevillea leptopoda McGill., New magnifica McGill., New Names Grevillea: 11. 1986.
Names Grevillea: 8. 1986. Hakea maherae (Makinson & M.D.Barrett) Christenh.
Hakea leucoclada (McGill.) Christenh. & Byng, comb. & Byng, comb. nov. Basionym: Grevillea maherae
nov. Basionym: Grevillea leucoclada McGill., New Makinson & M.D.Barrett, Fl. Australia 17A: 493.
Names Grevillea: 8. 1986. 2000.
Hakea leucopteris (Meisn.) Christenh. & Byng, comb. Hakea makinsonii (McGill.) Christenh. & Byng, comb.
nov. Basionym: Grevillea leucopteris Meisn., nov. Basionym: Grevillea makinsonii McGill., New
Hooker’s J. Bot. Kew Gard. Misc. 7: 76. 1855. Names Grevillea: 9. 1986.
Hakea levis (Olde & Marriott) Christenh. & Byng, comb. Hakea manglesii (Graham) Christenh. & Byng, comb.
nov. Basionym: Grevillea levis Olde & Marriott, nov. Basionym: Anadenia manglesii Graham,

85
Edinburgh New Phil. J. 27: 189. 1839. & Proc. Victorian Inst. Advancem. Sci. 1: 132. 1855.
Hakea manglesioides (Meisn.) Christenh. & Byng, Hakea mollis (Olde & Molyneux) Christenh. & Byng,
comb. nov. Basionym: Grevillea manglesioides comb. nov. Basionym: Grevillea mollis Olde &
Meisn., Pl. Preiss. (Lehmann) 1: 547. 1845. Molyneux, Telopea 5: 781. 1994.
Hakea mantalakifera Christenh. & M.W.Chase, nom. Hakea molyneuxii (McGill.) Christenh. & Byng, comb.
nov. Replaced synonym: Grevillea glauca Banks & nov. Basionym: Grevillea molyneuxii McGill., New
Sol. ex Knight, Cult. Prot.: 121. 1809, not: Hakea Names Grevillea: 10. 1986.
glauca Knight. Etymology: The common name is Hakea monslacana (Molyneux & Stajsic in Makinson)
bushman’s clothes peg, hence μανταλάκι meaning Christenh. & Byng, comb. nov. Basionym: Grevillea
clothes peg and φέρει, to bear. monslacana Molyneux & Stajsic in Makinson, Fl.
Hakea maritima (Makinson) Christenh. & Byng, comb. Australia 17A: 502. 2000.
& stat. nov. Basionym: Grevillea humilis subsp. Hakea montana (R.Br.) Christenh. & Byng, comb. nov.
maritima Makinson, Fl. Australia 17A: 500. 2000. Basionym: Grevillea montana R.Br., Trans. Linn. Soc.
Hakea marriottii (Olde) Christenh. & Byng, comb. London, Bot. 10: 172. 1810.
nov. Basionym: Grevillea marriottii Olde, Western Hakea monticola (Meisn.) Christenh. & Byng, comb.
Australian Naturalist Ed. 3, 18: 75. 1990. nov. Basionym: Grevillea monticola Meisn., Pl.
Hakea masonii (Olde & Marriott) Christenh. & Byng, Preiss. (Lehmann) 2: 259. 1848.
comb. nov. Basionym: Grevillea masonii Olde & Hakea montis-cole (R.V.Sm.) Christenh. & Byng, comb.
Marriott, Telopea 5: 720. 1994. nov. Basionym: Grevillea montis-cole R.V.Sm.,
Hakea maxwellii (McGill.) Christenh. & Byng, comb. Muelleria 5: 223. 1983.
nov. Basionym: Grevillea maxwellii McGill., New Hakea moroka (Molyneux & Stajsic) Christenh. & Byng,
Names Grevillea: 9. 1986. comb. & stat. nov. Basionym: Grevillea miqueliana
Hakea mcgillivrayi (I.M.Turner) Christenh. & Byng, subsp. moroka Molyneux & Stajsic in Makinson, Fl.
comb. nov. Basionym: Grevillea mcgillivrayi Australia 17A: 502. 200.
I.M.Turner, Ann. Bot. Fenn. 51: 307. 2014. Hakea mucronulata (R.Br.) Christenh. & Byng, comb.
Hakea meisneri (Montrouz.) Christenh. & Byng, comb. nov. Basionym: Grevillea mucronulata R.Br., Trans.
nov. Basionym: Grevillea meisneri Montrouz., Mém. Linn. Soc. London, Bot. 10: 173. 1810.
Acad. Roy. Sci., Lyon, n.s., 10: 248. 1860. Hakea muelleri (Benth.) Christenh. & Byng, comb. nov.
Hakea metamorpha (Makinson) Christenh. & Byng, Basionym: Grevillea muelleri Benth., Fl. Austral. 5:
comb. nov. Basionym: Grevillea metamorpha 479. 1870.
Makinson, Fl. Australia 17A: 506. 2000. Hakea murex (McGill.) Christenh. & Byng, comb. nov.
Hakea metaxa (Makinson) Christenh. & Byng, comb. & Basionym: Grevillea murex McGill., New Names
stat. nov. Basionym: Grevillea manglesioides subsp. Grevillea: 10. 1986.
metaxa Makinson, Fl. Australia 17A: 497. 2000. Hakea muricata (J.M.Black) Christenh. & Byng, comb.
Hakea micrantha (Meisn.) Christenh. & Byng, comb. nov. Basionym: Grevillea muricata J.M.Black, Trans.
nov. Basionym: Grevillea micrantha Meisn., Linnaea Roy. Soc. South Australia 63: 244. 1939.
26: 358. 1854. Hakea myosodes (McGill.) Christenh. & Byng, comb.
Hakea microstegia (Molyneux) Christenh. & Byng, nov. Basionym: Grevillea myosodes McGill., New
comb. nov. Basionym: Grevillea microstegia Names Grevillea: 10. 1986.
Molyneux, Muelleria 3: 141. 1975. Hakea nana (C.A.Gardner) Christenh. & Byng, comb.
Hakea microstyla (M.D.Barrett & Makinson) Christenh. nov. Basionym: Grevillea nana C.A.Gardner, J. Roy.
& Byng, comb. nov. Basionym: Grevillea microstyla Soc. Western Australia 27: 168. 1943.
M.D.Barrett & Makinson, Fl. Australia 17A: 495. Hakea nematophylla (F.Muell.) Christenh. & Byng,
2000. comb. nov. Basionym: Grevillea nematophylla
Hakea miniata (W.Fitzg.) Christenh. & Byng, comb. F.Muell., Fragm. Phytogr. Austral.1: 136. 1859.
nov. Basionym: Grevillea miniata W.Fitzg., Western Hakea neodissecta (I.M.Turner) Christenh. & Byng,
Mail: 21. 1906, and J. Proc. Roy. Soc. Western comb. nov. Basionym: Grevillea neodissecta
Australia 3: 131. 1918. I.M.Turner, Ann. Bot. Fenn. 51: 307. 2014. Replaced
Hakea minutiflora (McGill.) Christenh. & Byng, comb. synonym: Grevillea dissecta Olde & Marriott, Nuytsia
nov. Basionym: Grevillea minutiflora McGill., New 9: 282. 1993, nom. illeg.
Names Grevillea: 10. 1986. Hakea neorigida (I.M.Turner) Christenh. & Byng, comb.
Hakea miqueliana (F.Muell.) Christenh. & Byng, comb. nov. Basionym: Grevillea neorigida I.M.Turner,
nov. Basionym: Grevillea miqueliana F.Muell., Trans. Ann. Bot. Fenn. 51: 307. 2014. Replaced synonym:
Grevillea rigida Olde & Marriott, Grevillea Book 1:

186. 1995, nom. illeg. & Byng, comb. nov. Basionym: Grevillea oncogyne
Hakea newbeyi (McGill.) Christenh. & Byng, comb. Diels ex Diels & E.Pritz., Bot. Jahrb. Syst. 35: 149.
nov. Basionym: Grevillea newbeyi McGill., New 1904.
Names Grevillea: 10. 1986. Hakea ornithopoda (Meisn.) Christenh. & Byng, comb.
Hakea nivea (Olde & Marriott) Christenh. & Byng, nov. Basionym: Grevillea ornithopoda Meisn., Pl.
comb. nov. Basionym: Grevillea nivea Olde & Preiss. (Lehmann) 2: 256. 1848.
Marriott. Nuytsia 19: 237. 2009. Hakea oxyantha (Makinson) Christenh. & Byng, comb.
Hakea novae-austrocambriae Christenh. & M.W.Chase, nov. Basionym: Grevillea oxyantha Makinson.
nom. nov. Replaced synonym: Grevillea canescens Telopea 7: 130. 1997.
R.Br., Prodr. Fl. Nov. Holl., Suppl. 1: 18. 1830, not: Hakea pachylostyla (McGill.) Christenh. & Byng, comb.
Hakea canescens Link. Etymology: This species is & stat. nov. Basionym: Grevillea willisii subsp.
found in New South Wales, hence the name. pachylostyla McGill., New Names Grevillea: 16.
Hakea nudiflora (Meisn.) Christenh. & Byng, comb. 1986.
nov. Basionym: Grevillea nudiflora Meisn., Prodr. (A. Hakea pachypoda (O.Schwarz) Christenh. & Byng,
P. de Candolle) 14: 366. 1856. comb. nov. Basionym: Grevillea pachypoda
Hakea obliquistigma (C.A.Gardner) Christenh. & O.Schwarz, Repert. Spec. Nov. Regni Veg. 24: 83.
Byng, comb. nov. Basionym: Grevillea obliquistigma 1927. Replaced synonym: Grevillea mimosoides
C.A.Gardner, J. Roy. Soc. Western Australia 27: 169. R.Br., Trans. Linn. Soc. London, Bot. 10: 177. 1810,
1943. not: Hakea mimosoides A.Cunn. ex Meisn.
Hakea obtecta (Molyneux) Christenh. & Byng, comb. Hakea pagna (Cranfield) Christenh. & Byng, comb. &
nov. Basionym: Grevillea obtecta Molyneux, stat. nov. Basionym: Grevillea bipinnatifida R.Br.
Muelleria 6: 147. 1985. subsp. pagna Cranfield, Nuytsia 15: 190. 2004.
Hakea obtusiflora (R.Br.) Christenh. & Byng, comb. Hakea pallarupensis Christenh. & M.W.Chase, nom.
nov. Basionym: Grevillea obtusiflora R.Br., Prodr. Fl. nov. Replaced synonym: Grevillea prostrata
Nov. Holl., Suppl. 1: 19. 1830. C.AGardner & A.S.George, J. Roy. Soc. Western
Australia 46: 129, not: Hakea prostrata R.Br.
Hakea obtusifolia (Meisn.) Christenh. & Byng, comb.
Etymology: This species is named for the Pallarup
nov. Basionym: Grevillea obtusifolia Meisn., Prodr.
Nature Reserve in Western Australia, where it was
(A. P. de Candolle) 14: 356. 1856.
first discovered.
Hakea occidentalis (R.Br.) Christenh. & Byng, comb.
Hakea paludosa (Makinson & Albr.) Christenh. & Byng,
nov. Basionym: Grevillea occidentalis R.Br., Trans.
comb. & stat. nov. Basionym: Grevillea acanthifolia
Linn. Soc. London, Bot. 10: 173. 1810.
subsp. paludosa Makinson & Albr., Muelleria 7: 89.
Hakea oldei (McGill.) Christenh. & Byng, comb. nov. 1989.
Basionym: Grevillea oldei McGill., New Names
Hakea paniculata (Meisn.) Christenh. & Byng, comb.
Grevillea: 11. 1986.
nov. Basionym: Grevillea paniculata Meisn., Pl.
Hakea olearioides Christenh. & Byng, comb. nov. Preiss. (Lehmann) 1: 550. 1845.
Replaced synonym: Grevillea erinacea Meisn.,
Hakea papillosa (McGill.) Christenh. & Byng, comb. &
Hooker’s J. Bot. Kew Garden Misc. 7: 74. 1855, not:
stat. nov. Basionym: Grevillea manglesioides subsp.
Hakea erinacea Meisn. Etymology: This species
papillosa McGill., New Names Grevillea: 9. 1986.
resembles a member of the Asteraceae genus
Olearia, hence the name. Hakea papuana (Diels) Christenh. & Byng, comb. nov.
Basionym: Grevillea papuana Diels, Bot. Jahrb. Syst.
Hakea oleoides (Sieber ex Schult. & Schult.f.)
54: 205. 1916.
Christenh. & Byng, comb. nov. Basionym: Grevillea
oleoides Sieber ex Schult. & Schult.f., Mant. Ed. 16, Hakea paradoxa (F.Muell.) Christenh. & Byng, comb.
3: 277. 1827. nov. Basionym: Grevillea paradoxa F.Muell., Fragm.
Phytogr. Austral. 6: 246. 1868
Hakea oligantha (F.Muell.) Christenh. & Byng, comb.
nov. Basionym: Grevillea oligantha F.Muell., Fragm. Hakea parallela (Knight) Christenh. & Byng, comb.
Phytogr. Austral. 6: 206. 1868. nov. Basionym: Grevillea parallela Knight, Cult.
Prot.: 121. 1809.
Hakea oligomera (McGill.) Christenh. & Byng, comb. &
stat. nov. Basionym: Grevillea petrophiloides subsp. Hakea parallelinervis (Carrick) Christenh. & Byng,
oligomera McGill., New Names Grevillea: 12. 1986. comb. nov. Basionym: Grevillea parallelinervis
Carrick, Contrib. Herb. Austral. 15: 1. 1976.
Hakea olivacea (A.S.George) Christenh. & Byng, comb.
nov. Basionym: Grevillea olivacea A.S.George, Hakea parviflora (R.Br.) Christenh. & Byng, comb. nov.
Nuytsia 1: 373. 1974. Basionym: Grevillea parviflora R.Br., Trans. Linn. Soc.
Hakea oncogyne (Diels ex Diels & E.Pritz.) Christenh.

87

Hakea parvula (Molyneux & Stajsic) Christenh. & Byng, Hakea pluricaulis (McGill.) Christenh. & Byng, comb.
comb. nov. Basionym: Grevillea parvula Molyneux & & stat. nov. Basionym: Grevillea goodii subsp.
Stajsic, Fl. Australia 17A: 502. 2000. pluricaulis McGill., New Names Grevillea: 6. 1986.
Hakea pateana Christenh. & M.W.Chase, nom. nov. Hakea plurijuga (F.Muell.) Christenh. & Byng, comb.
Replaced synonym: Grevillea intricata Meisn., nov. Basionym: Grevillea plurijuga F.Muell., Fragm.
Hooker’s J. Bot. Kew Garden Misc. 7: 74. 1855, not: Phytogr. Austral. 4: 84. 1864. (Figure 11C)
Hakea intricata R.Br. Etymology: This species is Hakea polyacida (McGill.) Christenh. & Byng, comb.
named in honour of John S. Pate, who contributed nov. Basionym: Grevillea polyacida McGill., New
extensively to the ecology and taxonomy of Western Names Grevillea: 12. 1986.
Australian plants. Hakea polybotrya (Meisn.) Christenh. & Byng, comb.
Hakea patentiloba (F.Muell.) Christenh. & Byng, comb. nov. Basionym: Grevillea polybotrya Meisn. Prodr.
nov. Basionym: Grevillea patentiloba F.Muell., [A. P. de Candolle] 14: 386. 1856.
Fragm. Phytogr. Austral. 1: 137. 1859. Hakea polybractea (H.B.Will.) Christenh. & Byng,
Hakea patulifolia (Gand.) Christenh. & Byng, comb. comb. nov. Basionym: Grevillea polybractea
nov. Basionym: Grevillea patulifolia Gand., Bull. Soc. H.B.Will., Victorian Naturalist 44: 139. 1927.
Bot. France 66: 231. 1919. Hakea polychroma (Molyneux & Stajsic) Christenh.
Hakea pauciflora (R.Br.) Christenh. & Byng, comb. nov. & Byng, comb. & stat. nov. Basionym: Grevillea
Basionym: Grevillea pauciflora R.Br., Trans. Linn. Soc. brevifolia subsp. polychroma Molyneux & Stajsic in
London, Bot. 10: 171. 1810. Makinson, Fl. Australia 17A: 502. 2000.
Hakea phanerophlebia (Diels) Christenh. & Byng, Hakea pompona Christenh. & M.W.Chase, nom. nov.
comb. nov. Basionym: Grevillea phanerophlebia Replaced synonym: Grevillea incrassata Diels &
Diels & E.Pritz., Bot. Jahrb. Syst. 35: 157. 1904. E.Pritz., Bot. Jahrb. Syst. 35: 156. 1904, not: Hakea
Hakea phillipsiana (McGill.) Christenh. & Byng, comb. incrassata R.Br. Etymology: The inflorescence
nov. Basionym: Grevillea phillipsiana McGill., New resembles a pompon, hence the name.
Names Grevillea: 12. 1986. Hakea prasina (McGill.) Christenh. & Byng, comb. nov.
Hakea phylicoides (R.Br.) Christenh. & Byng, comb. Basionym: Grevillea prasina McGill., New Names
nov. Basionym: Grevillea phylicoides R.Br., Trans. Grevillea: 12. 1986.
Linn. Soc. London, Bot. 10: 174. 1810. Hakea prominens (Olde & Marriott) Christenh. & Byng,
Hakea pilosa (A.S.George) Christenh. & Byng, comb. comb. nov. Basionym: Grevillea prominens Olde &
nov. Basionym: Grevillea pilosa A.S.George, Western Marriott, Nuytsia 9: 262. 1993.
Australian Naturalist 10: 32. 1966. Hakea psilantha (McGill.) Christenh. & Byng, comb.
Hakea pimeleoides (W.Fitzg.) Christenh. & Byng, comb. nov. Basionym: Grevillea psilantha McGill., New
nov. Basionym: Grevillea pimeleoides W.Fitzg., J. Names Grevillea: 12. 1986.
Proc. Mueller Bot. Soc. Western Australia 1: 37. Hakea psilophylla (McGill.) Christenh. & Byng, comb.
1902. & stat. nov. Basionym: Grevillea pauciflora subsp.
Hakea pinaster (Meisn.) Christenh. & Byng, comb. nov. psilophylla McGill., New Names Grevillea: 11. 1986.
Basionym: Grevillea pinaster Meisn., Hooker’s J. Bot. Hakea pteridifolia (Knight) Christenh. & Byng, comb.
Kew Gard. Misc. 7: 76. 1855. nov. Basionym: Grevillea pteridifolia Knight, Cult.
Hakea pinnatifida (F.M.Bailey) Christenh. & Byng, Prot.: 121. 1809.
comb. nov. Basionym: Kermadecia pinnatifida Hakea pterosperma (F.Muell.) Christenh. & Byng,
F.M.Bailey, Cat. Woods Queensl.: 69. 1886. Syn.: comb. nov. Basionym: Grevillea pterosperma
Grevillea baileyana McGill., New Names Grevillea: 2. F.Muell., Trans. Phil. Soc. Victoria 1: 22. 1854.
1986, not: Grevillea pinnatifida Jacques. Hakea pulchella (R.Br.) Christenh. & Byng, comb. nov.
Hakea pinnatisecta (F.Muell.) Christenh. & Byng, Basionym: Anadenia pulchella R.Br., Trans. Linn. Soc.
comb. & stat. nov. Basionym: Grevillea commutata London 10: 167. 1810.
var. pinnatisecta F.Muell., Fragm. Phytogr. Austral. 6: Hakea punctata (Olde & Marriott) Christenh. & Byng,
208. 1868. comb. nov. Basionym: Grevillea punctata Olde &
Hakea pityophylla (F.Muell.) Christenh. & Byng, comb. Marriott, Grevillea Book 1: 182. 1994.
nov. Basionym: Grevillea pityophylla F.Muell., Hakea pungens (R.Br.) Christenh. & Byng, comb. nov.
Fragm. Phytogr. Austral. 6: 208. 1868. Basionym: Grevillea pungens R.Br., Trans. Linn. Soc.
Hakea platypoda (F.Muell.) Christenh. & Byng, comb. London, Bot. 10: 175. 1810.
nov. Basionym: Grevillea platypoda F.Muell., Fragm. Hakea pyramidalis (A.Cunn. ex R.Br.) Christenh. &
Byng, comb. nov. Basionym: Grevillea pyramidalis

A.Cunn. ex R.Br., Prodr. Fl. Nov. Holl., Suppl. 1: 25. nov. Basionym: Grevillea ripicola A.S.George,
1830. Nuytsia 1: 373. 1974.
Hakea pythara (Olde & Marriott) Christenh. & Byng, Hakea rivularis (McGill.) Christenh. & Byng, comb.
comb. nov. Basionym: Grevillea pythara Olde & nov. Basionym: Grevillea rivularis McGill., Telopea
Marriott, Nuytsia 9: 293. 1993. 1: 23. 1975.
Hakea quadricauda (Olde & Marriott) Christenh. & Hakea robusta (A.Cunn. ex R.Br.) Christenh. &
Byng, comb. nov. Basionym: Grevillea quadricauda M.W.Chase, comb. nov. Basionym: Grevillea robusta
Olde & Marriott, Telopea 5: 722. 1994. A.Cunn. ex R.Br., Prodr. Fl. Nov. Holl., Suppl. 1: 24.
Hakea quercifolia (R.Br.) Christenh. & Byng, comb. 1830. (Figure 11D)
nov. Basionym: Grevillea quercifolia R.Br., Prodr. Fl. Hakea rogersoniana (C.A.Gardner) Christenh. &
Nov. Holl., Suppl. 1: 23. 1830. Byng, comb. nov. Basionym: Grevillea rogersoniana
Hakea quinquenervis (J.M.Black) Christenh. & Byng, C.A.Gardner, J. Roy. Soc. Western Australia 47: 56.
comb. nov. Basionym: Grevillea quinquenervis 1964.
J.M.Black, Trans. Proc. Report Roy. Soc. South Hakea rosea Christenh. & M.W.Chase, nom. nov.
Australia 33: 225. 1909. Replaced synonym: Grevillea petrophiloides Meisn.,
Hakea ramosissima (Meisn.) Christenh. & Byng, comb. Pl. Preiss. (Lehmann) 2: 257. 1848, not: Hakea
nov. Basionym: Grevillea ramosissima Meisn., Prodr. petrophiloides Sweet. Etymology: This species,
[A. P. de Candolle] 14: 388. 1856. called ‘pink pokers’, has pink flowers, hence the
name. (Figure 11E)
Hakea rara (Olde & Marriott) Christenh. & Byng, comb.
nov. Basionym: Grevillea rara Olde & Marriott, Hakea rosieri (McGill.) Christenh. & Byng, comb. nov.
Nuytsia 9: 244. 1993. Basionym: Grevillea rosieri McGill., New Names
Hakea raybrownii (Olde & Marriott) Christenh. &
Byng, comb. nov. Basionym: Grevillea raybrownii Hakea rosmarinifolia (A.Cunn.) Christenh. & Byng,
Olde & Marriott. Telopea 5: 774. 1994. comb. nov. Basionym: Grevillea rosmarinifolia
A.Cunn., Geogr. Mem. New South Wales (Field): 328.
Hakea recedens (Olde & Marriott) Christenh. & Byng,
1825.
comb. & stat. nov. Basionym: Grevillea haplantha
subsp. recedens Olde & Marriott, Nuytsia 9: 290. Hakea roycei (McGill.) Christenh. & Byng, comb. nov.
1993. Basionym: Grevillea roycei McGill., New Names
Hakea refracta (R.Br.) Christenh. & Byng, comb. nov.
Basionym: Grevillea refracta R.Br., Trans. Linn. Soc. Hakea rubicunda (S.Moore) Christenh. & Byng, comb.
London, Bot. 10: 176. 1810. nov. Basionym: Grevillea rubicunda S.Moore, J. Linn.
Soc., Bot. 45: 210. 1920.
Hakea remota (Olde & Marriott) Christenh. & Byng,
comb. & stat. nov. Basionym: Grevillea magnifica Hakea rubiginosa (Brongn. & Gris) Christenh. & Byng,
subsp. remota Olde & Marriott, Grevillea Book 1: comb. nov. Basionym: Grevillea rubiginosa Brongn.
179. 1994. & Gris, Bull. Soc. Bot. France 12: 40. 1865.
Hakea renwickiana (F.Muell.) Christenh. & Byng, comb. Hakea rubra Christenh. & M.W.Chase, nom. nov.
nov. Basionym: Grevillea renwickiana F.Muell.. Proc. Replaced synonym: Grevillea sulcata C.AGardner ex
Linn. Soc. New South Wales, Ser. 2, 1: 1105. 1887. Olde & Marriott, Grevillea Book 1: 183. 1994, not:
Hakea sulcata R.Br. Etymology: This is one of the
Hakea repens (F.Muell. ex Meisn.) Christenh. & Byng,
species with red flowers, hence the name.
comb. nov. Basionym: Grevillea repens F.Muell. ex
Meisn., Linnaea 26: 355. 1854. Hakea rudis (Meisn.) Christenh. & Byng, comb. nov.
Basionym: Grevillea rudis Meisn., Hooker’s J. Bot.
Hakea reptans (Makinson) Christenh. & Byng, comb.
Kew Gard. Misc. 7: 73. 1855.
nov. Basionym: Grevillea reptans Makinson, Fl.
Australia 17A: 500. 2000. Hakea rufa (Warb.) Christenh. & Byng, comb. nov.
Basionym: Finschia rufa Warb., Bot. Jahrb. Syst.
Hakea rhizomatosa (Olde & Marriott) Christenh. &
13: 297. 1891. Synonym: Finschia carrii (Sleumer)
Byng, comb. nov. Basionym: Grevillea rhizomatosa
C.T.White
Olde & Marriott, Telopea 5: 724. 1994.
Hakea saccata (Benth.) Christenh. & Byng, comb. nov.
Hakea rhyolitica (Makinson) Christenh. & Byng, comb.
Basionym: Grevillea saccata Benth., Fl. Austral. 5:
nov. Basionym: Grevillea rhyolitica Makinson,
450. 1870. (Figure 11F)
Telopea 7: 134. 1997.
Hakea sarissa (S.Moore) Christenh. & Byng, comb.
Hakea riparia (R.Br.) Christenh. & Byng, comb. nov.
nov. Basionym: Grevillea sarissa S.Moore, J. Linn.
Basionym: Grevillea riparia R.Br., Trans. Linn. Soc.
Soc., Bot. 34: 222. 1899.
London, Bot. 10: 170. 1810.
Hakea saxatilis (McGill.) Christenh. & Byng, comb.
Hakea ripicola (A.S.George) Christenh. & Byng, comb.

89
& stat. nov. Basionym: Grevillea pauciflora subsp. comb. nov. Basionym: Grevillea shuttleworthiana
saxatilis McGill., New Names Grevillea : 1. 1986. Meisn., Pl. Preiss. (Lehmann) 2: 258. 1848.
Hakea saxicola (S.J.Dillon) Christenh. & Byng, comb. Hakea singuliflora (F.Muell.) Christenh. & Byng, comb.
nov. Basionym: Grevillea saxicola S.J.Dillon, Nuytsia nov. Basionym: Grevillea singuliflora F.Muell.,
24: 104. 2014. Fragm. Phytogr. Austral. 6: 92. 1867.
Hakea scabra (Meisn.) Christenh. & Byng, comb. Hakea sparsiflora (F.Muell.) Christenh. & Byng, comb.
nov. Basionym: Grevillea scabra Meisn., Pl. Preiss. nov. Basionym: Grevillea sparsiflora F.Muell., Fragm.
(Lehmann) 1: 541. 1845, in adnot. Phytogr. Austral. 6: 206. 1868.
Hakea scabrida (C.A.Gardner) Christenh. & Byng, comb. Hakea speciosa (Knight) Christenh. & Byng, comb. nov.
nov. Basionym: Grevillea scabrida C.A.Gardner, J. Basionym: Lysanthe speciosa Knight, Cult. Prot. 118.
Roy. Soc. Western Australia 22: 120. 1936. 1809.
Hakea scapigera (A.S.George) Christenh. & Byng, Hakea sphacelata (R.Br.) Christenh. & Byng, comb.
comb. nov. Basionym: Grevillea scapigera nov. Basionym: Grevillea sphacelata R.Br., Trans.
A.S.George, Nuytsia 1: 374. 1974. Linn. Soc. London, Bot. 10: 174. 1810.
Hakea scortechinii (F.Muell. ex Scort.) Christenh. Hakea spinigera Christenh. & Byng, nom. nov.
& Byng, comb. & stat. nov. Basionym: Grevillea Replaced synonym: Grevillea halmaturina Tate
ilicifolia var. scortechinii F.Muell. ex Scort., Proc. Linn. subsp. laevis Makinson, Fl. Australia 17A: 501.
Soc. New South Wales 8: 174. 1883. 2000, not: Hakea levis (Olde & Marriott) Christenh.
Hakea scottensis (Keighery) Christenh. & Byng, nom. & Byng. Etymology: This species has spiny leaves,
nov. Replaced synonym: Grevillea brachystylis subsp. hence its name.
australis Keighery, Nuytsia 7: 128. 1990, not: Hakea Hakea spinosa (McGill.) Christenh. & Byng, comb. nov.
australis (R.Br.) Christenh. & Byng. Etymology: The Basionym: Grevillea spinosa McGill., New Names
type was collected in Scott National Park. Grevillea: 14. 1986.
Hakea secunda (McGill.) Christenh. & Byng, comb. Hakea spinosissima (McGill.) Christenh. & Byng, comb.
nov. Basionym: Grevillea secunda McGill., New nov. Basionym: Grevillea spinosissima McGill., New
Names Grevillea: 14. 1986. Names Grevillea: 14. 1986.
Hakea seminuda Christenh. & Byng, nom. nov. Hakea squiresiae (Olde & Marriott) Christenh. & Byng,
Replaced synonym: Grevillea amplexans subsp. comb. nov. Basionym: Grevillea squiresiae Olde &
semivestita Makinson, Fl. Australia 17A: 505. 2000, Marriott, Nuytsia 15: 96. 2002, as ‘squiresae’.
not Hakea semivestita (Makinson) Christenh. & Hakea stajsiciana Christenh. & M.W.Chase, nom.
Byng. Etymology: The epithet means half naked, nov. Replaced synonym: Grevillea curviloba McGill.
which is effectively the same as semivestita, half subsp. incurva Olde & Marriott, Nuytsia 9: 243.
clothed. 1993, not: Hakea incurva (Diels) Christenh. & Byng.
Hakea semivestita (Makinson) Christenh. & Byng, Etymology: This species is named for Val Stajsic, for
comb. & stat. nov. Basionym: Grevillea rhyolitica his contributions to the knowledge of Australian
subsp. semivestita Makinson, Telopea 7: 137. 1997. Proteaceae.
Hakea ×semperflorens (F.E.Briggs ex Mulligan) Hakea steiglitziana (N.A.Wakef.) Christenh. & Byng,
Christenh. & Byng, comb. nov. Basionym: Grevillea comb. nov. Basionym: Grevillea steiglitziana
×semperflorens F.E.Briggs ex Mulligan, J. Roy. Hort. N.A.Wakef., Victorian Naturalist 73: 74. 1956.
Soc., London 62: 126. 1937 (= H. thelemanniana × Hakea stenobotrya (F.Muell.) Christenh. & Byng, comb.
H. juniperina). nov. Basionym: Grevillea stenobotrya F.Muell.,
Hakea senegalioides Christenh. & Byng, nom. nov. Fragm. Phytogr. Austral. 9: 3. 1875.
Replaced synonym: Grevillea acacioides C.Gardner Hakea stenogyne (Benth.) Christenh. & Byng, comb. &
ex McGill., New Names Grevillea: 1. 1986, not: stat. nov. Basionym: Grevillea vestita var. stenogyne
Hakea acacioides Domin. Etymology: Named for the Benth., Fl. Austral. 5: 488. 1870.
Fabaceae genus Senegalia, a segregate of Acacia. Hakea stenomera (F.Muell.) Christenh. & Byng, comb.
Hakea sessilis (C.T.White & W.D.Francis) Christenh. nov. Basionym: Grevillea stenomera F.Muell., Fragm.
& Byng, comb. nov. Basionym: Grevillea sessilis Phytogr. Austral. 4: 85. 1864.
C.T.White & W.D.Francis, Proc. Roy. Soc. Queensland Hakea stenostachya (C.A.Gardner) Christenh. &
37: 165. 1926. Byng, comb. nov. Basionym: Grevillea stenostachya
Hakea shiressii (Blakely) Christenh. & Byng, comb. nov. C.A.Gardner, J. Roy. Soc. Western Australia 22: 122.
Basionym: Grevillea shiressii Blakely, Proc. Linn. Soc. 1936.
New South Wales 50: 383. 1925. Hakea striata (R.Br.) Christenh. & Byng, comb. nov.
Hakea shuttleworthiana (Meisn.) Christenh. & Byng, Basionym: Grevillea striata R.Br., Trans. Linn. Soc.


Hakea subterlineata (Makinson) Christenh. & Byng, Hakea trifida (R.Br.) Christenh. & Byng, comb. nov.
comb. nov. Basionym: Grevillea subterlineata Basionym: Anadenia trifida R.Br. Trans. Linn. Soc.
Makinson in McGill., Grevillea, Proteac.: taxon. rev.: London, Bot. 10: 167. 1810.
443. 1993. Hakea triloba (Meisn.) Christenh. & Byng, comb. nov.
Hakea subtiliflora (McGill.) Christenh. & Byng, comb. Basionym: Grevillea triloba Meisn., Hooker’s J. Bot.
nov. Basionym: Grevillea subtiliflora McGill., New Kew Gard. Misc. 7: 74. 1855.
Names Grevillea: 14. 1986. Hakea tripartita (Meisn.) Christenh. & Byng, comb.
Hakea superba (Olde & Marriott) Christenh. & Byng, nov. Basionym: Grevillea tripartita Meisn., Prodr. [A.
comb. nov. Basionym: Grevillea superba Olde & P. de Candolle] 14: 373. 1856.
Marriott, Nuytsia 9: 298. 1993. Hakea triternata (R.Br.) Christenh. & Byng, comb. nov.
Hakea supplicans (Makinson) Christenh. & Byng, Basionym: Grevillea triternata R.Br., Prodr. Fl. Nov.
comb. & stat. nov. Basionym: Grevillea parviflora Holl., Suppl. 1: 21. 1830.
subsp. supplicans Makinson, Fl. Australia 17A: 501. Hakea umbellulata (J.M.Black) Christenh. & Byng,
200. comb. nov. Basionym: Grevillea umbellulata
Hakea synapheae (R.Br.) Christenh. & Byng, comb. J.M.Black, Trans. Roy. Soc. South Australia 71: 21.
nov. Basionym: Grevillea synapheae R.Br., Prodr. Fl. 1947.
Nov. Holl., Suppl. 1: 23. 1830. Hakea uncinulata (Diels & E.Pritz.) Christenh. & Byng,
Hakea telfordii (Makinson) Christenh. & Byng, comb. & comb. nov. Basionym: Grevillea uncinulata Diels &
stat. nov. Basionym: Grevillea decora subsp. telfordii E.Pritz., Bot. Jahrb. Syst. 35: 152. 1904.
Makinson, Fl. Australia 17A: 494. 2000. Hakea uniformis (McGill.) Christenh. & Byng, comb.
Hakea temulenta (Olde & Marriott) Christenh. & Byng, & stat. nov. Basionym: Grevillea acrobotrya subsp.
comb. & stat. nov. Basionym: Grevillea juncifolia uniformis McGill., New Names Grevillea: 1. 1986.
subsp. temulenta Olde & Marriott, Grevillea Book 1: Hakea variifolia (C.A.Gardner & A.S.George) Christenh.
185. 1994. & Byng, comb. nov. Basionym: Grevillea variifolia
Hakea tenuiloba (C.A.Gardner) Christenh. & C.A.Gardner & A.S.George, A.S., J. Roy. Soc. Western
Byng, comb. nov. Basionym: Grevillea tenuiloba Australia 46: 129. 1963.
C.A.Gardner, J. Roy. Soc. Western Australia 19: 81. Hakea velutinella (McGill.) Christenh. & Byng, comb.
1934. nov. Basionym: Grevillea velutinella McGill., New
Hakea tetragonoloba (Meisn.) Christenh. & Byng, Names Grevillea: 16. 1986.
comb. nov. Basionym: Grevillea tetragonoloba Hakea venusta (R.Br.) Christenh. & Byng, comb. nov.
Meisn., Prodr. [A. P. de Candolle] 14: 374. 1856. Basionym: Grevillea venusta R.Br., Trans. Linn. Soc.
Hakea tetrapleura (McGill.) Christenh. & Byng, comb. London 10: 175. 1811.
nov. Basionym: Grevillea tetrapleura McGill., New Hakea versicolor (McGill.) Christenh. & Byng, comb.
Names Grevillea: 15. 1986. nov. Basionym: Grevillea versicolor McGill., New
Hakea thelemanniana (Hügel ex Endl.) Christenh. Names Grevillea: 16. 1986.
& Byng, comb. nov. Basionym: Grevillea Hakea vestita (Endl.) Christenh. & Byng, comb. nov.
thelemanniana Hügel ex Endl., Nov. Stirp. Dec. 1: 6. Basionym: Manglesia vestita Endl., Nov. Stirp. Dec.
1839. 4: 26. 1839.
Hakea thyrsoides (Meisn.) Christenh. & Byng, comb. Hakea victoriae (F.Muell.) Christenh. & Byng, comb.
nov. Basionym: Grevillea thyrsoides Meisn., Hooker’s nov. Basionym: Grevillea victoriae F.Muell., Trans.
J. Bot. Kew Gard. Misc. 7: 77. 1855. Proc. Phil. Soc. Victoria 1: 107. 1855, not: Hakea
Hakea toodyay Christenh. & M.W.Chase, nom. nov. victoria J.Drumm.
Replaced synonym: Grevillea candolleana Meisn, Hakea viridiflava (Makinson) Christenh. & Byng, comb.
Pl. Preiss. (Lehmann) 1: 541. 1845, not: Hakea nov. Basionym: Grevillea viridiflava Makinson, Fl.
candolleana Meisn. Etymology: This species is Australia 17A: 500. 2000.
known as ‘toodyay grevillea’, hence the choice of Hakea westoniana Christenh. & M.W.Chase, nom. nov.
epithet. Replaced synonym: Grevillea virgata Makinson, Fl.
Hakea trachytheca (F.Muell.) Christenh. & Byng, comb. Australia 17A: 500. 2000, not: Hakea virgata R.Br.
nov. Basionym: Grevillea trachytheca F.Muell., Etymology: This species is named in honour of
Fragm. Phytogr. Austral. 6: 207. 1868. Peter Weston, for his contribution to Proteaceae
Hakea treueriana (F.Muell.) Christenh. & Byng, comb. systematics and taxonomy.
nov. Basionym: Grevillea treueriana F.Muell., Fragm. Hakea whiteana (McGill.) Christenh. & Byng, comb.
nov. Basionym: Grevillea whiteana McGill., New

91
Names Grevillea: 16. 1986. & Byng, comb. nov. Basionym: Haloragodendron
Hakea wickhamii (Meisn.) Christenh. & Byng, comb. glandulosum Orchard, Bull. Auckland Inst. Mus. 10:
nov. Basionym: Grevillea wickhamii Meisn., Prodr. 148. 1975.
[A. P. de Candolle] 14: 380. 1856. Glischrocaryon lucasii (Maid. & Betche) Christenh. &
Hakea wilkinsonii (Makinson) Christenh. & Byng, Byng, comb. nov. Basionym: Haloragis lucasii Maid.
comb. nov. Basionym: Grevillea wilkinsonii & Betche, Proc. Linnean Soc. New South Wales 34:
Makinson, Telopea 5: 351. 1993. 358. 1909.
Hakea willisii (R.V.Sm. & McGill.) Christenh. & Byng, Glischrocaryon monospermum (F.Muell.) Christenh. &
comb. nov. Basionym: Grevillea willisii R.V.Sm. & Byng, comb. nov. Basionym: Haloragis monosperma
McGill., Muelleria 3: 102. 1975. F.Muell., Proc. Linnean Soc. New South Wales 10:
197. 1885.
Hakea wilsonii (A.Cunn.) Christenh. & Byng, comb.
nov. Basionym: Grevillea wilsonii A.Cunn. in Wilson, Glischrocaryon racemosum (Labill.) Christenh. & Byng,
Narr. Voy. Round World: 273. 1835. comb. nov. Basionym: Haloragis racemosa Labill.,
Nov. Holl. Pl. 1: 100. 1805.
Hakea wiradjuri (Makinson) Christenh. & Byng, comb.
nov. Basionym: Grevillea wiradjuri Makinson, Fl.
Australia 17A: 501. 2000. 138. Zygophyllaceae. References: Sheahan & Chase
Hakea wirregaensis (Downing) Christenh. & Byng, (2000), Beier et al. (2003).
comb. & stat. nov. Basionym: Grevillea angustiloba Note: Augea Thunb., Fagonia L. and Tetraena Maxim. are
(F.Muell.) Downing subsp. wirregaensis Downing, embedded in Zygophyllum L.. We prefer to include all
Austral. Syst. Bot. 17: 340. 2004. in a single genus, because morphologically it would
Hakea wittweri (McGill.) Christenh. & Byng, comb. be difficult to find characters for the new genera that
nov. Basionym: Grevillea wittweri McGill., New would otherwise be required. Additionally, as more
Names Grevillea: 16. 1986. species are sequenced it is likely that these new
genera will be found to be polyphyletic and will need
Hakea xerophylla Christenh. & M.W.Chase, nom. nov.
to be split again without morphological support. Both
Basionym: Grevillea teretifolia Meisn., Pl. Preiss.
Zygophyllum and Fagonia are both Linnean names
(Lehmann) 2: 255 1848, not: Hakea teretifolia
but we chose Zygophyllum over Fagonia, because
(Salisb.) Britten. Etymology: Referring to the xeric
the first is a much larger genus and would require
leaves of this species.
a greater degree of nomenclatural change. Izozogia
Hakea xiphoidea (Olde & Marriott) Christenh. & Byng, G.Navarro shares all characters with Guaiacum L.
comb. nov. Basionym: Grevillea xiphoidea Olde & and is not separate from that genus. Some species of
Marriott, Grevillea Book 1: 175. 1994. Bulnesia Gay are better placed in Plectrocarpa Gillies
Hakea yorkrakinensis (C.A.Gardner) Christenh. & ex Hook. & Arn., because Bulnesia would otherwise
Byng, comb. nov. Basionym: Grevillea yorkrakinensis be biphyletic.
C.A.Gardner, J. Proc. Roy. Soc. Western Australia 9: Guaiacum nellii (Navarro) Christenh. & Byng, comb.
34. 1923. nov. Basionym: Izozogia nellii G.Navarro, Novon 7:
Hakea zygoloba (Olde & Marriott) Christenh. & Byng, 1. 1997.
comb. nov. Basionym: Grevillea zygoloba Olde & Plectrocarpa arborea (Jacq.) Christenh. & Byng, comb.
Marriott, Grevillea Book 1: 180. 1994. nov. Basionym: Zygophyllum arboreum Jacq., Enum.
Syst. Pl.: 20. 1760.
134. Haloragaceae. Reference: Chen et al. (2014). Plectrocarpa bonariensis (Groseb.) Christenh. & Byng,
Note: Glischrocaryon Endl. is embedded in comb. nov. Basionym: Bulnesia bonariensis Griseb.,
Haloragodendron Orchard, together forming a clade Abh. Königl. Gesell. Wiss. Göttingen 19: 105. 1874.
sister to the rest of Haloragaceae. As Glischrocaryon Plectrocarpa carrapo (Killip & Dugand) Christenh. &
has nomenclatural priority, Haloragodendron is Byng, comb. nov. Basionym: Bulnesia carrapo Killip
hereby transferred into that genus. & Dugand, Caldasia 3: 35. 1944.
Glischrocaryon baeuerlenii (F.Muell) Christenh. & Plectrocarpa sarmientoi (Lorentz ex Griseb.)
Byng, comb. nov. Basionym: Haloragis baeuerlenii Christenh. & Byng, comb. nov. Basionym: Bulnesia
F.Muell., Bot. Centralbl. 8: 147. 1887. sarmientoi Lorentz ex Griseb., Abh. Königl. Gesell.
Glischrocaryon gibsonii (Peter G. Wilson & Wiss. Göttingen 24: 75. 1879, as ‘Sarmienti’.
M.L.Moody) Christenh. & Byng, comb. nov. Zygophyllum acerosum (Boiss.) Christenh. & Byng,
Basionym: Haloragodendron gibsonii Peter G.Wilson comb. nov. Basionym: Fagonia acerosa Boiss.,
& M.L.Moody, Telopea 11: 142. 2006. Diagn. Pl. Orient. 8: 124. 1849.
Glischrocaryon glandulosum (Orchard) Christenh. Zygophyllum arabicum (L.) Christenh. & Byng, comb.

nov. Basionym: Fagonia arabica L., Sp. Pl.: 386. 1753. Beier & Thulin, Syst. Biodivers. 3: 243. 2005.
Zygophyllum augea Christenh. & Byng, nom. nov. Zygophyllum luntii (Baker) Christenh. & Byng, comb.
Replaced synonym: Augea capensis Thunb., Prodr. nov. Basionym: Fagonia luntii Baker, Bull. Misc.
Pl. Cap. 1[viii]: 80. 1794, not: Zygophyllum capense Inform. Kew 1894: 330. 1894.
Lam. Etymology: Named for the former genus Zygophyllum mahranum (Beier) Christenh. & Byng,
Augea, a synonym of Zygophyllum comb. nov. Basionym: Fagonia mahrana Beier,
Zygophyllum bruguieri (DC.) Christenh. & Byng, comb. Nordic J. Bot. 21: 451. 2001 publ. 2002.
nov. Basionym: Fagonia bruguieri DC., Prodr. 1: 704. Zygophyllum mayanum Christenh. & Byng, nom. nov.
1824. Basionym: Fagonia mayana Schltdl, Bot. Zeitung
Zygophyllum californicum (Benth.) Christenh. & Byng, (Berlin) 9: 844 1851. Replaced synonym: Fagonia
comb. nov. Basionym: Fagonia californica Benth., latifolia Delile, Descr. Egypte, Hist. Nat.: 230. 1813,
Bot. Voy. Sulphur: 10. 1844. not: Zygophyllum latifolium Schrenk.
Zygophyllum charoides (Chiov.) Christenh. & Byng, Zygophyllum minutistipulum (Engl.) Christenh. &
comb. nov. Basionym: Fagonia charoides Chiov., Fl. Byng, comb. nov. Basionym: Fagonia minutistipula
Somala 1: 112. 1929. Engl., Bot. Jahrb. Syst. 10: 33. 1889.
Zygophyllum chilense (Hook. & Arn.) Christenh. & Zygophyllum molle (Delile) Christenh. & Byng, comb.
Byng, comb. nov. Basionym: Fagonia chilensis Hook. nov. Basionym: Fagonia mollis Delile, Descr. Egypte,
& Arn., Bot. Misc. 3: 165. 1833. Hist. Nat.: 220. 1813.
Zygophyllum creticum (L.) Christenh. & Byng, comb. Zygophyllum mongolicum (Maxim.) Christenh. &
nov. Basionym: Fagonia cretica L., Sp. Pl.: 386. 1753. Byng, comb. nov. Basionym: Tetraena mongolica
Zygophyllum densispinum (Beier & Thulin) Christenh. Maxim., Enum. Pl. Mongolia: 129. 1889.
& Byng, comb. nov. Basionym: Fagonia densispina Zygophyllum olivieri (DC.) Christenh. & Byng, comb.
Beier & Thulin, Syst. Biodivers. 3: 235. 2005. nov. Basionym: Fagonia olivieri DC., Prodr. 1: 704.
Zygophyllum densum (I.M.Johnst.) Christenh. & Byng, 1824.
comb. nov. Basionym: Fagonia densa I.M.Johnst., Zygophyllum orientale (C.Presl) Christenh. & Byng,
Proc. Calif. Acad. Sci. IV, 12: 1052. 1924. comb. nov. Basionym: Fagonia orientalis C.Presl,
Zygophyllum glutinosum (Delile) Christenh. & Byng, Delic. Prag.: 23. 1822.
comb. nov. Basionym: Fagonia glutinosa Delile, Zygophyllum pachyacanthum (Rydb.) Christenh. &
Descr. Egypte, Hist. Nat.: 230. 1813. Byng, comb. nov. Basionym: Fagonia pachyacantha
Zygophyllum gypsophilum (Beier & Thulin) Christenh. Rydb., N. Amer. Fl. 25: 105. 1910.
& Byng, comb. nov. Basionym: Fagonia gypsophila Zygophyllum palmeri (Vasey & Rose) Christenh. &
Beier & Thulin, Nordic J. Bot. 21: 452. 2001 publ. Byng, comb. nov. Basionym: Fagonia palmeri Vasey
2002. & Rose, Contr. U.S. Natl. Herb. 1: 82. 1890.
Zygophyllum hadramauticum (Beier & Thulin) Zygophyllum paulayanum (J.Wagner & Vierh.)
Christenh. & Byng, comb. nov. Basionym: Fagonia Christenh. & Byng, comb. nov. Basionym: Fagonia
hadramautica Beier & Thulin, Syst. Biodivers. 3: 238. paulayana J.Wagner & Vierh., Oesterr. Bot. Z. 54: 34.
2005. 1904.
Zygophyllum harpago (Emb. & Maire) Christenh. & Zygophyllum scabrum (Forssk.) Christenh. & Byng,
Byng, comb. nov. Basionym: Fagonia harpago Emb. comb. nov. Basionym: Fagonia scabra Forssk., Fl.
& Maire, Bull. Soc. Hist. Nat. Afrique N. 22: 284. Aegypt.-Arab.: 88. 1775.
1931. Zygophyllum scoparium (Brandegee) Christenh. &
Zygophyllum indicum (Burm.f.) Christenh. & Byng, Byng, comb. nov. Basionym: Fagonia scoparia
comb. nov. Basionym: Fagonia indica Burm.f., Fl. Brandegee, Univ. Calif. Publ. Bot. 4: 181. 1911.
Indica: 102. 1768. Zygophyllum spinosissimum (Blatt. & Hallb.) Christenh.
Zygophyllum laeve (Standl.) Christenh. & Byng, comb. & Byng, comb. nov. Basionym: Fagonia spinosissima
nov. Basionym: Fagonia laevis Standl., Proc. Biol. Blatt. & Hallb., J. Indian Bot. 1: 89. 1919.
Soc. Wash. 24: 249. 1911. Zygophyllum subinerme (Boiss.) Christenh. & Byng,
Zygophyllum lahovarii (Volkens & Schweinf.) comb. nov. Basionym: Fagonia subinermis Boiss.,
Christenh. & Byng, comb. nov. Basionym: Fagonia Diagn. Pl. Orient. 1: 62. 1843.
lahovarii Volkens & Schweinf., Liste Pl. Somalis: 204. Zygophyllum villosum (D.M.Porter) Christenh. & Byng,
1897. comb. nov. Basionym: Fagonia villosa D.M.Porter,
Zygophyllum latistipulatum (Beier & Thulin) Christenh. Contr. Gray Herb. 192: 120. 1963.
& Byng, comb. nov. Basionym: Fagonia latistipulata Zygophyllum zilloides (Humbert) Christenh. & Byng,

93
comb. nov. Basionym: Fagonia zilloides Humbert, Wiss. Z. Martin-Luther-Univ. Halle-Wittenberg,

Bull. Soc. Hist. Nat. Afrique N. 18: 144. 1927. Math.-Naturwiss. Kl. 15: 532. 1966.
Pyrus ailanthoides M.F.Fay & Christenh., nom. nov.
142. Polygalaceae. Reference: Pastore et al. (2017). Replaced synonym: Sorbus hupehensis C.K.Schneid.,
Bull. Herb. Boissier, sér. 2, 6: 316. 1906, not:
Note: Balgoya Morat & Meijden is embedded in
Pyrus hupehensis Pamp. Note: Pyrus hupehensis
Moutabea Aubl. and is hereby transferred.
(C.K.Schneid.) Bean is an illegitimate homonym.
Moutabea pacifica (Morat & Meijden) Byng & Etymology: The foliage of this species resembles
Christenh., comb. nov. Basionym: Balgoya pacifica Ailanthes (Simaroubaceae), hence the name.
Morat & Meijden, Bull. Mus. Natl. Hist. Nat., B,
Pyrus aitchisonii (C.K.Schneid.) M.F.Fay & Christenh.,
Adansonia Sér. 4, 13: 3. 1991.
comb. nov. Basionym: Cotoneaster aitchisonii
C.K.Schneid., Illustr. Handb. Laubholzk. 1: 749. 1906.
143. Rosaceae. Reference: Li et al. (2012). Pyrus alabaster M.F.Fay & Christenh., nom. nov.
Note: Sorbus L. is polyphyletic and even though several Replaced synonym: Eriobotrya wardii C.E.C.Fisch.,
solutions have been suggested, it complicates the Bull. Misc. Inform. Kew 1929(6): 205. 1929, not:
taxonomy, with many unfamiliar genera and with Pyrus wardii (W.W.Sm.) M.F.Fay & Christenh.
numerous complex intergeneric hybrids. To show Etymology: This species have flower petals that are
that these are closely related taxa, Pyrus L. is hereby fragile and white, like alabaster, hence the name.
expanded to include most of Maleae, e.g. the genera Pyrus alashanensis (J.Fryer & B.Hylmö) M.F.Fay &
Aronia Medik., Chaenomeles Lindl., Chamaemeles Christenh., comb. nov. Basionym: Cotoneaster
Lindl., Chamaemespilus Medik., Cotoneaster Medik., alashanensis J.Fryer & B.Hylmö, Cotoneasters
Cydonia Mill., Docynia Decne., Eriobotrya Lindl., Compreh. Guide: 307. 2009.
Heteromeles M.Roem., Malus Mill., Micromeles
Pyrus alatavica (M.Popov) M.F.Fay & Christenh., comb.
Decne., Osteomeles Lindl., Photinia Lindl., Pourthiaea
nov. Basionym: Cotoneaster alatavicus M.Popov,
Decne., Pseudocydonia (C.K.Schneid.) C.K.Schneid.,
Bull. Soc. Imp. Naturalistes Moscou 47: 86. 1938.
Rhaphiolepis Lindl., Sorbus s.l. and Stranvaesia Lindl.
returning it to the Linnaean concept of the genus Pyrus alaunica (Golitsin) M.F.Fay & Christenh., comb.
Pyrus. nov. Basionym: Cotoneaster alaunicus Golitsin,
Novosti Sist. Vyssh. Rast. 1964: 145. 1964.
Pyrus ×abscondita (Kovanda) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus ×abscondita Kovanda, Pyrus albensis (M.Lepší, K.Boublík, P.Lepší & P.Vít)
Verh. Zool.-Bot. Ges. Österreich 133: 339. 1996. (= P. M.F.Fay & Christenh., comb. nov. Basionym: Sorbus
aucuparia × P. danubialis). albensis M.Lepší, K.Boublík, P.Lepší & P.Vít, Preslia
81: 76. 2009.
Pyrus acuminata (Lindl.) M.F.Fay & Christenh., comb.
nov. Basionym: Cotoneaster acuminatus Lindl., Pyrus albokermesina (J.Fryer & B.Hylmö) M.F.Fay
Trans. Linn. Soc. London 13: 101. 1822. & Christenh., comb. nov. Basionym: Cotoneaster
albokermesinus J.Fryer & B.Hylmö, New Plantsman
Pyrus acutifolia (Turcz.) M.F.Fay & Christenh., comb.
5: 132. 1998.
nov. Basionym: Cotoneaster acutifolius Turcz., Bull.
Soc. Imp. Naturalistes Moscou 4: 190. 1832. Pyrus albopilosa (T.T.Yu & L.T.Lu) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus albopilosa T.T.Yu &
Pyrus adamii (Kárpáti) M.F.Fay & Christenh., comb.
L.T.Lu, Acta Phytotax. Sin. 18: 495. 1980.
nov. Basionym: Sorbus adamii Kárpáti, Hung. Acta
Biol. 1: 112. 1949, as ‘adami’. Pyrus albovii (Zinserl.) M.F.Fay & Christenh., comb.
nov. Basionym: Sorbus albovii Zinserl., Fl. URSS 9:
Pyrus adeana (N.Mey.) M.F.Fay & Christenh., comb.
390. 1939.
nov. Basionym: Sorbus adeana N.Mey., Ber. Bayer.
Bot. Ges. Sonderband : 139. 2005. Pyrus algoviensis (N.Mey.) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus algoviensis N.Mey.,
Pyrus admonitor (M.Proctor) M.F.Fay & Christenh.,
Ber. Bayer. Bot. Ges. 86: 228. 2016.
comb. nov. Basionym: Sorbus admonitor M.Proctor,
Watsonia 27: 207. 2009. Pyrus allochroa (Pojark.) M.F.Fay & Christenh., comb.
nov. Basionym: Cotoneaster allochrous Pojark., Bot.
Pyrus adpressa (Bois) M.F.Fay & Christenh., comb. nov.
Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R.
Basionym: Cotoneaster adpressus Bois, Bull. Soc.
21: 171. 1961. Replaced synonym: Cotoneaster
Bot. France 51: 149. 1904.
fontanesii Spach var. songoricus Regel, Trudy Imp. S.-
Pyrus affinis (Lindl.) M.F.Fay & Christenh., comb. nov. Peterburgsk. Bot. Sada 2: 313. 1873, as ‘soongarica’,
Basionym: Cotoneaster affinis Lindl., Trans. Linn. Soc. not: Pyrus songorica Fisch. ex Juz., and orthographic
London 13: 101. 1822. variant.
Pyrus afghanica (G.Klotz) M.F.Fay & Christenh., comb. Pyrus alma M.F.Fay & Christenh., nom. nov. Replaced
nov. Basionym: Cotoneaster afghanicus G.Klotz,

synonym: Sorbus fedorovii Zaik., Bot. Zhurn. (Sargent) 1(3): 465. 1913.
(Moscow & Leningrad) 59(11): 1605. 1974, not: Pyrus aphroditae M.F.Fay & Christenh., nom. nov.
Pyrus fedorovii Kuth. Etymology: From the Kazakh Replaced synonym: Cotoneaster saxatilis Pojark.,
word алма, apple. Not. Syst. Inst. Bot. Acad. Sci. USSR 8: 7. 1938, not:
Pyrus alnifrons (Kovanda) M.F.Fay & Christenh., comb. Pyrus saxatilis Schltdl. ex Ledeb. Etymology: Named
nov. Basionym: Sorbus alnifrons Kovanda, Verh. for Ἀφροδίτη, Greek goddess of love and beauty.
Zool.-Bot. Ges. Österreich 133: 356. 1996. The myth goes that at the marriage of Peleus and
Pyrus altaica (Koehne) M.F.Fay & Christenh., comb. Thetis, Eris, the goddess of discord, was not invited.
nov. Basionym: Sorbus altaica Koehne, Repert. Spec. She was not amused, so she gave Paris a golden
Nov. Regni Veg. 10: 505. 1912. apple inscribed with the word καλλίστῃ (kallistēi,
“for the fairest”). Aphrodite, Hera, and Athena all
Pyrus ×ambigua (Decne.) M.F.Fay & Christenh., comb.
claimed to be the fairest, and thus the rightful owner
nov. Basionym: Aria ambigua Decne., Nouv. Arch.
of the apple.
Mus. Hist. Nat. 10: 165. 1874.
Pyrus apiculata (Rehder & E.H.Wilson) M.F.Fay &
Pyrus amici-petri (Mikoláš) M.F.Fay & Christenh.,
Christenh., comb. nov. Basionym: Cotoneaster
comb. nov. Basionym: Sorbus amici-petri Mikoláš,
apiculatus Rehder & E.H.Wilson, Pl. Wilson. (Sargent)
Thaiszia 13: 128. 2004.
1: 156. 1912.
Pyrus amphidoxa (C.K.Schneid.) M.F.Fay & Christenh.,
Pyrus arbuscula (G.Klotz) M.F.Fay & Christenh.,
comb. nov. Basionym: Stranvaesia amphidoxa
comb. nov. Basionym: Cotoneaster arbusculus
C.K.Schneid., Bull. Herb. Boissier Ser. 2, 6: 319. 1906.
G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-
Pyrus anadyrensis (Komarov) M.F.Fay & Christenh., Naturwiss. Reihe 17: 337. 1968.
comb. nov. Basionym: Sorbus anadyrensis Komarov,
Pyrus arctostaphylos M.F.Fay & Christenh., nom.
Bot. Mater. Gerb. Glavn. Bot. Sada R.S.F.S.R. 2: 134.
nov. Replaced synonym: Cotoneaster congestus
1921.
Baker, Saunder’s Ref. Bot. 1: pl. 51. 1869, not:
Pyrus anatolii (Teimurov & Taisumov) M.F.Fay & Pyrus congesta Rehder. Etymology: This species
Christenh., comb. nov. Basionym: Cotoneaster superficically resembles a species of Arctostaphylos
anatolii Teimurov & Taisumov, Byull. Moskovsk. (Ericaceae).
Obshch. Isp. Prir., Otd. Biol. 115: 71. 2010.
Pyrus argentea (G.Klotz) M.F.Fay & Christenh., comb.
Pyrus andersonii (G.N.Jones) M.F.Fay & Christenh., nov. Basionym: Cotoneaster argenteus G.Klotz,
comb. nov. Basionym: Sorbus andersonii G.N.Jones, Wiss. Z. Martin-Luther-Univ. Halle-Wittenberg,
Rhodora 47: 220. 1945. Math.-Naturwiss. Reihe 12: 775. 1963.
Pyrus andina M.F.Fay & Christenh., nom. nov. Replaced Pyrus arguta (Wall. ex Lindl.) M.F.Fay & Christenh.,
synonym: Cratageus ferruginea Pers., Syn. Pl. 2: 37. comb. nov. Basionym: Photinia arguta Wall. ex
1806, not: Pyrus ferruginea Hook.f. Etymology: This Lindl., Edwards’s Bot. Reg. 23: sub t. 1956. 1837.
species occurs in the Andes.
Pyrus armena (Pojark.) M.F.Fay & Christenh., comb.
Pyrus andreanszkyana (Kárpáti) M.F.Fay & Christenh., nov. Basionym: Cotoneaster armenus Pojark., Not.
comb. nov. Basionym: Sorbus andreanszkyana Syst. Inst. Bot. Acad. Sci. USSR 17: 202. 1955.
Kárpáti, Agrártud. Egyet. Kert- Szölögazdaságtud.
Pyrus armeniaca (Hedl.) M.F.Fay & Christenh., comb.
Karának Évk. 1: 34. 1950.
nov. Basionym: Sorbus armeniaca Hedl., Kongl.
Pyrus angusta (T.T.Yu ex T.T.Yu & K.C.Kuan) M.F.Fay Svenska Vetensk.-Akad. Handl. 35: 69. 1901.
& Christenh., comb. nov. Basionym: Cotoneaster
Pyrus ×arnoldiana (Rehder) M.F.Fay & Christenh.,
salicifolius Franch. var. angustus T.T.Yu ex T.T.Yu &
comb. nov. Basionym: Sorbus ×arnoldiana Rehder, J.
K.C.Kuan, Acta Phytotax. Sin. 8: 219. 1963.
Arnold Arbor. 2: 42. 1920. Note: = Pyrus aucuparia ×
Pyrus angustissima (Hook.f.) M.F.Fay & Christenh., Pyrus discolor.
comb. nov. Basionym: Eriobotrya angustissima
Pyrus aronioides (Rehder) M.F.Fay & Christenh., comb.
Hook.f., Fl. Brit. India 2: 372. 1878.
nov. Basionym: Sorbus aronioides Rehder, Pl. Wilson.
Pyrus anlungensis (T.T.Yu) M.F.Fay & Christenh., comb. (Sargent) 2(2): 268. 1915.
nov. Basionym: Photinia anlungensis T.T.Yu, Acta
Pyrus arvonicola (P.D.Sell) M.F.Fay & Christenh., comb.
Phytotax. Sin. 8: 228. 1963.
nov. Basionym: Sorbus arvonicola P.D.Sell in Sell &
Pyrus antoninae (Juz. ex Orlova) M.F.Fay & Christenh., Murrell, Fl. Gr. Brit. Ireland 2: 522. 2014.
comb. nov. Basionym: Cotoneaster antoninae Juz. ex
Pyrus ascendens (Flinck & B.Hylmö) M.F.Fay &
Orlova, Bot. Mater. Gerb. Bot. Inst. Komarova Akad.
Nauk S.S.S.R. 13: 33. 1950.
ascendens Flinck & B.Hylmö, Bot. Not. (Lund) 119:
Pyrus aperta (Koehne) M.F.Fay & Christenh., comb.
nov. Basionym: Sorbus aperta Koehne, Pl. Wilson.

95
453. 1966. Replaced synonym: Cotoneaster atropurpureus

Pyrus asiatica (Nakai) M.F.Fay & Christenh., comb. nov. Flinck & B.Hylmö, Watsonia 18: 311. 1991, not:
Basionym: Malus asiatica Nakai, Icon. Pl. Koisikav. 3: Pyrus atropurpurea (Britton) L.H.Bailey. Etymology:
19. 1915. This species has dark coloured branches, hence the
name, from Latin atro, dark and ramosus, branched.
Pyrus assadii (Khat.) M.F.Fay & Christenh., comb. nov.
Basionym: Cotoneaster assadii, Iran. J. Bot. 4: 116. Pyrus atrosanguinea (T.T.Yu & Tsai) M.F.Fay &
1988. Christenh., comb. nov. Basionym: Sorbus
atrosanguinea T.T.Yu & Tsai, Bull. Fan Mem. Inst. Biol.
Pyrus assamensis (G.Klotz) M.F.Fay & Christenh.,
Bot. 7: 119. 1936.
comb. nov. Basionym: Cotoneaster assamensis
G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.- Pyrus atrovinacea (J.Fryer & B.Hylmö) M.F.Fay &
Naturwiss. Reihe 21: 996. 1972. Christenh., comb. nov. Basionym: Cotoneaster
atrovinaceus J.Fryer & B.Hylmö, Cotoneasters
Pyrus asterope M.F.Fay & Christenh., nom. nov.
Compreh. Guide : 307. 2009.
Replaced synonym: Crataegus integrifolia Roxb.,
Fl. Ind. 2: 509. 1832, not: Pyrus integrifolia (Lindl.) Pyrus atrovirens (J.Fryer & B.Hylmö) M.F.Fay &
M.F.Fay & Christenh. Etymology: Named for Christenh., comb. nov. Basionym: Cotoneaster
Ἀστεροπή, a Hesperid nymph. Her name means atrovirens J.Fryer & B.Hylmö, Cotoneasters Compreh.
‘starry faced’, but also means ‘lightning’. Guide : 307. 2009.
Pyrus astrophora (J.Fryer & E.C.Nelson) M.F.Fay & Pyrus atuntzensis (J.Fryer & B.Hylmö) M.F.Fay &
Christenh., comb. nov. Basionym: Cotoneaster Christenh., comb. nov. Basionym: Cotoneaster
astrophoros J.Fryer & E.C.Nelson, Glasra 2: 128. atuntzensis J.Fryer & B.Hylmö, Cotoneasters
1995. Compreh. Guide : 308. 2009.
Pyrus ataensis (J.Fryer & B.Hylmö) M.F.Fay & Pyrus aurantiaca (J.Fryer & B.Hylmö) M.F.Fay &
Christenh., comb. nov. Basionym: Cotoneaster Christenh., comb. nov. Basionym: Cotoneaster
ataensis J.Fryer & B.Hylmö, Cotoneasters Compreh. aurantiacus J.Fryer & B.Hylmö, Cotoneasters
Guide : 307 (221; pl. 130). 2009. Compreh. Guide : 308. 2009.
Pyrus atalantae M.F.Fay & Christenh., nom. nov. Pyrus aurelia-aquensis M.F.Fay & Christenh., nom.
Replaced synonym: Photinia lanuginosa T.T.Yu, Acta nov. Replaced synonym: Sorbus badensis Düll in
Phytotax. Sin. 8: 227. 1963, not: Pyrus lanuginosa N.Mey., Ber. Bayer. Bot. Ges. Sonderband: 131. 2005,
DC. Etymology: Named for Ἀταλάντη, a virgin not: Pyrus badensis hort. ex K.Koch Etymology:
huntress, who did not want to get married, but got Named for Baden-Baden in Germany, or Aurelia-
distracted by three golden apples and lost her race Aquensis in Latin.
to Hippomanes, whom she then had to marry. Pyrus avalon M.F.Fay & Christenh., nom. nov. Replaced
Pyrus athenae M.F.Fay & Christenh., nom. nov. synonym: Eriobotrya lasiogyna Franch., Pl. Delavay.
Replaced synonym: Hiptage cavaleriei H.Lév., 1: 225. 1890, not: Pyrus lasiogyna Koidz. Etymology:
Repert. Spec. Nov. Regni Veg. 10: 372. 1912, not: from Welsh Ynys Afallon or Ynys Afallach, meaning
Pyrus cavaleriei H.Lév. Etymology: Named for ‘the island of apple trees’ as in ‘Avalon’ the mythical
Ἀθηνᾶ, Greek goddess of wisdom and war. The island featured in the legends of King Arthur.
myth goes that at the marriage of Peleus and Thetis, Pyrus avonensis (T.C.G.Rich) M.F.Fay & Christenh.,
Eris, the goddess of discord, was not invited. She comb. nov. Basionym: Sorbus avonensis T.C.G.Rich,
was not amused, so she gave Paris a golden apple Watsonia 27: 370. 2009.
inscribed with the word καλλίστῃ (kallistēi, “for the Pyrus bacillaris (Wall. ex Lindl.) M.F.Fay & Christenh.,
fairest”). Aphrodite, Hera, and Athena all claimed comb. nov. Basionym: Cotoneaster bacillaris Wall. ex
to be the fairest, and thus the rightful owner of the Lindl., Edward’s Bot. Reg. 15: t. 1229. 1829.
apple. Note: This was described in Hiptage Gaertn. Pyrus bakeri (G.Klotz) M.F.Fay & Christenh., comb.
(Malpighiaceae), but later transferred to Eriobotrya nov. Basionym: Cotoneaster bakeri G.Klotz, Wiss. Z.
Lindl. as E. cavaleriei (H.Lév.) Rehder. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe
Pyrus atlantica (G.Klotz) M.F.Fay & Christenh., comb. 21: 970. 1972. Replaced synonym: Cotoneaster
nov. Basionym: Cotoneaster atlanticus G.Klotz, Wiss. sanguineus T.T.Yu, Bull. Brit. Mus. (Nat. Hist.), Bot. 1:
Z. Martin-Luther-Univ. Halle-Wittenberg, Math.- 130. 1954, not: Pyrus sanguinea Pursh.
Naturwiss. Reihe 12: 759. 1963. Pyrus bakonyensis (Jáv.) M.F.Fay & Christenh.,
Pyrus atrimontis (Bernátová & Májovský) M.F.Fay & comb. nov. Basionym: Sorbus franconica Bornm. f.
Christenh., comb. nov. Basionym: Sorbus atrimontis bakonyensis Jáv., Magyar Bot. Lapok 25: 87. 1927.
Bernátová & Májovský, Biologia (Bratislava) 58: 786. Pyrus balatonica (Kárpáti) M.F.Fay & Christenh., comb.
2003. nov. Basionym: Sorbus balatonica Kárpáti, Hung.
Pyrus atroramosa M.F.Fay & Christenh., nom. nov.

A B
C D
E F
Figure 12: Angiosperms (Eudicots). A, Pyrus bibas (Lour.) M.F.Fay & Christenh.; B, Pyrus lancastriensis (E.F.Warb.)
M.F.Fay & Christenh.; C, Pyrus mandshurica (Maxim.) M.F.Fay & Christenh.; D, Pyrus microphylla (Wall. ex Lindl.)
M.F.Fay & Christenh.; E, Pyrus morrisonensis (Hayata) M.F.Fay & Christenh.; and F, Pyrus photinia M.F.Fay &
Christenh.. All images © Maarten Christenhusz.

97
Acta Biol. 1: 121. 1949. C.K.Schneid, Ill. Handb. Laubholzk. 1: 709. 1906.
Pyrus baldaccii (C.K.Schneid.) M.F.Fay & Christenh., Pyrus bibas (Lour.) M.F.Fay & Christenh., comb. nov.
comb. nov. Basionym: Sorbus umbellata (Desf.) Basionym: Crataegus bibas Lour., Fl. Cochinch. 1:
Fritsch var. baldaccii C.K.Schneid., Ill. Handb. 319. 1790. Replaced synonym: Mespilus japonica
Laubholzk. 1: 691. 1906. Thunb., Nova Acta Regiae Soc. Sci. Upsal. 3: 208.
Pyrus barabitsii (C.Németh) M.F.Fay & Christenh., 1780; syn.: Eriobotrya japonica (Thunb.) Lindl., not:
comb. nov. Basionym: Sorbus barabitsii C.Németh, Pyrus japonica Thunb. (Figure 12A)
Acta Bot. Hung. 54: 138. 2012. Pyrus bisramianus (G.Klotz) M.F.Fay & Christenh.,
Pyrus barrandienica (P.Vít, M.Lepší & P.Lepší) M.F.Fay comb. nov. Basionym: Cotoneaster bisramianus
& Christenh., comb. nov. Basionym: Sorbus G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-
barrandienica P.Vít, M.Lepší & P.Lepší, Preslia 84: 82. Naturwiss. Reihe 21: 994. 1972, as ‘bis-ramianus’.
2012. Pyrus bissetii (McAll.) M.F.Fay & Christenh., comb.
Pyrus barthae (Kárpáti) M.F.Fay & Christenh., comb. nov. Basionym: Sorbus bissetii McAll., Gen. Sorbus:
nov. Basionym: Sorbus barthae Kárpáti, Agrártud. 234. 2005.
Egyet. Kert- Szölögazdaságtud. Karának Évk. 1(14): Pyrus blinii (H.Lév.) M.F.Fay & Christenh., comb. nov.
37. 1950. Basionym: Cotoneaster blinii H.Lév., Cat. Pl. Yun-
Pyrus beauverdiana (C.K.Schneid.) M.F.Fay & Nan: 229. 1917.
Christenh., comb. nov. Basionym: Photinia Pyrus bodajkensis (Barabits) M.F.Fay & Christenh.,
beauverdiana C.K.Schneid., Bull. Herb. Boissier Ser. comb. nov. Basionym: Sorbus bodajkensis Barabits,
2, 6: 319. 1906. Tilia 13: 19. 2007.
Pyrus beckii (C.K.Schneid.) M.F.Fay & Christenh., Pyrus bohemica (Kovanda) M.F.Fay & Christenh.,
comb. nov. Basionym: Photinia beckii C.K.Schneid., comb. nov. Basionym: Sorbus bohemica Kovanda,
Ill. Handb. Laubholzk. 1: 707. 1906. Acta Univ. Carol., Biol. 1961: 77. 1961.
Pyrus beijingensis M.F.Fay & Christenh., nom. nov. Pyrus boisiana (G.Klotz) M.F.Fay & Christenh., comb.
Replaced synonym: Cotoneaster acutifolius Turcz. nov. Basionym: Cotoneaster boisianus G.Klotz, Wiss.
f. pekinensis Koehne, Deut. Dendrol.: 225. 1893, Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss.
not: Pyrus pekinensis (Koehne) Cardot. Etymology: Reihe 21: 987. 1972.
Named for Beijing, the modern name for the capital Pyrus boissieri (C.K.Schneid.) M.F.Fay & Christenh.,
of China formerly known as Peking. comb. nov. Basionym: Sorbus boissieri C.K.Schneid.,
Pyrus beimashanensis (J.Fryer & B.Hylmö) M.F.Fay Bull. Herb. Boissier Ser. 2, 6: 312. 1906.
& Christenh., comb. nov. Basionym: Cotoneaster Pyrus bombycilla M.F.Fay & Christenh., nom.
beimashanensis J.Fryer & B.Hylmö, Cotoneasters nov. Replaced synonym: Cotoneaster amoenus
Compreh. Guide: 308. 2009. E.H.Wilson, Gard. Chron. 51: 2. 1912, not: Pyrus
Pyrus belladonna M.F.Fay & Christenh., nom. nov. amoena Poit. & Turpin. Etymology: Named for the
Replaced synonym: Cotoneaster altaicus J.Fryer & waxwing (Bombycilla garrulus L.), a bird that likes
B.Hylmö, Cotoneasters Compreh. Guide: 307. 2009, fruits of this and other small-fruited Pyrus species.
not: Pyrus altaica (Koehne) M.F.Fay & Christenh. Pyrus borbasii (Jáv.) M.F.Fay & Christenh., comb. nov.
Etymology: This species has black fruits that Basionym: Sorbus borbasii Jáv., Bot. Közlem. 14: 99.
superficially resemble those of deadly nightshade, 1915.
Atropa belladonna (Solanaceae). The name means Pyrus borealichinensis (Hurus.) M.F.Fay & Christenh.,
‘beautiful lady’. comb. nov. Basionym: Cotoneaster multiflorus
Pyrus bengalensis (Roxb.) M.F.Fay & Christenh., comb. Bunge var. borealichinensis Hurus., Acta Phytotax.
nov. Basionym: Mespilus bengalensis Roxb., Fl. Ind. Geobot. 13: 230. 1943.
2: 510. 1832. Pyrus borosiana (Kárpáti) M.F.Fay & Christenh., comb.
Pyrus benthamiana (Hance) M.F.Fay & Christenh., nov. Basionym: Sorbus borosiana Kárpáti, Agrártud.
comb. nov. Basionym: Photinia benthamiana Hance, Egyet. Kert- Szölögazdaságtud. Karának Közlem. 12:
Ann. Sci. Nat., Bot. sér. 5, 5: 213. 1866. 144. 1948.
Pyrus berberidifolia (Rehder & E.H.Wilson) M.F.Fay Pyrus bosniaca (Hajrud., Frajman, Schönsw. & Bogunić)
& Christenh., comb. nov. Basionym: Photinia M.F.Fay & Christenh., comb. nov. Basionym: Sorbus
berberidifolia Rehder & E.H.Wilson, Pl. Wilson. bosniaca Hajrud., Frajman, Schönsw. & Bogunić, Bot.
(Sargent) 1(2): 191. 1912. J. Linn. Soc. 178: 682. 2015.
Pyrus bergerae (C.K.Schneid.) M.F.Fay & Christenh., Pyrus bradyi (E.C.Nelson & J.Fryer) M.F.Fay &
comb. nov. Basionym: Photinia bergerae Christenh., comb. nov. Basionym: Cotoneaster

bradyi E.C.Nelson & J.Fryer, Glasra 2: 131. 1995. ex Lind., Edward’s Bot. Reg. 15: t. 1229. 1829.
Pyrus brandisii (G.Klotz) M.F.Fay & Christenh., comb. Pyrus caeruleomontana (Bernátová & Májovský)
nov. Basionym: Cotoneaster brandisii G.Klotz, Wiss. M.F.Fay & Christenh., comb. nov. Basionym: Sorbus
Z. Martin-Luther-Univ. Halle-Wittenberg, Math.- caeruleomontana Bernátová & Májovský, Biologia
Naturwiss. Reihe 15: 537. 1966. (Bratislava) 58: 786. 2003.
Pyrus breviramea (Rehder & E.H.Wilson) M.F.Fay Pyrus calcicola (W.B.Liao & W.Guo) M.F.Fay &
& Christenh., comb. nov. Basionym: Cotoneaster Christenh., comb. nov. Basionym: Sorbus calcicola
brevirameus Rehder & E.H.Wilson, Pl. Wilson. W.B.Liao & W.Guo, Phytotaxa 261: 261. 2016.
(Sargent) 1: 177. 1912. Pyrus californica (Greene) M.F.Fay & Christenh., comb.
Pyrus brickellii (J.Fryer & B.Hylmö) M.F.Fay & nov. Basionym: Sorbus californica Greene, Pittonia
Christenh., comb. nov. Basionym: Cotoneaster 4: 131. 1900.
brickellii J.Fryer & B.Hylmö, New Plantsman 8: 236. Pyrus decaisnei M.F.Fay & Christenh., nom. nov.
2001. Replaced synonym: Pourthiaea calleryana Decne.,
Pyrus bristoliensis (Wilmott) M.F.Fay & Christenh., Nouv. Arch. Mus. Par. Ser. 1, 10: 147. 1874., not:
comb. nov. Basionym: Sorbus bristoliensis Wilmott, Pyrus calleryana Decne. Etymology: Named for
Proc. Linn. Soc. Lond. 146: 76. 1934. Joseph Decaisne who originally described this
Pyrus britannica M.F.Fay & Christenh., nom. nov. species.
Replaced synonym: Sorbus anglica Hedl., Nyt Mag. Pyrus callistei M.F.Fay & Christenh., nom. nov.
Naturvidensk. 52: 258. 1914, not: Pyrus anglica Poit. Replaced synonym: Cotoneaster rockii G.Klotz, Wiss.
& Turpin (note: Pyrus anglica (Hedl.) Druce is an Z. Martin-Luther-Univ. Halle-Wittenberg, Math.-
illegitimate homonym). Etymology: This is a species Naturwiss. Reihe 12: 775. 1963, not: Pyrus rockii
found in Britain, hence the name. (Rehder) M.F.Fay & Christenh. Etymology: In Greek
Pyrus buekkensis (Soó) M.F.Fay & Christenh., comb. mythology (Judgment of Paris), Eris was not invited
nov. Basionym: Sorbus aria L. subsp. buekkensis Soó, at the marriage of Peleus and Thetis. She was not
Tisia 2: 222. 1937, as ’bükkensis’. amused, so she gave Paris a golden apple inscribed
with the word καλλίστῃ (kallistēi, “for the fairest”) to
Pyrus bullatus (Bois) M.F.Fay & Christenh., comb. nov.
take to the wedding and give to the most beautiful
Basionym: Cotoneaster bullatus Bois, Bull. Soc. Bot.
of all goddesses, which did not go down well.
France 51: 153. 1904.
Pyrus callosa (Chun ex K.C.Kuan) M.F.Fay & Christenh.,
Pyrus bulleyana (McAll.) M.F.Fay & Christenh., comb.
comb. nov. Basionym: Photinia callosa Chun ex
nov. Basionym: Sorbus bulleyana McAll., Gen.
K.C.Kuan, Acta Phytotax. Sin. 8: 229. 1963.
Sorbus: 232. 2005.
Pyrus calocarpa (Rehder & E.H.Wilson) M.F.Fay &
Pyrus bumthangensis (J.Fryer & B.Hylmö) M.F.Fay
& Christenh., comb. nov. Basionym: Cotoneaster
multiflorus Bunge var. calocarpus Rehder &
bumthangensis J.Fryer & B.Hylmö, Cotoneasters
E.H.Wilson, Pl. Wilson. (Sargent) 1: 170. 1912.
Pyrus caloneura (Stapf) M.F.Fay & Christenh., comb.
Pyrus burmanica (G.Klotz) M.F.Fay & Christenh.,
nov. Basionym: Micromeles caloneura Stapf, Bull.
comb. nov. Basionym: Cotoneaster burmanicus
Misc. Inform. Kew 1910: 192. 1910
Naturwiss. Reihe 21: 1004. 1972. Pyrus cambrensis (M.Proctor) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus cambrensis M.Proctor,
Pyrus burtonsmithiorum (Rushforth) M.F.Fay
Watsonia 27: 208. 2009.
& Christenh., comb. nov. Basionym: Sorbus
burtonsmithiorum Rushforth, Int. Dendrol. Soc. Year Pyrus cambrica (J.Fryer & B.Hylmö) M.F.Fay &
Book 2009: 85. 2010. Christenh., comb. nov. Basionym: Cotoneaster
cambrica J.Fryer & B.Hylmö, Watsonia 20: 62. 1994.
Pyrus busambarensis (G.Castellano, P.Marino,
Raimondo & Spadaro) M.F.Fay & Christenh., comb. Pyrus camilli-schneideri (Pojark.) M.F.Fay & Christenh.,
nov. Basionym: Sorbus busambarensis G.Castellano, comb. nov. Basionym: Cotoneaster camilli-schneideri
P.Marino, Raimondo & Spadaro, Pl. Biosystems Pojark., Not. Syst. Inst. Bot. Acad. Sci. USSR 17: 180.
146(Suppl.): 339. 2012. 1955.
Pyrus buschiana (Zinserl.) M.F.Fay & Christenh., comb. Pyrus campanulata (J.Fryer & B.Hylmö) M.F.Fay &
nov. Basionym: Sorbus buschiana Zinserl., Fl. URSS Christenh., comb. nov. Basionym: Cotoneaster
9: 392. 1939. campanulatus J.Fryer & B.Hylmö, Cotoneasters
Pyrus buxifolia (Wall. ex Lindl.) M.F.Fay & Christenh.,
comb. nov. Basionym: Cotoneaster buxifolius Wall. Pyrus capsicina (J.Fryer & B.Hylmö) M.F.Fay &

99
capsicinus J.Fryer & B.Hylmö, Cotoneasters Compreh. comb. nov. Basionym: Photinia chingiana Hand.-
Guide : 309. 2009. Mazz., Sinensia 2: 125. 1932.
Pyrus carmesina (McAll.) M.F.Fay & Christenh., comb. Pyrus chingshuiensis (T.Shimizu) M.F.Fay & Christenh.,
nov. Basionym: Sorbus carmesina McAll., Gen. comb. nov. Basionym: Pourthiaea chingshuiensis
Sorbus 231. 2005. T.Shimizu, J. Fac. Text. Sci. & Technol., Shinshu Univ.
Pyrus carniolica (Kárpáti) M.F.Fay & Christenh., comb. no 36 (ser. A, Biol.) 12: 36. 1963.
nov. Basionym: Sorbus carniolica Kárpáti, Biol. Vestn. Pyrus chitralensis (Vassilcz.) M.F.Fay & Christenh.,
16: 21. 1968. comb. nov. Basionym: Malus chitralensis Vassilcz.,
Pyrus cascadensis (G.N.Jones) M.F.Fay & Christenh., Fl. Iranica [Rechinger] 66: 38. 1969.
comb. nov. Basionym: Sorbus cascadensis G.N.Jones, Pyrus chuana (J.Fryer & B.Hylmö) M.F.Fay & Christenh.,
Univ. Wash. Publ. Biol. 7: 108. 1938. comb. nov. Basionym: Cotoneaster chuanus J.Fryer
Pyrus cashmiriensis (G.Klotz) M.F.Fay & Christenh., & B.Hylmö, Cotoneasters Compreh. Guide: 309.
comb. nov. Basionym: Cotoneaster cashmiriensis 2009.
G.Klotz, Wiss. Z. Martin-Luther-Univ. Halle- Pyrus chulingensis (J.Fryer & B.Hylmö) M.F.Fay &
Wittenberg, Math.-Naturwiss. Reihe 12: 778. 1963. Christenh., comb. nov. Basionym: Cotoneaster
Pyrus cavei (G.Klotz) M.F.Fay & Christenh., comb. nov. chulingensis J.Fryer & B.Hylmö, Cotoneasters
Basionym: Cotoneaster cavei G.Klotz, Bull. Bot. Surv. Compreh. Guide : 309. 2009.
India 5: 213. 1963. Replaced synonym: Cotoneaster Pyrus chungtiensis (J.Fryer & B.Hylmö) M.F.Fay &
cordifolius G.Klotz, Bull. Bot. Surv. India 5: 212. 1963, Christenh., comb. nov. Basionym: Cotoneaster
not: Pyrus cordifolia Zamani & Attar. chungtiensis J.Fryer & B.Hylmö, Cotoneasters
Pyrus centralasiatica M.F.Fay & Christenh., nom. Compreh. Guide : 108. 2009.
nov. Replaced synonym: Cotoneaster turcomanicus Pyrus cinerascens (Rehder) M.F.Fay & Christenh.,
Pojark., Not. Syst. Inst. Bot. Acad. Sci USSR 21: 162. comb. nov. Basionym: Cotoneaster franchetii Bois
1961, not: Pyrus turcomanica Maleev. Etymology: var. cinerascens Rehder, J. Arnold Arbor. 4: 114. 1923.
This species occurs in Central Asia, hence the name. Pyrus cinereopubescens (McAll.) M.F.Fay & Christenh.,
Pyrus chadwellii (J.Fryer & B.Hylmö) M.F.Fay & comb. nov. Basionym: Sorbus cinereopubescens
Christenh., comb. nov. Basionym: Cotoneaster McAll., Gen. Sorbus: 233. 2005.
chadwellii J.Fryer & B.Hylmö, Cotoneasters Compreh. Pyrus cinovskisii (J.Fryer & B.Hylmö) M.F.Fay &
Guide : 309. 2009. Christenh., comb. nov. Basionym: Cotoneaster
Pyrus cheddarensis (L.Houston & Ashley Robertson) cinovskisii J.Fryer & B.Hylmö, Cotoneasters Compreh.
M.F.Fay & Christenh., comb. nov. Basionym: Sorbus Guide : 310. 2009.
cheddarensis L.Houston & Ashley Robertson, Pyrus ×clarkiana (C.Weber) M.F.Fay & Christenh.,
Watsonia 27: 288. 2009. comb. nov. Basionym: Cotoneaster ×clarkiana
Pyrus chelseana M.F.Fay & Christenh., nom. nov. C.Weber, Arnoldia 23: 53. 1963. (note: = Pyrus
Replaced synonym: Mespilus tomentosa Aiton, japonica × Pyrus cathayensis).
Hort. Kew. Cat. 2: 174. 1789, not: Pyrus tomentosa Pyrus coadunata (J.Fryer & B.Hylmö) M.F.Fay &
Moench. Etymology: Named for Chelsea Physic Christenh., comb. nov. Basionym: Cotoneaster
Garden in London, where the type specimen was coadunatus J.Fryer & B.Hylmö, Cotoneasters
cultivated. Compreh. Guide : 310. 2009.
Pyrus chengii (C.J.Qi) M.F.Fay & Christenh., comb. nov. Pyrus cochleariformis (Meierott) M.F.Fay & Christenh.,
Basionym: Sorbus chengii C.J.Qi, J. Nanjing Technol. comb. nov. Basionym: Sorbus cochleariformis
Coll. Forest Prod. 1981: 124. 1981. Meierott, Ber. Bayer. Bot. Ges. Sonderband: 176.
Pyrus chengkangensis (T.T.Yu) M.F.Fay & Christenh., 2005.
comb. nov. Basionym: Cotoneaster chengkangensis Pyrus cochleata (Franch.) M.F.Fay & Christenh., comb.
T.T.Yu, Acta Phytotax. Sin. 8: 220. 1963. nov. Basionym: Cotoneaster buxifolius Wall. ex Lindl.
Pyrus chihsiniana (K.C.Kuan) M.F.Fay & Christenh., f. cochleatus Franch., Pl. Delavay. : 224. 1890.
comb. nov. Basionym: Photinia chihsiniana K.C.Kuan, Pyrus colchica (Zinserl.) M.F.Fay & Christenh., comb.
Acta Phytotax. Sin. 8: 227. 1963. nov. Basionym: Sorbus colchica Zinserl., Fl. URSS 9:
Pyrus china M.F.Fay & Christenh., nom. nov. Replaced 391. 1939.
synonym: Sorbus forrestii McAll. & Gillham, Bot. Pyrus collina (M.Lepší, P.Lepší & N.Mey.) M.F.Fay &
Mag. 183: t. 792. 1980, not: Pyrus forrestii (G.Klotz) Christenh., comb. nov. Basionym: Sorbus collina
M.F.Fay & Christenh. Etymology: This is a Chinese M.Lepší, P.Lepší & N.Mey., Preslia 87: 145. 2015.
species, hence the name. Pyrus concavifolia (C.Németh) M.F.Fay & Christenh.,
Pyrus chingiana (Hand.-Mazz.) M.F.Fay & Christenh., comb. nov. Basionym: Sorbus concavifolia

C.Németh, Stud. Bot. Hung. 47: 299. 2016. Bot. Zhurn. (Moscow & Leningrad) 65: 1351. 1980.
Pyrus confusa (G.Klotz) M.F.Fay & Christenh., comb. Pyrus cucullifera (M.Lepší & P.Lepší) M.F.Fay &
nov. Basionym: Cotoneaster confusus G.Klotz, Wiss. Christenh., comb. nov. Basionym: Sorbus cucullifera
Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. M.Lepší & P.Lepší, Preslia 87: 125. 2015.
Reihe 19: 340. 1970. Pyrus cunonioides M.F.Fay & Christenh., nom.
Pyrus conspicua (J.B.Comber ex C.Marquand) M.F.Fay nov. Replaced synonym: Sorbus rufoferruginea
& Christenh., comb. nov. Basionym: Cotoneaster C.K.Schneid., Ill. Handb. Laubholzk. 2: 996. 1912 =
conspicuus J.B.Comber ex C.Marquand, Bull. Misc. Pyrus rufoferruginea (C.K.Schneid.) Cardot, nom.
Inform. Kew 1937: 119. 1937. illeg., not: Pyrus rufoferruginea Koidz. Etymology:
Pyrus convexa (J.Fryer & B.Hylmö) M.F.Fay & This species superficially resembles a species of
Christenh., comb. nov. Basionym: Cotoneaster Cunonia (Cunoniaceae), hence the name.
convexus J.Fryer & B.Hylmö, Cotoneasters Compreh. Pyrus dacica (Borbás) M.F.Fay & Christenh., comb. nov.
Guide : 310. 2009. Basionym: Sorbus dacica Borbás, Oesterr. Bot. Z. 37:
Pyrus cooperi (C.Marquand) M.F.Fay & Christenh., 404. 1887. Replaced synonym: Sorbus semipinnata
comb. nov. Basionym: Cotoneaster cooperi Borbás, Math. Természettud. Értes. 1: 85. 1883, not:
C.Marquand, Hooker’s Icon. Pl. 32: t. 3146. 1930. Pyrus semipinnata Bechst., nor: Pyrus semipinnata
Roth.
Pyrus cordigastensis (N.Mey.) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus cordigastensis N.Mey., Pyrus daliensis (J.Fryer & B.Hylmö) M.F.Fay &
Ber. Bayer. Bot. Ges. Sonderband : 143. 2005. Christenh., comb. nov. Basionym: Cotoneaster
daliensis J.Fryer & B.Hylmö, Cotoneasters Compreh.
Pyrus coriacea (Franch.) M.F.Fay & Christenh., comb.
Guide: 310. 2009.
nov. Basionym: Cotoneaster coriaceus Franch., Pl.
Delavay. 3: 222. 1890. Pyrus dammeri (C.K.Schneid.) M.F.Fay & Christenh.,
comb. nov. Basionym: Cotoneaster dammeri
Pyrus cornifolia (Rehder & E.H.Wilson) M.F.Fay &
C.K.Schneid., Illustr. Handb. Laubholzk. 1: 761.1906.
obscurus Rehder & E.H.Wilson var. cornifolius Rehder Pyrus danubialis (Jav.) M.F.Fay & Christenh., comb.
& E.H.Wilson, Pl. Wilson. (Sargent) 1: 162. 1912. nov. Basionym: Sorbus cretica Dippel f. danubialis
Jáv., Bot. Közlem. 14: 104. 1915.
Pyrus brevipetiolata M.F.Fay & Christenh., nom.
nov. Replaced synonym: Sorbus brevipetiolata Pyrus davidiana (Decne.) M.F.Fay & Christenh., comb.
T.H.Nguyên & Yakovlev, Bot. Zhurn. (Moscow & nov. Basionym: Stranvaesia davidiana Decne., Nouv.
Leningrad) 66: 1189. 1981. Arch. Mus. Par. Ser. 1, 10: 179. 1874.
Pyrus coxii (McAll.) M.F.Fay & Christenh., comb. nov. Pyrus davidsoniae (Rehder & E.H.Wilson) M.F.Fay
Basionym: Sorbus coxii McAll., Gen. Sorbus: 232. & Christenh., comb. nov. Basionym: Photinia
2005. davidsoniae Rehder & E.H.Wilson, Pl. Wilson.
(Sargent) 1: 185. 1912.
Pyrus crassifolia (H.Lév.) M.F.Fay & Christenh., comb.
nov. Basionym: Photinia crassifolia H.Lév., Fl. Kouy- Pyrus decandra (J.Fryer & B.Hylmö) M.F.Fay &
Tchéou: 349. 1915. Christenh., comb. nov. Basionym: Cotoneaster
decandrus J.Fryer & B.Hylmö, Cotoneasters
Pyrus crataegaster M.F.Fay & Christenh., nom. nov.
Replaced synonym: Cotoneaster serotinus Hutch.,
Bot. Mag. 146: t. 8854. 1920, not: Pyrus serotina Pyrus decipientiformis (Kárpáti) M.F.Fay & Christenh.,
Rehder. Etymology: From Crataegus, a genus in comb. nov. Basionym: Sorbus decipientiformis
Rosaceae, and Greek αστέρ, a star. Kárpáti, Agrártud. Egyet. Kert- Szölögazdaságtud.
Karának Évk. 1: 36. 1950.
Pyrus crispii (Exell) M.F.Fay & Christenh., comb. nov.
Basionym: Cotoneaster crispii Exell, Gard. Chron. 83: Pyrus declinata (J.Fryer & B.Hylmö) M.F.Fay &
44. 1928. Christenh., comb. nov. Basionym: Cotoneaster
declinatus J.Fryer & B.Hylmö, Cotoneasters
Pyrus croatica (Kárpáti) M.F.Fay & Christenh., comb.
nov. Basionym: Sorbus austriaca (Beck) Prain subsp.
croatica Kárpáti, Feddes Repert. Spec. Nov. Regni Pyrus deflexa (Hemsl.) M.F.Fay & Christenh., comb.
Veg. 62: 177. 1960. nov. Basionym: Photinia deflexa Hemsl., Ann. Bot.
(Oxford) 9: 153. 1895.
Pyrus croceocarpa (P.D.Sell) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus croceocarpa P.D.Sell, Pyrus degenii (Jáv.) M.F.Fay & Christenh., comb. nov.
Watsonia 17: 392. 1989. Basionym: Sorbus degenii Jáv., Magyar Bot. Lapok
25: 85. 1927.
Pyrus cucphuongensis (T.H.Nguyên & Yakovlev)
M.F.Fay & Christenh., comb. nov. Basionym: Pyrus ×delacourii (André) M.F.Fay & Christenh., comb.
Photinia cucphuongensis T.H.Nguyên & Yakovlev, nov. Basionym: Rhaphiolepis ×delacourii André, Rev.

101
Hort. (Paris) 72: 698. 1900. Naturwiss. Reihe 21: 974. 1972.
Pyrus delavayana (G.Klotz) M.F.Fay & Christenh., Pyrus dolichofoliolatus (X.F.Gao & Meng Li) M.F.Fay
comb. nov. Basionym: Cotoneaster delavayanus & Christenh., comb. nov. Basionym: Sorbus
G.Klotz, Wiss. Z. Martin-Luther-Univ. Halle- dolichofoliolatus X.F.Gao & Meng Li, Phytotaxa 227:
Wittenberg, Math.-Naturwiss. Reihe 12: 774. 1963. 191. 2015.
Pyrus delicata M.F.Fay & Christenh., nom. nov. Pyrus dolomiticola (Mikoláš) M.F.Fay & Christenh.,
Replaced synonym: Cotoneaster gracilis Rehder & comb. nov. Basionym: Sorbus dolomiticola Mikoláš,
E.H.Wilson, Pl. Wilson. (Sargent) 1: 167. 1912, not: Thaiszia 6: 2. 1997.
Pyrus gracilis Siebold & Zucc. Etymology: This is an Pyrus dominii (Kárpáti) M.F.Fay & Christenh., comb.
elegant, delicate species, hence the name. nov. Basionym: Sorbus dominii Kárpáti, Bot. Közlem.
Pyrus delphinensis (Chatenier) M.F.Fay & Christenh., 52: 140. 1966.
comb. nov. Basionym: Cotoneaster delphinensis Pyrus domugledica (Kárpáti) M.F.Fay & Christenh.,
Chatenier, Bull. Soc. Bot. France 69: 712. 1923. comb. nov. Basionym: Sorbus domugledica Kárpáti
Pyrus detergibilis (Merr.) M.F.Fay & Christenh., comb. in Németh, Acta Bot. Hung. 52: 383. 2010.
nov. Basionym: Sorbus detergibilis Merr., Brittonia 4: Pyrus dracofolia (C.Németh) M.F.Fay & Christenh.,
76. 1941. comb. nov. Basionym: Sorbus dracofolia C.Németh,
Pyrus devoniensis (E.F.Warb.) M.F.Fay & Christenh., Kitaibelia 14: 93. 2009, as ‘dracofolius’.
comb. nov. Basionym: Sorbus devoniensis E.F.Warb., Pyrus draconis M.F.Fay & Christenh., nom. nov.
Watsonia 4: 46. 1957. Replaced synonym: Sorbus stenophylla M.Proctor,
Pyrus dielsiana (E.Pritz.) M.F.Fay & Christenh., comb. Watsonia 27: 214. 2009, not: Pyrus stenophylla
nov. Basionym: Cotoneaster dielsianus E.Pritz., Bot. (Hand.-Mazz.) M.F.Fay & Christenh. Etymology:
Jahrb. Syst. 29: 385. 1900. Named for the dragon on the flag of Wales, the
Pyrus difficilis (G.Klotz) M.F.Fay & Christenh., comb. country to which this species is endemic. From Latin
nov. Basionym: Cotoneaster difficilis G.Klotz, Wiss. Z. draco.
Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe Pyrus drogochia (J.Fryer & B.Hylmö) M.F.Fay &
21: 1017. 1972. Christenh., comb. nov. Basionym: Cotoneaster
Pyrus ×dippelii (Zabel) M.F.Fay & Christenh., comb. drogochius J.Fryer & B.Hylmö, Cotoneasters
nov. Basionym: Aronia ×dippelii Zabel, Handb. Compreh. Guide : 311. 2009.
Laubholzben. [Beissner & al.]: 193. 1903. (note: Pyrus dunnii (Rehder) M.F.Fay & Christenh., comb.
Pyrus aria × Pyrus melanocarpa). nov. Basionym: Sorbus dunnii Rehder, Pl. Wilson.
Pyrus disaster M.F.Fay & Christenh., nom. nov. (Sargent) 2: 273. 1915.
Replaced synonym: Cotoneaster orientalis Kerner, Pyrus duthieana (C.K.Schneid.) M.F.Fay & Christenh.,
Oesterr. Bot. Z. 19: 270. 1869, not: Pyrus orientalis comb. nov. Basionym: Cotoneaster distichus Lange
Pall. Etymology: From Greek δισεκατομμύριο, var. duthieanus C.K.Schneid., Ill. Handb. Laubholzk.
abbreviated here as δισ, a billion, and αστέρ, a star. 1: 745. 1906, as ‘duthieana’.
Pyrus dissimilis (G.Klotz) M.F.Fay & Christenh., comb. Pyrus eburnea (McAll.) M.F.Fay & Christenh., comb.
nov. Basionym: Cotoneaster dissimilis G.Klotz, Wiss. nov. Basionym: Sorbus eburnea McAll., Gen. Sorbus
Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. : 233. 2005.
Reihe 21: 984. 1972. Pyrus elata (G.Klotz) M.F.Fay & Christenh., comb.
Pyrus divaricata (Rehder & E.H.Wilson) M.F.Fay & nov. Basionym: Cotoneaster elatus G.Klotz, Wiss.
Christenh., comb. nov. Basionym: Cotoneaster Z. Martin-Luther-Univ. Halle-Wittenberg, Math.-
divaricatus Rehder & E.H.Wilson, Pl. Wilson. Naturwiss. Reihe 12: 781. 1963.
(Sargent) 1: 157. 1912. Pyrus elegans (Rehder & E.H.Wilson) M.F.Fay &
Pyrus diversicolor (Bernátová & Májovský) M.F.Fay & Christenh., comb. nov. Basionym: Cotoneaster
Christenh., comb. nov. Basionym: Sorbus diversicolor dielsianus Pritzel var. elegans Rehder & E.H.Wilson,
Bernátová & Májovský, Biologia (Bratislava) 58: 788. Pl. Wilson. (Sargent) 1: 166. 1912.
2003. Pyrus eleonorae (Aldasoro, Aedo & C.Navarro)
Pyrus doerriana (N.Mey.) M.F.Fay & Christenh., comb. M.F.Fay & Christenh., comb. nov. Basionym: Sorbus
nov. Basionym: Sorbus doerriana N.Mey., Ber. Bayer. eleonorae Aldasoro, Aedo & C.Navarro, Syst. Bot.
Bot. Ges. 86: 228. 2016. Monogr. 69: 51. 2004.
Pyrus dokeriensis (G.Klotz) M.F.Fay & Christenh., Pyrus ellipsoidalis (McAll.) M.F.Fay & Christenh.,
comb. nov. Basionym: Cotoneaster dokeriensis comb. nov. Basionym: Sorbus ellipsoidalis McAll.,
G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.- Gen. Sorbus: 232. 2005.
Pyrus elliptica (Lindl.) M.F.Fay & Christenh., comb.

nov. Basionym: Eriobotrya elliptica Lindl., Trans. H.Lév. Etymology: From Greek εύρηκα, I found.
Linn. Soc. London 13: 102. 1821. Pyrus evae M.F.Fay & Christenh., nom. nov. Replaced
Pyrus emeiensis (J.Fryer & B.Hylmö) M.F.Fay & synonym: Mespilus cotoneaster L. var. nigra Wahlb.,
Christenh., comb. nov. Basionym: Cotoneaster Fl. Gothob.: 53. 1820, not: Pyrus nigra (Willd.) Sarg.
emeiensis J.Fryer & B.Hylmö, Cotoneasters Compreh. Etymology: This species is named for Eve of the book
Guide : 311. 2009. of Genesis, who ate the apple of paradise.
Pyrus eminens (E.F.Warb.) M.F.Fay & Christenh., comb. Pyrus evansii (T.C.G.Rich) M.F.Fay & Christenh., comb.
nov. Basionym: Sorbus eminens E.F.Warb., Watsonia nov. Basionym: Sorbus evansii T.C.G.Rich, New J. Bot.
4: 44. 1957. 4: 2. 2014.
Pyrus eminentiformis (T.C.G.Rich) M.F.Fay & Christenh., Pyrus eximia (Kovanda) M.F.Fay & Christenh., comb.
comb. nov. Basionym: Sorbus eminentiformis nov. Basionym: Sorbus eximia Kovanda, Preslia 56:
T.C.G.Rich, Watsonia 27: 210. 2009. 170. 1984.
Pyrus eminentoides (L.Houston) M.F.Fay & Christenh., Pyrus eyola M.F.Fay & Christenh., nom. nov. Replaced
comb. nov. Basionym: Sorbus eminentoides synonym: Photinia parviflora Card., Notul. Syst.
L.Houston, Watsonia 27: 290. 2009. (Paris) 3: 378. 1918, not: Pyrus parviflora Desf.
Pyrus encavei (J.Fryer & B.Hylmö) M.F.Fay & Christenh., Etymology: Named for Dame Eyola, a plant taking
comb. nov. Basionym: Cotoneaster encavei J.Fryer & the form of a motherly woman who lives in the
B.Hylmö, Cotoneasters Compreh. Guide: 311. 2009. House of Change from The Neverending Story (Ende,
1979) who carried apples on her head.
Pyrus epidendron (Hand.-Mazz.) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus epidendron Hand.- Pyrus eystettensis (N.Mey.) M.F.Fay & Christenh.,
Mazz., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Anz. comb. nov. Basionym: Sorbus eystettensis N.Mey.,
60: 135. 1923. Ber. Bayer. Bot. Ges. Sonderband: 165. 2005.
Pyrus eris M.F.Fay & Christenh., nom. nov. Replaced Pyrus falconeri (G.Klotz) M.F.Fay & Christenh., comb.
synonym: Sorbus salwinensis T.T.Yu & L.T.Lu, Kew nov. Basionym: Cotoneaster falconeri G.Klotz, Wiss.
Bull. 64: 574. 2009, not: Pyrus salwinensis (Hand.- Z. Martin-Luther-Univ. Halle-Wittenberg, Math.-
Mazz.) M.F.Fay & Christenh. Etymology: Named Naturwiss. Reihe 15: 535. 1966.
for Ἔρις, Greek goddess of discord. The myth goes Pyrus fangiana (T.T.Yu) M.F.Fay & Christenh., comb.
that at the marriage of Peleus and Thetis, Eris, was nov. Basionym: Cotoneaster fangianus T.T.Yu, Acta
not invited. She was not amused, so she gave Paris Phytotax. Sin. 8: 219. 1963.
a golden apple inscribed with the word καλλίστῃ Pyrus fansipanensis (McAll.) M.F.Fay & Christenh.,
(kallistēi, “for the fairest”) to take to the wedding. comb. nov. Basionym: Sorbus fansipanensis McAll.,
Pyrus errata (J.Fryer & B.Hylmö) M.F.Fay & Christenh., Gen. Sorbus: 233. 2005.
comb. nov. Basionym: Cotoneaster errata J.Fryer & Pyrus fantabulosa M.F.Fay & Christenh., nom. nov.
B.Hylmö, Cotoneasters Compreh. Guide : 311. 2009. Replaced synonym: Pourthiaea parvifolia E.Pritz.,
Pyrus escalloniifolia (Schltdl.) M.F.Fay & Christenh., Bot. Jahrb. Syst. 29: 389. 1900, not: Pyrus parvifolia
comb. nov. Basionym: Crataegus escalloniifolia Blatt. Etymology: From Polari, fantabulosa, meaning
Schltdl., Linnaea 28: 468. 1857. wonderful.
Pyrus esfandiarii (Khat.) M.F.Fay & Christenh., comb. Pyrus farreri (G.Klotz) M.F.Fay & Christenh., comb.
nov. Basionym: Cotoneaster esfandiarii Khat., Iran. nov. Basionym: Cotoneaster farreri G.Klotz, Wiss. Z.
J. Bot. 5: 2. 1991. Friedrich-Schiller-Univ. Jena Math.-Naturwiss. Reihe
Pyrus esserteauiana (Koehne) M.F.Fay & Christenh., 21: 1006. 1972.
comb. nov. Basionym: Sorbus esserteauiana Koehne, Pyrus fastigiata (J.Fryer & B.Hylmö) M.F.Fay &
Pl. Wilson. (Sargent) 1: 459–460. 1913. Christenh., comb. nov. Basionym: Cotoneaster
Pyrus estiensis (J.Fryer & B.Hylmö) M.F.Fay & fastigiatus J.Fryer & B.Hylmö, Cotoneasters
Christenh., comb. nov. Basionym: Cotoneaster Compreh. Guide : 312. 2009.
estiensis J.Fryer & B.Hylmö, Cotoneasters Compreh. Pyrus favargeri (J.Fryer & B.Hylmö) M.F.Fay &
Guide: 311. 2009. Christenh., comb. nov. Basionym: Cotoneaster
Pyrus eugenii-kelleri (Kárpáti) M.F.Fay & Christenh., favargeri J.Fryer & B.Hylmö, Cotoneasters Compreh.
comb. nov. Basionym: Sorbus eugenii-kelleri Kárpáti, Guide: 312. 2009.
Hung. Acta Biol. 1: 113. 1949. Pyrus filipes (Hand.-Mazz.) M.F.Fay & Christenh.,
Pyrus eureka M.F.Fay & Christenh., nom. nov. Replaced comb. nov. Basionym: Sorbus filipes Hand.-Mazz.,
synonym: Photinia bodinieri H.Lév., Repert. Spec. Symb. Sin. Pt. 7: 472. 1933.
Nov. Regni Veg. 4: 334. 1907, not: Pyrus bodinieri Pyrus fischeri (N.Mey.) M.F.Fay & Christenh., comb.
nov. Basionym: Sorbus fischeri N.Mey., Ber. Bayer.

103
Bot. Ges. Sonderband: 172. 2005. nov. Basionym: Sorbus frutescens McAll., Gen.
Pyrus flinckii (J.Fryer & B.Hylmö) M.F.Fay & Christenh., Sorbus: 233. 2005.
comb. nov. Basionym: Cotoneaster flinckii J.Fryer & Pyrus fruticosa (Steud.) M.F.Fay & Christenh., comb.
B.Hylmö, New Plantsman 5: 140. 1998. nov. Basionym: Sorbus fruticosa Steud., Nomencl.
Pyrus floccosa (Rehder & E.H.Wilson) M.F.Fay & Bot. : 800. 1821.
Christenh., comb. nov. Basionym: Cotoneaster Pyrus fryerae M.F.Fay & Christenh., nom. nov. Replaced
salicifolius Franch. var. floccosus Rehder & synonym: Cotoneaster tardiflorus J.Fryer & B.Hylmö,
E.H.Wilson, Pl. Wilson. (Sargent) 1: 173. 1912. New Plantsman 5: 139. 1998, not: Pyrus tardiflora
Pyrus florida (J.Fryer & B.Hylmö) M.F.Fay & Christenh., Poit. & Turpin. Etymology: This species is named for
comb. nov. Basionym: Cotoneaster floridus J.Fryer & Jeanette Fryer, who first described it.
B.Hylmö, J. Bot. Res. Inst. Texas 2: 55. 2008. Pyrus fulvida (W.W.Sm.) M.F.Fay & Christenh., comb.
Pyrus florifera M.F.Fay & Christenh., nom. nov. nov. Basionym: Cotoneaster hebephyllus Diels var.
Replaced synonym: Cotoneaster racemiflorus Booth fulvidus W.W.Sm., Not. Roy. Bot. Gard., Edinburgh
ex Bosse var. veitchii Rehder & E.H.Wilson, Pl. Wilson. 10: 22. 1917, as ‘fulvida’.
(Sargent) 3(3): 431. 1917, not Pyrus ×veitchii (Trabut) Pyrus futakiana (Kárpáti) M.F.Fay & Christenh., comb.
M.F.Fay & Christenh. Etymology: This species bears nov. Basionym: Sorbus futakiana Kárpáti in Bertová,
abundant flowers, hence its name. Fl. Slovenska IV/3: 441. 1992.
Pyrus fokienensis (Finet & Franch.) M.F.Fay & Pyrus gabrieljanae M.F.Fay & Christenh., nom.
Christenh., comb. nov. Basionym: Photinia glabra nov. Replaced synonym: Sorbus tamamschjanae
(Thunb.) Maxim. var. fokienensis Finet & Franch., Gabrieljan, Biol. Zhurn. Armenii 22(2): 42. 1969, not:
Bull. Soc. Bot. France 46: 207. 1899. Pyrus tamamschianae Fed. (orth. var.). Etymology:
Pyrus formosensis M.F.Fay & Christenh., nom. nov. This species is named for Elenora Gabrieljan, who
Replaced synonym: Cotoneaster taiwanensis J.Fryer first described it.
& B.Hylmö, Cotoneasters Compreh. Guide: 316. Pyrus gallica M.F.Fay & Christenh., nom. nov. Replaced
2009, not: Pyrus taiwanensis Iketani & H.Ohashi. synonym: Sorbus remensis Cornier, Bull. Mens. Soc.
Etymology: This species is found on Taiwan, which Linn. Lyon 78: 36. 2009, not: Pyrus remensis Poit. &
was formerly called Formosa, hence this choice of Turpin. Etymology: This species is found in France,
epithet. hence the epithet chosen.
Pyrus forrestii (G.Klotz) M.F.Fay & Christenh., comb. Pyrus gamblei (G.Klotz) M.F.Fay & Christenh., comb.
nov. Basionym: Cotoneaster forrestii G.Klotz, Bull. nov. Basionym: Cotoneaster gamblei G.Klotz, Wiss.
Bot. Surv. India 5: 209. 1963. Z. Martin-Luther-Univ. Halle-Wittenberg, Math.-
Pyrus foveolata (Rehder & E.H.Wilson) M.F.Fay & Naturwiss. Reihe 15: 530. 1966.
Christenh., comb. nov. Basionym: Cotoneaster Pyrus ganghobaensis (J.Fryer & B.Hylmö) M.F.Fay
foveolatus Rehder & E.H.Wilson, Pl. Wilson. & Christenh., comb. nov. Basionym: Cotoneaster
(Sargent) 1: 162. 1912. ganghobaensis J.Fryer & B.Hylmö, New Plantsman
Pyrus franchetii (Bois) M.F.Fay & Christenh., comb. 5: 136. 1998.
nov. Basionym: Cotoneaster franchetii Bois, Rev. Pyrus gansuana M.F.Fay & Christenh., nom. nov.
Hortic. 2: 380. 1902. Replaced synonym: Cotoneaster kansuensis
Pyrus franconica (Bornm.) M.F.Fay & Christenh., comb. G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena,
nov. Basionym: Sorbus franconica Bornm., Beih. Bot. Math.-Naturwiss. Reihe 21: 1001. 1972, not: Pyrus
Centralbl. 36: 186. 1918. kansuensis Batalin. Etymology: This species is only
known from the Chinese province Gansu.
Pyrus fraxinus M.F.Fay & Christenh., nom. nov.
Replaced synonym: Sorbus cashmiriana Hedl., Kongl. Pyrus ganymedes M.F.Fay & Christenh., nom. nov.
Svenska Vetensk.-Akad. Handl. 35(1): 35. 1901, not: Replaced synonym: Stranvaesia tomentosa T.T.Yu
Pyrus cashmiriana Cardot. Etymology: This species & T.C.Ku, Acta Phytotax. Sin. 13: 102. 1975, not:
has leaves that superficially resemble those of an Pyrus tomentosa Moench. Etymology: Named for
ash tree, Fraxinus (Oleaceae). Γανυμήδης, a prince of Troy who was kidnapped
by Zeus (in the form of an eagle) because of his
Pyrus frigida (Wall. ex Lindl.) M.F.Fay & Christenh.,
outstanding beauty.
comb. nov. Basionym: Cotoneaster frigidus Wall. ex
Lind., Edward’s Bot. Reg. 15: t. 1229. 1829. Pyrus garhwalensis (G.Klotz) M.F.Fay & Christenh.,
comb. nov. Basionym: Cotoneaster garhwalensis
Pyrus froebelii (Sax ex Vilm.) M.F.Fay & Christenh.,
G.Klotz, Wiss. Z. Martin-Luther-Univ. Halle-
comb. nov. Basionym: Cotoneaster froebelii Sax ex
Wittenberg, Math.-Naturwiss. Reihe 15: 543. 1966.
Vilm., J. Arnold Arbor.. 35: 340. 1954.
Pyrus garraudiana M.F.Fay & Christenh., nom. nov.
Pyrus frutescens (McAll.) M.F.Fay & Christenh., comb.
Replaced synonym: Cotoneaster intermedius

H.J.Coste, Bull. Soc. Bot. France 40: 122. 1894, not: Franch., Pl. Delavay. 3: 222. 1890.
Pyrus intermedia Bechst., nor: Pyrus intermedia Pyrus globosa (T.T.Yu & H.T.Tsai) M.F.Fay & Christenh.,
Ten. Etymology: Named for L. Garraud, for his comb. nov. Basionym: Sorbus globosa T.T.Yu &
contributions to Cotoneaster taxonomy. H.T.Tsai, Bull. Fan Mem. Inst. Biol. Bot. 7: 121. 1936.
Pyrus gauckleri (N.Mey.) M.F.Fay & Christenh., comb. Pyrus glomerata (Rehder & E.H.Wilson) M.F.Fay
nov. Basionym: Sorbus gauckleri N.Mey., Ber. Bayer. & Christenh., comb. nov. Basionym: Photinia
Bot. Ges. Sonderband: 96. 2005. glomerata Rehder & E.H.Wilson, Pl. Wilson.
Pyrus gaultherioides M.F.Fay & Christenh., nom. nov. (Sargent) 1: 190. 1912.
Replaced synonym: Cotoneaster fruticosus J.Fryer & Pyrus godiva M.F.Fay & Christenh., nom. nov.
B.Hylmö, Cotoneasters Compreh. Guide: 312. 2009, Replaced synonym: Rhaphiolepis integerrima Hook
not: Pyrus fruticosa (Steud.) M.F.Fay & Christenh. & Arn., Bot. Beechey Voy.: 263. 1838, not: Pyrus
Etymology: This species resembles a wintergreen, integerrima Wall. ex D.Don Etymology: Named for
Gaultheria (Ericaceae). Lady Godgifu (Old English, meaning ‘gift of god’,
Pyrus gayana (Decne.) M.F.Fay & Christenh., comb. Latinised as Godiva), Countess of Mercia in the 11th
nov. Basionym: Osteomeles gayana Decne., Nouv. Century, who rode a horse naked to be pardoned for
Arch. Mus. Par. Ser. 1, 10: 183. 1874. her husband’s oppressive taxes.
Pyrus gayeriana (Kárpáti) M.F.Fay & Christenh., comb. Pyrus goloskokovii (Pojark.) M.F.Fay & Christenh.,
nov. Basionym: Sorbus gayeriana Kárpáti, Agrártud. comb. nov. Basionym: Cotoneaster goloskokovii
Egyet. Kert- Szölögazdaságtud. Karának Évk. 1(14): Pojark., Opred. Rast. Sred. Azii 5: 245. 1976.
35. 1950. Pyrus gomorrana M.F.Fay & Christenh., nom. nov.
Pyrus gemella (Kovanda) M.F.Fay & Christenh., comb. Replaced synonym: Rhaphiolepis salicifolia Lind.,
nov. Basionym: Sorbus gemella Kovanda, Verh. Zool.- Collect. Bot. 1: t. 3. 1821, not: Pyrus salicifolia
Bot. Ges. Österreich 133: 329. 1996. Pall. Etymology: Named for the Biblical sinful city
Pyrus genesis M.F.Fay & Christenh., nom. nov. Replaced mentioned in Genesis.
synonym: Malus muliensis T.C.Ku, Acta Phytotax. Sin. Pyrus gonggashanensis (J.Fryer & B.Hylmö) M.F.Fay
29: 83. 1991, not: Pyrus muliensis (G.Klotz) M.F.Fay & Christenh., comb. nov. Basionym: Cotoneaster
& Christenh. Etymology: This species is named for gonggashanensis J.Fryer & B.Hylmö, Cotoneasters
the Book of Genesis, in which an appletree played Compreh. Guide: 312. 2009.
an important role. Pyrus gonggashanica (McAll.) M.F.Fay & Christenh.,
Pyrus genitana (Hurus.) M.F.Fay & Christenh., comb. comb. nov. Basionym: Sorbus gonggashanica McAll.,
nov. Basionym: Cotoneaster genitanus Hurus., Acta Gen. Sorbus: 234. 2005.
Phytotax. Geobot. 13: 233. 1943. Pyrus gotlandica M.F.Fay & Christenh., nom. nov.
Pyrus gerecseensis (Boros & Kárpáti) M.F.Fay Replaced synonym: Cotoneaster canescens Vestergr.
& Christenh., comb. nov. Basionym: Sorbus ex B.Hylmö, Svensk Bot. Tidskr. 87: 319. 1993, not:
gerecseensis Boros & Kárpáti, Hung. Acta Biol. 1: Pyrus canescens Spach. Etymology: This is a species
107. 1949. found on Gotland, Sweden, hence the name.
Pyrus gilgitana (McAll.) M.F.Fay & Christenh., comb. Pyrus gracia (I.F.Schneid.) M.F.Fay & Christenh., comb.
nov. Basionym: Sorbus gilgitana McAll., Gen. Sorbus: nov. Basionym: Cotoneaster gracia I.F.Schneid.,
235. 2005. Dendroflora 1: 35. 1964. (note: originally described
Pyrus giraldii (Flinck & B.Hylmö ex G.Klotz) M.F.Fay as a cultivar of hybrid origin, this is more likely to be
& Christenh., comb. nov. Basionym: Cotoneaster a species and should be treated as such).
giraldii Flinck & B.Hylmö ex G.Klotz, Wiss. Z. Pyrus granatensis (Boiss.) M.F.Fay & Christenh., comb.
Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe nov. Basionym: Cotoneaster granatensis Boiss.,
21: 1018. 1972. Elench. Pl. Nov. Hisp. Austr.: 71. 1838.
Pyrus glabriuscula (McAll.) M.F.Fay & Christenh., Pyrus grandifructa M.F.Fay & Christenh., nom. nov.
comb. nov. Basionym: Sorbus glabriuscula McAll., Replaced synonym: Cotoneaster megalocarpus
Gen. Sorbus: 232. 2005. M.Popov, Bull. Soc. Imp. Naturalistes Moscou 44:
Pyrus glacialis (Hook.f. ex Wenz.) M.F.Fay & Christenh., 128. 1935, not: Pyrus megalocarpa (Rehder) Bean.
comb. nov. Basionym: Cotoneaster microphyllus Etymology: This species has large fruits, hence the
Wall. ex Lindl. f. glacialis Hook.f. ex Wenz, Linnaea name.
38: 195. 1874. Pyrus greenii (T.C.G.Rich) M.F.Fay & Christenh., comb.
Pyrus glaucophylla (Franch.) M.F.Fay & Christenh., nov. Basionym: Sorbus greenii T.C.G.Rich, New J. Bot.
comb. nov. Basionym: Cotoneaster glaucophyllus 4: 3. 2014.
Pyrus griffithiana M.F.Fay & Christenh., nom. nov.

105
Replaced synonym: Photinia griffithii Decne., Nouv. Christenh., comb. nov. Basionym: Cotoneaster
Arch. Mus. Par. Ser. 1, 10: 142. 1874, not: Pyrus hedegaardii J.Fryer & B.Hylmö, Cotoneasters
griffithii Hook.f. Etymology: Named for William Compreh. Guide : 312. 2009.
Griffith, following the lead of Joseph Descaine, who Pyrus helenae (Koehne) M.F.Fay & Christenh., comb.
originally named this species in his honour. nov. Basionym: Sorbus helenae Koehne, Pl. Wilson.
Pyrus guanii (Rushforth) M.F.Fay & Christenh., comb. (Sargent) 1: 462. 1913.
nov. Basionym: Sorbus guanii Rushforth, Int. Pyrus hellas M.F.Fay & Christenh., nom. nov. Replaced
Dendrol. Soc. Year Book 2009: 88. 2010. synonym: Crataegus graeca Spach, Hist. Nat. Vég.
Pyrus guanxianensis (T.C.Ku) M.F.Fay & Christenh., 2: 102. 1834, not: Pyrus graeca hort. ex Steud.
comb. nov. Basionym: Sorbus guanxianensis T.C.Ku, Etymology: After Ελλας the modern Greek name for
Bull. Bot. Res., Harbin 10(3): 22. 1990. Greece.
Pyrus gulabi M.F.Fay & Christenh., nom. nov. Replaced Pyrus henryi (Nakai) M.F.Fay & Christenh., comb. nov.
synonym: Sorbus rosea McAll., Gen. Sorbus: 236. Basionym: Eriobotrya henryi Nakai, J. Arnold Arbor.
2005, not: Pyrus rosea (Edgew.) M.F.Fay & Christenh. 5: 70. 1924.
Etymology: The name means ‘pink’ in Urdu, the Pyrus heracleotica M.F.Fay & Christenh., nom. nov.
language spoken where this species occurs. It has Replaced synonym: Cotoneaster creticus J.Fryer
pink flowers and pink fruit. & B.Hylmö, Cotoneasters Compreh. Guide: 310.
Pyrus guerreris (Phipps) M.F.Fay & Christenh., comb. 2009, not: Pyrus cretica Willd. Etymology: Named
nov. Basionym: Photinia guerreris Phipps, Canad. J. for the hero Ἡρακλῆς, god of strength, health and
Bot. 70: 2158. 1992. agriculture in Greek mythology.
Pyrus haesitans (Meierott) M.F.Fay & Christenh., Pyrus herae M.F.Fay & Christenh., nom. nov. Replaced
comb. nov. Basionym: Sorbus haesitans Meierott, synonym: Eriobotrya latifolia Hook.f., Fl. Brit. India
Ber. Bayer. Bot. Ges. Sonderband : 180. 2005. 2: 370. 1878, not: Pyrus ×latifolia Lindl. ex Steud.
Pyrus hajastana (Gabrieljan) M.F.Fay & Christenh., Etymology: Named for Ἥρᾱ, Greek goddess of
comb. nov. Basionym: Sorbus hajastana Gabrieljan, marriage and femininity. The myth goes that at the
Dokl. Akad. Nauk Armyanskoi S.S.R. 22: 87. 1956. marriage of Peleus and Thetis, Eris, the goddess of
discord, was not invited. She was not amused, so she
Pyrus hajastanica (Nersesian) M.F.Fay & Christenh.,
gave Paris a golden apple inscribed with the word
comb. nov. Basionym: Cotoneaster hajastanicus
καλλίστῃ (kallistēi, “for the fairest”). Aphrodite,
Nersesian, Novosti Sist. Vyssh. Rast. 44: 106. 2013.
Hera, and Athena all claimed to be the fairest, and
Pyrus haljamovae (Bernátová & Májovský) M.F.Fay the rightful owner of the apple.
& Christenh., comb. nov. Basionym: Sorbus
Pyrus herbipolitana (Meierott) M.F.Fay & Christenh.,
haljamovae Bernátová & Májovský, Biologia
comb. nov. Basionym: Sorbus herbipolitana
(Bratislava) 58: 784. 2003.
Meierott, Ber. Bayer. Bot. Ges. Sonderband: 183.
Pyrus hammelii M.F.Fay & Christenh., nom. nov. 2005.
Replaced synonym: Sorbus meyeri S.Hammel &
Pyrus herefordensis (D.Green) M.F.Fay & Christenh.,
Haynold, Kochia 8: 2. 2014, not: Pyrus meyeri
comb. nov. Basionym: Sorbus herefordensis D.Green,
(Pojark.) M.F.Fay & Christenh. Etymology: This
New J. Bot. 4: 4. 2014.
species was named for Steffen Hammel, who first
described it. Pyrus hersiana (J.Fryer & B.Hylmö) M.F.Fay &
Pyrus harrysmithii (Flinck & B.Hylmö) M.F.Fay &
hersianus J.Fryer & B.Hylmö, Cotoneasters Compreh.
Guide : 312. 2009.
harrysmithii Flinck & B.Hylmö, Bot. Not. (Lund) 115.
29. 1962. Pyrus heseltinei (Rushforth) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus heseltinei Rushforth,
Pyrus harziana (N.Mey.) M.F.Fay & Christenh., comb.
Int. Dendrol. Soc. Year Book 2009: 83. 2010.
nov. Basionym: Sorbus harziana N.Mey., Ber. Bayer.
Bot. Ges. Sonderband : 100. 2005. Pyrus hesperidea M.F.Fay & Christenh., nom. nov.
Replaced synonym: Hesperomeles cuneata Lindl.,
Pyrus hazslinszkyana (Soó) M.F.Fay & Christenh., comb.
Edwards’s Bot. Reg. 23: sub t. 1956. 1837, not: Pyrus
nov. Basionym: Sorbus aria L. var. hazslinszkyana
cuneata Koidz. Etymology: Named for the Garden of
Soó, Tisia 2: 218. 1937, not: Pyrus ×hazslinszkyana
the Hesperides, Ἑσπερίδες, where the golden apples
Terpó, nom. inval., nom. nud.
of immortality grow.
Pyrus hebephylla (Diels) M.F.Fay & Christenh., comb.
Pyrus hibernica (E.F.Warb.) M.F.Fay & Christenh.,
nov. Basionym: Cotoneaster hebephyllus Diels, Not.
comb. nov. Basionym: Sorbus hibernica E.F.Warb.,
Roy. Bot. Gard. Edinburgh 5: 273. 1912.
Watsonia 4: 44. 1957.
Pyrus hedegaardii (J.Fryer & B.Hylmö) M.F.Fay &
Pyrus hicksii (J.Fryer & B.Hylmö) M.F.Fay & Christenh.,

comb. nov. Basionym: Cotoneaster hicksii J.Fryer & for the Garden of Eden.
B.Hylmö, Cotoneasters Compreh. Guide : 313. 2009. Pyrus ×houstoniae (T.C.G.Rich) M.F.Fay & Christenh.,
Pyrus hillieri (J.Fryer & B.Hylmö) M.F.Fay & Christenh., comb. nov. Basionym: Sorbus ×houstoniae
comb. nov. Basionym: Cotoneaster hillieri J.Fryer & T.C.G.Rich, Watsonia 27: 370. 2009. (note: Pyrus
B.Hylmö, Cotoneasters Compreh. Guide : 313. 2009. bristoliensis × Pyrus aria)
Pyrus himalaica (Gabrieljan) M.F.Fay & Christenh., Pyrus hsingshangensis (J.Fryer & B.Hylmö) M.F.Fay
comb. nov. Basionym: Sorbus himalaica Gabrieljan, & Christenh., comb. nov. Basionym: Cotoneaster
Bot. Zhurn. (Moscow & Leningrad) 56: 658. 1971. hsingshangensis J.Fryer & B.Hylmö, Watsonia 21:
Pyrus hippomenes M.F.Fay & Christenh., nom. nov. 335. 1997.
Replaced synonym: Mespilus prunifolia Marshall, Pyrus huahongdongensis (J.Fryer & B.Hylmö) M.F.Fay
Arbust. Amer.: 90. 1785, not: Pyrus prunifolia Willd. & Christenh., comb. nov. Basionym: Cotoneaster
Etymology: Named for Ἱππομένης, a Calydonian huahongdongensis J.Fryer & B.Hylmö, Cotoneasters
hunstman in Greek mythology, who used golden Compreh. Guide : 313. 2009.
apples to distract Atalanta so he could win the race Pyrus hualiensis (J.Fryer & B.Hylmö) M.F.Fay &
and marry her. Christenh., comb. nov. Basionym: Cotoneaster
Pyrus hirsuta (Hand.-Mazz.) M.F.Fay & Christenh., hualiensis J.Fryer & B.Hylmö, New Plantsman 8: 236.
comb. nov. Basionym: Photinia hirsuta Hand.-Mazz., 2001.
Symb. Sin. Pt. 7: 481. 1933. Pyrus hubeiana M.F.Fay & Christenh., nom. nov.
Pyrus hissarica (Pojark.) M.F.Fay & Christenh., comb. Replaced synonym: Cotoneaster hupehensis Rehder
nov. Basionym: Cotoneaster hissaricus Pojark., Not. & E.H.Wilson, Pl. Wilson. (Sargent) 1: 169. 1912, not:
Syst. Inst. Bot. Acad. Sci. USSR 16: 124. 1954. Pyrus hupehensis Pamp. Etymology: Named for the
Pyrus hjelmqvistii (Flinck & B.Hylmö) M.F.Fay & Chinese province of Hubei, formerly Hupeh, where
Christenh., comb. nov. Basionym: Cotoneaster this species occurs.
hjelmqvistii Flinck & B.Hylmö, Watsonia 18: 312. Pyrus hudsonii (Rushforthš) M.F.Fay & Christenh.,
1991. comb. nov. Basionym: Sorbus hudsonii Rushforth,
Pyrus hodjingensis (G.Klotz) M.F.Fay & Christenh., Int. Dendrol. Soc. Year Book 2009: 89. 2010.
comb. nov. Basionym: Cotoneaster hodjingensis Pyrus hughmcallisteri (Mikoláš) M.F.Fay & Christenh.,
G.Klotz, Wiss. Z. Martin-Luther-Univ. Jalle- comb. nov. Basionym: Sorbus hughmcallisteri
Wittenberg, Math.-Naturwiss. Reihe 12: 774. 1963. Mikoláš, Thaiszia 18: 66. 2009, as ‘hugh-mcallisteri’.
Pyrus hohenesteri (N.Mey.) M.F.Fay & Christenh., Pyrus humilis (Dunn) M.F.Fay & Christenh., comb. nov.
comb. nov. Basionym: Sorbus hohenesteri N.Mey., Basionym: Cotoneaster humilis Dunn, Bull. Misc.
Ber. Bayer. Bot. Ges. Sonderband : 103. 2005. Inform. Kew 1924(10): 384. 1924.
Pyrus holubyana (Kárpáti) M.F.Fay & Christenh., comb. Pyrus hummelii (J.Fryer & B.Hylmö) M.F.Fay &
nov. Basionym: Sorbus holubyana Kárpáti, Bot. Christenh., comb. nov. Basionym: Cotoneaster
Közlem. 52: 137. 1966. hummelii J.Fryer & B.Hylmö, Watsonia 21: 336.
Pyrus honanensis (Rehder) M.F.Fay & Christenh., 1997.
comb. nov. Basionym: Malus honanensis Rehder, J. Pyrus hungarica (Bornm.) M.F.Fay & Christenh., comb.
Arnold Arbor. 2: 51. 1920. nov. Basionym: Sorbus aria L. f. hungarica Bornm.,
Pyrus hookeriana (Decne.) M.F.Fay & Christenh., comb. Mitth. Thüring. Bot. Vereins 30: 54. 1913.
nov. Basionym: Eriobotrya hookeriana Decne., Nouv. Pyrus hurusawana (G.Klotz) M.F.Fay & Christenh.,
Arch. Mus. Par. Ser. 1, 10: 146. 1874. comb. nov. Basionym: Cotoneaster hurusawanus
Pyrus hoppeana (N.Mey.) M.F.Fay & Christenh., comb. G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-
nov. Basionym: Sorbus hoppeana N. Mey., Ber. Naturwiss. Reihe 21: 984. 1972.
Bayer. Bot. Ges. Sonderband: 150. 2005. Pyrus hylanderi (J.Fryer & B.Hylmö) M.F.Fay &
Pyrus horizontalis (Decne.) M.F.Fay & Christenh., Christenh., comb. nov. Basionym: Cotoneaster
comb. nov. Basionym: Cotoneaster horizontalis hylanderi J.Fryer & B.Hylmö, Cotoneasters Compreh.
Decne., Fl. Ser. Jard. Europe 22: 168. 1877. Guide : 313. 2009.
Pyrus hornadensis (Mikoláš) M.F.Fay & Christenh., Pyrus hylmoei (Flinck & J.Fryer) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus hornadensis Mikoláš, comb. nov. Basionym: Cotoneaster hylmoei Flinck &
Thaiszia 25: 22. 2015. J.Fryer, Plantsman 15: 26. 1993.
Pyrus horti-edeni M.F.Fay & Christenh., nom. nov. Pyrus hypocarpa (J.Fryer & B.Hylmö) M.F.Fay &
Replaced synonym: Sorbus serotina Koehne, Mitt. Christenh., comb. nov. Basionym: Cotoneaster
Deutsch. Dendrol. Ges. 15: 59. 1906, not: Pyrus hypocarpus J.Fryer & B.Hylmö, Cotoneasters
serotina Rehder. Etymology: This species is named

107
Compreh. Guide 313. 2009. formerly called Persia, hence the choice of name.
Pyrus ichangensis (G.Klotz) M.F.Fay & Christenh., Pyrus istriaca (Kárpáti) M.F.Fay & Christenh., comb.
comb. nov. Basionym: Cotoneaster ichangensis nov. Basionym: Sorbus istriaca Kárpáti, Biol. Vestn.
G.Klotz, Mitt. Deutsch. Dendrol. Ges. 82: 72. 1996. 16: 19. 1968.
Pyrus ignava (E.L.Wolf) M.F.Fay & Christenh., comb. Pyrus javorkana (Somlyay, Sennikov & Vojtkó)
nov. Basionym: Cotoneaster ignavus E.L.Wolf, Bull. M.F.Fay & Christenh., comb. nov. Basionym: Sorbus
Imp. For. Inst. St. Petersburg 15: 240. 1907. javorkana Somlyay, Sennikov & Vojtkó, Ann. Bot.
Pyrus ignescens (J.Fryer & B.Hylmö) M.F.Fay & Fenn. 54: 230. 2017.
Christenh., comb. nov. Basionym: Cotoneaster Pyrus jingii M.F.Fay & Christenh., nom. nov. Replaced
ignescens J.Fryer & B.Hylmö, New Plantsman 8: 236. synonym: Sorbus prunifolia W.B.Liao & H.J.Jing,
2001. Phytotaxa 258: 172. 2016, not: Pyrus prunifolia
Pyrus ignota (G.Klotz) M.F.Fay & Christenh., comb. Willd. Etymology: Named for Hui Juan Jing, who first
nov. Basionym: Cotoneaster ignotus G.Klotz, Wiss. Z. described this species.
Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe Pyrus jinxianensis (J.Q.Deng & J.Y.Hong) M.F.Fay &
19: 339. 1970. Christenh., comb. nov. Basionym: Malus jinxianensis
Pyrus immortalis M.F.Fay & Christenh., nom. nov. J.Q.Deng & J.Y.Hong, Acta Phytotax. Sin. 25: 326.
Replaced synonym: Cotoneaster marginatus Lindl. 1987.
ex Loudon, Encycl. Trees & Shrubs: 411. 1842, not: Pyrus jiulongjiangensis (P.C.Huang & K.M.Li) M.F.Fay
Pyrus marginata Kit. Etymology: This is a hardy & Christenh., comb. nov. Basionym: Rhaphiolepis
species that can live for a long time. However, its jiulongjiangensis P.C.Huang & K.M.Li, J. Nanjing
fruits will not convey immortality. Forest. Univ. 13: 85. 1989.
Pyrus impressivena (Hayata) M.F.Fay & Christenh., Pyrus joannis (Kárpáti) M.F.Fay & Christenh., comb.
comb. nov. Basionym: Photinia impressivena Hayata, nov. Basionym: Sorbus joannis Kárpáti, Bot. Közlem.
Icon. Pl. Formosan. 5: 67. 1915. 52: 140. 1966;
Pyrus incana (W.W.Sm.) M.F.Fay & Christenh., comb. Pyrus juno M.F.Fay & Christenh., nom. nov. Replaced
nov. Basionym: Cotoneaster hebephyllus Diels var. synonym: Sorbus kongboensis McAll., Gen. Sorbus:
incanus W.W.Sm., Not. Roy. Bot. Gard. Edinburgh 235. 2005, not: Pyrus kongboensis (G.Klotz) M.F.Fay
10(46): 22. 1917. & Christenh. Etymology: Named for Juno, an Ancient
Pyrus indurata (J.Fryer & B.Hylmö) M.F.Fay & Roman goddesss of childbirth and marriage.
Christenh., comb. nov. Basionym: Cotoneaster Pyrus jupiter M.F.Fay & Christenh., nom. nov. Replaced
induratus J.Fryer & B.Hylmö, Watsonia 21: 339. synonym: Photinia oblongifolia Standl., Publ. Field
1997. Columb. Mus., Bot. Ser. 4: 210. 1929, not: Pyrus
Pyrus inexpectata (G.Klotz) M.F.Fay & Christenh., oblongifolia Spach. Etymology: This species is
comb. nov. Basionym: Cotoneaster inexpectatus named for the planet Jupiter and the Roman God of
G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.- the same name.
Naturwiss. Reihe 27: 23. 1978. Pyrus jurana (Gand.) M.F.Fay & Christenh., comb. nov.
Pyrus insculpta (Diels) M.F.Fay & Christenh., comb. Basionym: Cotoneaster juranus Gand., Fl. Lyon.: 87.
nov. Basionym: Cotoneaster insculptus Diels, Not. 1875.
Roy. Bot. Gard. Edinburgh 5: 273. 1912. Pyrus kaganensis (G.Klotz) M.F.Fay & Christenh.,
Pyrus insignoides hort. ex M.F.Fay & Christenh., comb. nov. Basionym: Cotoneaster kaganensis
nom. nov. Replaced synonym: Cotoneaster insignis G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-
Pojark., Fl. USSR 9: 330. 1939, not: Pyrus insignis Naturwiss. Reihe 15: 541. 1966.
Hook.f. Pyrus kamschatcensis (Komarov) M.F.Fay & Christenh.,
Pyrus insolita (G.Klotz) M.F.Fay & Christenh., comb. comb. nov. Basionym: Sorbus kamschatcensis
nov. Basionym: Cotoneaster insolitus G.Klotz, Wiss. Komarov, Bot. Mater. Gerb. Glavn. Bot. Sada
Z. Martin-Luther-Univ. Halle-Wittenberg, Math.- R.S.F.S.R. 2: 135. 1921.
Naturwiss. Reihe 15: 536. 1966. Pyrus kangdingensis (J.Fryer & B.Hylmö) M.F.Fay
Pyrus integrifolia (Lindl.) M.F.Fay & Christenh., comb. & Christenh., comb. nov. Basionym: Cotoneaster
nov. Basionym: Photinia integrifolia Lindl., Trans. kangdingensis J.Fryer & B.Hylmö, Cotoneasters
Linn. Soc. London 13: 103. 1821. Compreh. Guide : 314. 2009.
Pyrus iranica M.F.Fay & Christenh., nom. nov. Replaced Pyrus karatavica (Pojark.) M.F.Fay & Christenh., comb.
synonym: Cotoneaster persicus Pojark., Not. Syst. nov. Basionym: Cotoneaster karatavicus Pojark.,
Inst. Bot. Acad. Sci. USSR 16: 118. 1954, not: Pyrus Nov. Syst. Inst. Bot. Acad. Sci. USSR 21: 184. 1961.
persica Pers. Etymology: This species occurs in Iran, Pyrus karchungii (Rushforth) M.F.Fay & Christenh.,

comb. nov. Basionym: Sorbus karchungii Rushforth, 21: 992. 1972, not: Pyrus ×vilmoriniana (C.Weber)
Int. Dendrol. Soc. Year Book 2009: 81. 2010. M.F.Fay & Christenh. Etymology: This species is
Pyrus karpatii (Boros) M.F.Fay & Christenh., comb. named for Gerhard Klotz, who first described it.
nov. Basionym: Sorbus karpatii Boros, Agrártud. Pyrus kmetiana (Kárpáti) M.F.Fay & Christenh., comb.
Egyet. Kert- Szölögazdaságtud. Karának Közlem. 13: nov. Basionym: Sorbus kmetiana Kárpáti in Bertová,
153. 1949. Fl. Slovenska IV/3: 440. 1992.
Pyrus kaschkarovii (Pojark.) M.F.Fay & Christenh., Pyrus komarovii (H.Lév.) M.F.Fay & Christenh., comb.
comb. nov. Basionym: Cotoneaster kaschkarovii nov. Basionym: Viburnum komarovii H.Lév. & Vaniot,
Pojark., Not. Syst. Inst. Bot. Acad. Sci USSR 21: 194. Repert. Spec. Nov. Regni Veg. 9: 78. 1910. Etymology:
1961. This species occurs in North Korea.
Pyrus ×kawashiroi (Murata) M.F.Fay & Christenh., Pyrus kongboensis (G.Klotz) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus kawashiroi Murata, comb. nov. Basionym: Cotoneaster kongboensis
Acta Phytotax. Geobot. 35: 36. 1984. G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-
Pyrus keenanii (Rushforth) M.F.Fay & Christenh., Naturwiss. Reihe 21: 997. 1972.
comb. nov. Basionym: Sorbus keenanii Rushforth, Pyrus konishii (Hayata) M.F.Fay & Christenh., comb.
Int. Dendrol. Soc. Year Book 2011: 92. 2012. nov. Basionym: Cotoneaster konishii Hayata, Icon. Pl.
Pyrus kerstanii (G.Klotz) M.F.Fay & Christenh., comb. Formosa 3: 100. 1913.
nov. Basionym: Cotoneaster kerstanii G.Klotz, Wiss. Pyrus kopetdagensis M.F.Fay & Christenh., nom. nov.
Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Replaced synonym: Cotoneaster discolor Pojark.,
Reihe 21: 1010. 1972. Not. Syste. Inst. Bot. Acad. USSR 16: 116. 1954, not:
Pyrus keszthelyensis (Somlyay & Sennikov) M.F.Fay Pyrus discolor Maxim. Etymology: This species is
& Christenh., comb. nov. Basionym: Sorbus only found on the Kopet Dag mountain range on the
keszthelyensis Somlyay & Sennikov, Ann. Bot. Fenn. border of Iran and Turkmenistan.
53: 368. 2016. Pyrus koreana M.F.Fay & Christenh., comb. nov.
Pyrus ×kewensis (K.J.W.Hensen) M.F.Fay & Christenh., Basionym: Crataegus komarovii Sarg., Pl. Wilson.
comb. nov. Basionym: Sorbus ×kewensis 1(2): 183. 1912, not: Pyrus komarovii (H.Lév.) M.F.Fay
K.J.W.Hensen, Dendroflora 3: 67. 1966. & Christenh. Etymology: This species occurs in North
Korea, hence the name.
Pyrus khumbuensis (McAll.) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus khumbuensis McAll., Pyrus ×kovalevii (Mezhenskyj) M.F.Fay & Christenh.,
Gen. Sorbus : 234. 2005. comb. nov. Basionym: ×Sorbaronia kovalevii
Mezhenskyj, Ukrayins’k. Bot. Zhurn. 70(4): 514. 2013.
Pyrus kingdonii (J.Fryer & B.Hylmö) M.F.Fay &
(note: Pyrus melanocarpa × Pyrus sambucifolia)
kingdonii J.Fryer & B.Hylmö, New Plantsman 8: 237. Pyrus krasnovii (Pojark.) M.F.Fay & Christenh., comb.
2001. nov. Basionym: Cotoneaster krasnovii Pojark., Not.
Syst. Inst. Bot. Acad. Sci. USSR 21: 187. 1961.
Pyrus kirghisorum (Al.Fed. & Fed.) M.F.Fay &
Christenh., comb. nov. Basionym: Malus kirghisorum Pyrus krymica M.F.Fay & Christenh., nom. nov.
Al.Fed. & Fed., Plodovye lesa juzhnoi Kirgisii i ich Replaced synonym: Cotoneaster tauricus Pojark.,
ispolsovanie: 220. 1949. Not. Syst. Inst. Bot. Acad. Sci. USSR 8: 138. 1940,
not: Pyrus taurica (Zinserl.) M.F.Fay & Christenh.
Pyrus kitaibeliana (Baksay & Kárpáti) M.F.Fay &
Etymology: This species if only found in Crimea
Christenh., comb. nov. Basionym: Sorbus kitaibeliana
(Krym).
Baksay & Kárpáti, Feddes Repert. 62: 299. 1960.
Pyrus kuanensis (J.Fryer & B.Hylmö) M.F.Fay &
Pyrus kitaibelii (J.Fryer & B.Hylmö) M.F.Fay &
kuanensis J.Fryer & B.Hylmö, Cotoneasters Compreh.
kitaibelii J.Fryer & B.Hylmö, Cotoneasters Compreh.
Guide: 314. 2009.
Guide: 314. 2009.
Pyrus kullensis (B.Hylmö) M.F.Fay & Christenh., comb.
Pyrus kiukiangensis (T.T.Yu) M.F.Fay & Christenh.,
nov. Basionym: Cotoneaster kullensis B.Hylmö,
comb. nov. Basionym: Sorbus kiukiangensis T.T.Yu,
Svensk Bot. Tidskr. 87: 322. 1993.
Acta Phytotax. Sin. 8: 225. 1963.
Pyrus kusnetzovii (Zinserl.) M.F.Fay & Christenh.,
Pyrus klasterskyana (Kárpáti) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus kusnetzovii Zinserl., Fl.
comb. nov. Basionym: Sorbus klasterskyana Kárpáti
URSS 9: 397. 1939.
in Bertová, Fl. Slovenska IV/3: 437. 1992.
Pyrus kwangsiensis (H.L.Li) M.F.Fay & Christenh.,
Pyrus klotzii M.F.Fay & Christenh., nom. nov. Replaced
comb. nov. Basionym: Photinia kwangsiensis H.L.Li,
synonym: Cotoneaster vilmorianus G.Klotz, Wiss. Z.
Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe

109
J. Arnold Arbor. 26: 62. 1945. Guihaia 9: 305. 1989.

Pyrus kweitschoviensis (G.Klotz) M.F.Fay & Christenh., Pyrus leptophylla (E.F.Warb.) M.F.Fay & Christenh.,
comb. nov. Basionym: Cotoneaster kweitschoviensis comb. nov. Basionym: Sorbus leptophylla E.F.Warb.,
G.Klotz, Wiss. Z. Martin-Luther-Univ. Halle- Watsonia 4: 44. 1957.
Wittenberg, Math.-Naturwiss. Reihe 12: 785. 1963. Pyrus leveillei (J.Fryer & B.Hylmö) M.F.Fay & Christenh.,
Pyrus lacei (G.Klotz) M.F.Fay & Christenh., comb. nov. comb. nov. Basionym: Cotoneaster leveillei J.Fryer &
Basionym: Cotoneaster lacei G.Klotz, Wiss. Z. Martin- B.Hylmö, Cotoneasters Compreh. Guid: 314. 2009.
Luther-Univ. Halle-Wittenberg, Math.-Naturwiss. Pyrus leyana (Wilmott) M.F.Fay & Christenh., comb.
Reihe 15: 851. 1966. nov. Basionym: Sorbus leyana Wilmott, Proc. Linn.
Pyrus lactea (W.W.Sm.) M.F.Fay & Christenh., comb. Soc. Lond. 146: 78. 1934.
nov. Basionym: Cotoneaster lacteus W.W.Sm., Not. Pyrus lidjiangensis (G.Klotz) M.F.Fay & Christenh.,
Roy. Bot. Gard. Edinburgh 10: 23. 1917. comb. nov. Basionym: Cotoneaster lidjiangensis
Pyrus laetevirens (Rehder & E.H.Wilson) M.F.Fay & G.Klotz, Wiss. Z. Martin-Luther-Univ. Halle-
Christenh., comb. nov. Basionym: Cotoneaster Wittenberg, Math.-Naturwiss. Reihe 12: 773. 1963.
acutifolius Turcz. var. laetevirens Rehder & Pyrus ×liljeforsii (T.C.G.Rich) M.F.Fay & Christenh.,
E.H.Wilson, Pl. Wilson. (Sargent) 1: 159. 1912. comb. nov. Basionym: Sorbus ×liljeforsii T.C.G.Rich,
Pyrus lambertii (G.Klotz) M.F.Fay & Christenh., comb. Nordic J. Bot. 25: 339. 2007.
nov. Basionym: Cotoneaster lambertii G.Klotz, Wiss. Pyrus lindleyana (Wight & Arn.) M.F.Fay & Christenh.,
Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. comb. nov. Basionym: Photinia lindleyana Wight &
Reihe 19: 343. 1970. Arn., Prodr. Fl. Ind. Orient. 1: 302. 1834.
Pyrus lancasteri (J.Fryer & B.Hylmö) M.F.Fay & Pyrus lingshiensis (Rushforth) M.F.Fay & Christenh.,
Christenh., comb. nov. Basionym: Cotoneaster comb. nov. Basionym: Sorbus lingshiensis Rushforth,
lancasteri J.Fryer & B.Hylmö, New Plantsman 8: 237. Plantsman 13: 121. 1991.
2001. Pyrus linnaeoides M.F.Fay & Christenh., nom. nov.
Pyrus lancastriensis (E.F.Warb.) M.F.Fay & Christenh., Replaced synonym: Cotoneaster rubens W.W.Sm.
comb. nov. Basionym: Sorbus lancastriensis var. minimus T.T.Yu, Acta Phytotax. Sin. 8: 220. 1963,
E.F.Warb., Watsonia 4: 45. 1957. (Figure 12B) not: Pyrus minima Ley. Etymology: Named for its
Pyrus lanceolata (Hu) M.F.Fay & Christenh., comb. vegetative resemblance to Linnaea (Caprifoliaceae).
nov. Basionym: Rhaphiolepis lanceolata Hu, J. Pyrus lithomelo M.F.Fay & Christenh., nom. nov.
Arnold Arbor. 13: 335. 1932. Replaced synonym: Osteomeles glabrata Kunth,
Pyrus langei (G.Klotz) M.F.Fay & Christenh., comb. Nov. Gen. Sp. (ed. quarto) 6: 210. 1815, not: Pyrus
nov. Basionym: Cotoneaster langei G.Klotz, Wiss. Z. glabrata (Rehder) L.H.Bailey. Etymology: From
Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe Greek λίθος, stone and μήλο, apple.
21: 1000. 1972. Pyrus llangollenica M.F.Fay & Christenh., nom. nov.
Pyrus lantana M.F.Fay & Christenh., nom. nov. Replaced synonym: Sorbus cuneifolia T.C.G.Rich,
Replaced synonym: Cotoneaster harrovianus Watsonia 27: 209. 2009, not: Pyrus cuneifolia Guss.
E.H.Wilson, Gard. Chron. 51: 3. 1912, not: Pyrus Etymology: This species occurs near Llangollen,
harrowiana Balf.f. & W.W.Sm. (orthographic variant). Wales, hence the name.
Etymology: This species resembles Viburnum Pyrus lochengensis (T.T.Yu) M.F.Fay & Christenh.,
lantana (Adoxaceae) in leaf and fruit. comb. nov. Basionym: Photinia lochengensis T.T.Yu,
Pyrus lasiopetala (Hayata) M.F.Fay & Christenh., comb. Acta Phytotax. Sin. 8: 226. 1963.
nov. Basionym: Photinia lasiopetala Hayata, Icon. Pl. Pyrus lonetalensis (S.Hammel & Haynold) M.F.Fay
Formosan. 6: 17. 1916. & Christenh., comb. nov. Basionym: Sorbus
Pyrus laxiflora (Jacq. ex Lindl.) M.F.Fay & Christenh., lonetalensis S.Hammel & Haynold, Jahresh. Ges.
comb. nov. Basionym: Cotoneaster laxiflorus Jacq. Naturk. Württemberg 171: 79. 2015.
ex Lindl., Edwards’s Bot. Reg. 15: t. 1229. 1829. Pyrus longifolia (Decne.) M.F.Fay & Christenh., comb.
Pyrus legrei (Cornier) M.F.Fay & Christenh., comb. nov. nov. Basionym: Photinia longifolia Decne., Nouv.
Basionym: Sorbus legrei Cornier, Bull. Mens. Soc. Arch. Mus. Par. Ser. 1, 10: 142. 1874.
Linn. Lyon 78: 29. 2009. Pyrus longii (Rushforth) M.F.Fay & Christenh., comb.
Pyrus leighensis (T.C.G.Rich) M.F.Fay & Christenh., nov. Basionym: Sorbus longii Rushforth, Plantsman
comb. nov. Basionym: Sorbus leighensis T.C.G.Rich, 13: 117. 1991.
Watsonia 27: 370. 2009. Pyrus loquat M.F.Fay & Christenh., nom. nov. Replaced
Pyrus leiocalyca (S.Z.Huang) M.F.Fay & Christenh., synonym: Photinia dubia Lindl., Trans. Linn. Soc.
comb. nov. Basionym: Malus leiocalyca S.Z.Huang, London 13: 104. 1821, not: Pyrus dubia E.H.L.Krause.

Etymology: from Cantonese, lou gwat, the name for 2012.

the fruit (‘lu orange’). Pyrus majuscula (W.W.Sm.) M.F.Fay & Christenh.,
Pyrus loriformis (W.W.Sm.) M.F.Fay & Christenh., comb. nov. Basionym: Cotoneaster hebephyllus
comb. nov. Basionym: Photinia loriformis W.W.Sm., Diels var. majusculus W.W.Sm., Not. Roy. Bot. Gard.
Notes Roy. Bot. Gard. Edinburgh 10: 60. 1917. Edinburgh 10(46): 22. 1917.
Pyrus lucida (Schltdl.) M.F.Fay & Christenh., comb. Pyrus malipoensis (K.C.Kuan) M.F.Fay & Christenh.,
nov. Basionym: Cotoneaster lucidus Schltdl., Linnaea comb. nov. Basionym: Eriobotrya malipoensis
27: 541. 1854. K.C.Kuan, Acta Phytotax. Sin. 8: 231. 1963.
Pyrus ludlowii (G.Klotz) M.F.Fay & Christenh., comb. Pyrus mamajevii (Knjaz.) M.F.Fay & Christenh., comb.
nov. Basionym: Cotoneaster ludlowii G.Klotz, Wiss. nov. Basionym: Cotoneaster mamajevii Knjaz., Bot.
Z. Martin-Luther-Univ. Halle-Wittenberg, Math.- Zhurn. (Moscow & Leningrad) 92: 423. 2007.
Naturwiss. Reihe 12: 775. 1963. Pyrus mandshurica (Maxim.) M.F.Fay & Christenh.,
Pyrus lui M.F.Fay & Christenh., nom. nov. Replaced comb. nov. Basionym: Pyrus baccata L. var.
synonym: Sorbus yunnanensis L.T.Lu, Acta Phytotax. mandshurica Maxim., Bull. Acad. Imp. Sci. Saint-
Sin. 38: 279. 2000, not: Pyrus yunnanensis Franch. Pétersbourg 19: 170. 1873. (Figure 12C)
Etymology: This species is named for Ling Ti Lu, who Pyrus margaretae (M.Proctor) M.F.Fay & Christenh.,
first described it. comb. nov. Basionym: Sorbus margaretae M.Proctor,
Pyrus luristanica (G.Klotz) M.F.Fay & Christenh., comb. Watsonia 27: 210. 2009.
nov. Basionym: Cotoneaster luristanicus G.Klotz, Pyrus margittaiana (Jáv.) M.F.Fay & Christenh., comb.
Feddes Repert. 76: 201. 1967. nov. Basionym: Sorbus hostii (J.Jacq. ex Host) Heynh.
Pyrus luzoniensis (Merr.) M.F.Fay & Christenh., comb. subsp. margittaiana Jáv., Sched. Fl. Hung. Exs. 8: 27.
nov. Basionym: Photinia luzoniensis Merr., Publ. Bur. 1927.
Sci. Gov. Lab., 17: 18. 1904. Pyrus marquandii (G.Klotz) M.F.Fay & Christenh.,
Pyrus macallisteri (Rushforth) M.F.Fay & Christenh., comb. nov. Basionym: Cotoneaster marquandii
comb. nov. Basionym: Sorbus macallisteri Rushforth, G.Klotz, Bull. Bot. Surv. India 5: 209. 1963.
Plantsman 13: 122. 1991. Pyrus marronina (J.Fryer & B.Hylmö) M.F.Fay &
Pyrus macrantha (Merr.) M.F.Fay & Christenh., comb. Christenh., comb. nov. Basionym: Cotoneaster
nov. Basionym: Sorbus macrantha Merr., Brittonia 4: marroninus J.Fryer & B.Hylmö, Cotoneasters
78. 1941. Compreh. Guide: 314. 2009.
Pyrus maderensis (Lowe) M.F.Fay & Christenh., comb. Pyrus mars M.F.Fay & Christenh., nom. nov. Replaced
nov. Basionym: Sorbus aucuparia L. var. maderensis synonym: Crataegus subfusca Ledeb. ex Nordm.,
Lowe, Man. Fl. Madeira 1: 259. 1868. Bull. Sci. Acad. Imp. Sci. St-Pétersbourg 2: 313. 1837,
Pyrus madoniensis (Raimondo, G.Castellano, not: Pyrus subfusca Regel. Etymology: This species
Bazan & Schicchi) M.F.Fay & Christenh., comb. is named for the planet Mars, in turn named for the
nov. Basionym: Sorbus madoniensis Raimondo, Roman god of war.
G.Castellano, Bazan & Schicchi, Pl. Biosystems Pyrus matudae (Lundell) M.F.Fay & Christenh., comb.
146(Suppl.): 347. 2012. nov. Basionym: Photinia matudae Lundell, Contr.
Pyrus magnifica (J.Fryer & B.Hylmö) M.F.Fay & Univ. Michigan Herb. 4: 7. 1940.
Christenh., comb. nov. Basionym: Cotoneaster Pyrus medogensis (L.T.Lu & T.C.Ku) M.F.Fay &
magnificus J.Fryer & B.Hylmö, New Plantsman 5: Christenh., comb. nov. Basionym: Sorbus
138. 1998. medogensis L.T.Lu & T.C.Ku, Acta Phytotax. Sin. 40:
Pyrus magocsyana (Kárpáti) M.F.Fay & Christenh., 475. 2002.
comb. nov. Basionym: Sorbus magocsyana Kárpáti Pyrus megacarpa (Li H.Zhou & C.Y.Wu) M.F.Fay &
in Németh, Acta Bot. Hung. 52: 388. 2010. Christenh., comb. nov. Basionym: Pleiosorbus
Pyrus maia M.F.Fay & Christenh., nom. nov. Replaced megacarpa Li H.Zhou & C.Y.Wu, Acta Bot. Yunnan.
synonym: Sorbus ovalis McAll., Gen. Sorbus: 236. 22: 384. 2000, as ‘megacarpus’.
2005, not: Pyrus ovalis Willd. Etymology: From Greek Pyrus megaphylla (T.T.Yu & T.C.Ku) M.F.Fay &
Μάιος, May, the month when this species flowers. Christenh., comb. nov. Basionym: Photinia
Pyrus major (Cardot) M.F.Fay & Christenh., comb. nov. megaphylla T.T.Yu & T.C.Ku, Acta Phytotax. Sin. 18:
Basionym: Rhaphiolepis major Cardot, Notul. Syst. 493. 1980.
(Paris) 3: 380. 1918. Pyrus meierottii (N.Mey.) M.F.Fay & Christenh., comb.
Pyrus majoricensis (L.Sáez & Rosselló) M.F.Fay & nov. Basionym: Sorbus meierottii N.Mey., Ber. Bayer.
Christenh., comb. nov. Basionym: Cotoneaster Bot. Ges. Sonderband: 169. 2005.
majoricensis L.Sáez & Rosselló, Candollea 67: 245. Pyrus meinichii (Lindeb.) M.F.Fay & Christenh., comb.

111
nov. Basionym: Sorbus aucuparia L. subsp. meinichii nom. nud. (Figure 12D)
Lindeb. ex Hartman, Handb. Skand. Fl., ed. 11: 271. Pyrus migarica (Zinserl.) M.F.Fay & Christenh., comb.
1879 nov. Basionym: Sorbus migarica Zinserl., Fl. URSS 9:
Pyrus meiophylla (W.W.Sm.) M.F.Fay & Christenh., 398, 1939.
comb. nov. Basionym: Cotoneaster glaucophyllus Pyrus milensis (M.Lepší, K.Boublík, P.Lepší & P.Vít)
Franch. var meiophyllus W.W.Sm., Not. Roy. Bot. M.F.Fay & Christenh., comb. nov. Basionym: Sorbus
Gard., Edinburgh 10: 21. 1917, as ‘meiophylla’. milensis M.Lepší, K.Boublík, P.Lepší & P.Vít, Preslia
Pyrus mekongensis (Cardot) M.F.Fay & Christenh., 80: 235. 2008.
comb. nov. Basionym: Rhaphiolepis indica (L.) Lindl. Pyrus milkedandaensis (J.Fryer & B.Hylmö) M.F.Fay
var. mekongensis Cardot, Notul. Syst. (Paris) 3: 380. & Christenh., comb. nov. Basionym: Cotoneaster
1918. milkedandaensis J.Fryer & B.Hylmö, Cotoneasters
Pyrus melanion M.F.Fay & Christenh., nom. nov. Compreh. Guide: 314. 2009.
Replaced synonym: Cotoneaster vulgaris Lindl. var. Pyrus miniata (Rehder & E.H.Wilson) M.F.Fay &
melanocarpus Ledeb., Fl. Altaic. 2: 219. 1830, as Christenh., comb. nov. Basionym: Cotoneaster
‘melanocarpa’, not: Pyrus melanocarpa (Michx.) zabelii C.K.Schneid. var. miniatus Rehder &
Willd. Etymology: Named for Μελανίων (also known E.H.Wilson, Pl. Wilson. (Sargent) 3(3): 430. 1917.
as Hippomenes), a Calydonian hunstman in Greek Pyrus minuta (G.Klotz) M.F.Fay & Christenh., comb.
mythology, who used golden apples to distract nov. Basionym: Cotoneaster minutus G.Klotz, Wiss.
Atalanta so he could win the race and marry her. Z. Martin-Luther-Univ. Halle-Wittenberg, Math.-
Pyrus melanotricha (Franch.) M.F.Fay & Christenh., Naturwiss. Reihe 12: 763. 1963.
comb. nov. Basionym: Cotoneaster buxifolius Lindl. f. Pyrus mirabilis (G.Klotz & Krügel) M.F.Fay & Christenh.,
melanotrichus Franch., Pl. Delavay.: 224. 1890. comb. nov. Basionym: Cotoneaster mirabilis G.Klotz
Pyrus mercurius M.F.Fay & Christenh., nom. nov. & Krügel, Wiss. Z. Friedrich-Schiller-Univ. Jena,
Replaced synonym: Pyrus aria (L.) Ehrh. var. Math.-Naturwiss. Reihe 32: 909. 1983, as hybrid.
obtusifolia DC., Prodr. 2: 636. 1825, not: Pyrus Pyrus mitschurinii (A.K.Skvortsov & Maitul.) M.F.Fay
obtusifolia (Pers.) M.F.Fay & Christenh. Etymology: & Christenh., comb. nov. Basionym: Aronia
Named for the planet Mercury. mitschurinii A.K.Skvortsov & Maitul., Byull. Glavn.
Pyrus mergenthaleriana (N.Mey.) M.F.Fay & Christenh., Bot. Sada (Moscow) 126: 40. 1982.
comb. nov. Basionym: Sorbus mergenthaleriana Pyrus mongolica (Pojark.) M.F.Fay & Christenh., comb.
N.Mey., Ber. Bayer. Bot. Ges. Sonderband : 154. nov. Basionym: Cotoneaster mongolicus Pojark.,
2005. Not. Syst. Inst. Bot. Acad. Sci. USSR 17: 196. 1955.
Pyrus merguiensis (J.E.Vidal) M.F.Fay & Christenh., Pyrus monopyrena (W.W.Sm.) M.F.Fay & Christenh.,
comb. nov. Basionym: Eriobotrya merguiensis comb. nov. Basionym: Cotoneaster hebephyllus Diels
J.E.Vidal, Adansonia sér. 2, 5: 563. 1965. var. monopyrenus W.W.Sm., Not. Roy. Bot. Gard.
Pyrus merula M.F.Fay & Christenh., nom. nov. Replaced Edinburgh 10: 23. 1917.
synonym: Sorbus rockii P.D.Sell, Fl. Gr. Brit. Ireland Pyrus ×monstrosa (Zabel) M.F.Fay & Christenh., comb.
2: 521. 2014, not: Pyrus rockii (Rehder) M.F.Fay & nov. Basionym: Aronia ×monstrosa Zabel in Beissn.
Christenh. Etymology: From Latin for the blackbird & al., Handb. Laubholzben. 1: 677. 1903. (note:
(Turdus merula). Pyrus arbutifolia × Pyrus americana).
Pyrus meuselii (G.Klotz) M.F.Fay & Christenh., comb. Pyrus montisalpae (Bernátová & Májovský) M.F.Fay
nov. Basionym: Cotoneaster meuselii G.Klotz, Wiss. & Christenh., comb. nov. Basionym: Sorbus
Z. Martin-Luther-Univ. Halle-Wittenberg, Math.- montisalpae Bernátová & Májovský, Biologia
Naturwiss. Reihe 12: 777. 1963. (Bratislava) 58: 788. 2003.
Pyrus mexicana (Baill.) M.F.Fay & Christenh., comb. Pyrus moravica (M.Lepší & P.Lepší) M.F.Fay &
nov. Basionym: Chamaemeles mexicana Baill., Christenh., comb. nov. Basionym: Sorbus moravica
Adansonia 9: 148. 1869. M.Lepší & P.Lepší, Preslia 87: 133. 2015.
Pyrus meyeri (Pojark.) M.F.Fay & Christenh., comb. Pyrus morrisiana M.F.Fay & Christenh., nom. nov.
nov. Basionym: Cotoneaster meyeri Pojark., Not. Replaced synonym: Sorbus japonica Siebold, Verh.
Syst. Inst. Bot. Acad. Sci. USSR 17: 185. 1955. Batav. Genootsch. Kunst. 12: 67. 1830, not: Pyrus
Pyrus microphylla (Wall. ex Lindl.) M.F.Fay & japonica Thunb. Etymology: This species is named
Christenh., comb. nov. Basionym: Cotoneaster in memory of Amy Morris, who encouraged Michael
microphyllus Wall. ex Lindl., Edwards’s Bot. Reg. 13: Fay to pursue a career in botany.
t. 1114. 1827 = Pyrus microphylla Wall., nom. inval. Pyrus morrisonensis (Hayata) M.F.Fay & Christenh.,
comb. nov. Basionym: Cotoneaster morrisonensis

Hayata, Icon. Pl. Formosa 5: 62. 1915. (Figure 12E) B.Hylmö, Cotoneasters Compreh. Guide: 315. 2009.
Pyrus morula (Pojark.) M.F.Fay & Christenh., comb. Pyrus natmataungensis (J.Fryer, B.Hylmö &
nov. Basionym: Cotoneaster morulus Pojark., Not. E.C.Nelson) M.F.Fay & Christenh., comb. nov.
Syst. Inst. Bot. Acad. Sci. USSR 21: 177 1961. Basionym: Cotoneaster natmataungensis J.Fryer,
Pyrus ×motleyi (T.C.G.Rich) M.F.Fay & Christenh., B.Hylmö & E.C.Nelson, Cotoneasters Compreh.
comb. nov. Basionym: Sorbus ×motleyi T.C.G.Rich, Guide : 315. 2009.
Watsonia 27: 211. 2009. (note: Pyrus aucuparia × Pyrus nedoluzhkoi (Tzvelev) M.F.Fay & Christenh.,
Pyrus leyana). comb. nov. Basionym: Cotoneaster nedoluzhkoi
Pyrus moupinensis (Franch.) M.F.Fay & Christenh., Tzvelev, Bot. Zhurn. (Moscow & Leningrad) 87: 115.
comb. nov. Basionym: Cotoneaster moupinensis 2002.
Franch., Nouv. Arch. Mus. Hist., ser. 2, 8: 224. 1885. Pyrus needhamii (Rushforth) M.F.Fay & Christenh.,
Pyrus mucronata (Franch.) M.F.Fay & Christenh., comb. nov. Basionym: Sorbus needhamii Rushforth,
comb. nov. Basionym: Cotoneaster mucronatus Bot. Mag. 27: 378. 2010.
Franch., Pl. Delavay. 3: 223. 1890. Pyrus nefedovii (Galushko) M.F.Fay & Christenh.,
Pyrus mulica M.F.Fay & Christenh., nom. nov. Replaced comb. nov. Basionym: Cotoneaster nefedovii
synonym: Sorbus muliensis McAll., Gen. Sorbus: Galushko, Derev. Kustarnik. Severn. Kavkaza : 216.
231. 2005, not: Pyrus muliensis (G.Klotz) M.F.Fay & 1967.
Christenh. Etymology: This species occurs in Muli, Pyrus neglecta (Hedl.) M.F.Fay & Christenh., comb.
China. nov. Basionym: Sorbus arranensis [unranked]
Pyrus muliensis (G.Klotz) M.F.Fay & Christenh., comb. neglecta Hedl., Skr. Vidensk.-Selsk. Kristiania. Mat.-
nov. Basionym: Cotoneaster muliensis G.Klotz, Wiss. Naturvidensk. Kl. 1914: 181. 1915.
Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Pyrus neimongolica M.F.Fay & Christenh., nom. nov.
Reihe 17: 336. 1968. Replaced synonym: Cotoneaster dielsianus E.Pritz.
Pyrus multiflora (Bunge) M.F.Fay & Christenh., comb. ex Diels var. globosus Hurus., Acta Phytotax. Geobot.
nov. Basionym: Cotoneaster multiflorus Bunge in 13: 233. 1943, not: Pyrus globosa (T.T.Yu & H.T.Tsai)
Ledeb., Fl. Altaica 2: 220. 1830. M.F.Fay & Christenh. Etymology: Named for the
Chinese autonomous region of Nei Mongol or Inner
Pyrus multijuga (Koehne) M.F.Fay & Christenh., comb.
Mongolia, where this species is found.
nov. Basionym: Sorbus multijuga Koehne, Pl. Wilson.
(Sargent) 1: 472. 1913. Pyrus neoantoninae (A.Vassil.) M.F.Fay & Christenh.,
comb. nov. Basionym: Cotoneaster neoantoninae
Pyrus munda (Koehne) M.F.Fay & Christenh., comb.
A.Vassil., Fl. Kazakhstana 6: 444. 1963.
nov. Basionym: Sorbus munda Koehne, Pl. Wilson.
(Sargent) 1: 469. 1913. Pyrus neopinnatifida (P.D.Sell) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus neopinnatifida P.D.Sell
Pyrus nagaensis (G.Klotz) M.F.Fay & Christenh., comb.
in Sell & Murrell, Fl. Gr. Brit. Ireland 2: 523. 2014.
nov. Basionym: Cotoneaster nagaensis G.Klotz, Bull.
Bot. Surv. India 5: 212. 1963. Pyrus neopopovii (Czerep.) M.F.Fay & Christenh.,
comb. nov. Basionym: Cotoneaster neopopovii
Pyrus naninitens (J.Fryer & B.Hylmö) M.F.Fay &
Czerep., Sosud. Rast. SSSR: 433. 1981.
naninitens J.Fryer & B.Hylmö, Cotoneasters Pyrus nepalensis (André) M.F.Fay & Christenh., comb.
Compreh. Guide : 315. 2009. nov. Basionym: Cotoneaster nepalensis André, Illust.
Hort. 22: 95. 1875, not: Pyrus nepalensis Loudon,
Pyrus nanshan (M.Vilm. ex Mottet) M.F.Fay &
nom. inval., nom. nud., nor: Pyrus nepalensis Hook.f.,
nom inval. pro syn.
nanshan M.Vilm. ex Mottet, Arbres Arbust. Ornem.
Pleine Terre 207: 1925, as ‘Nan-Shan’. Pyrus neptunus M.F.Fay & Christenh., nom. nov.
Replaced synonym: Sorbus arguta T.T.Yu, Acta
Pyrus nantouensis (J.Fryer & B.Hylmö) M.F.Fay &
Phytotax. Sin. 8: 223. 1963, not: Pyrus arguta (Wall.
nantouensis J.Fryer & B.Hylmö, Cotoneasters ex Lindl.) M.F.Fay & Christenh. Etymology: This
Compreh. Guide : 315. 2009. species is named for the planet Neptune.
Pyrus naoujanensis (J.Fryer & B.Hylmö) M.F.Fay & Pyrus newryensis (Barbier) M.F.Fay & Christenh.,
Christenh., comb. nov. Basionym: Cotoneaster comb. nov. Basionym: Cotoneaster newryensis
naoujanensis J.Fryer & B.Hylmö, Cotoneasters Barbier, Barbier & Son nursery trade list, Orléans,
Compreh. Guide : 315. 2009. France: 121. 1908.
Pyrus narynensis (Tkatsch. ex J.Fryer & B.Hylmö) Pyrus newtoniana M.F.Fay & Christenh., nom. nov.
M.F.Fay & Christenh., comb. nov. Basionym: Replaced synonym: Sorbus aestivalis Koehne, Pl.
Cotoneaster narynensis Tkatsch. ex J.Fryer & Wilson. (Sargent) 1: 469. 1913, not: Pyrus aestivalis

113
Lamotte. Etymology: Named for Isaac Newton, who Roy. Bot. Gard. Edinburgh 10: 29. 1917.
is said to have been inspired to formulate his theory Pyrus obscura (Rehder & E.H.Wilson) M.F.Fay &
of gravitation by watching an apple fall from a tree. Christenh., comb. nov. Basionym: Cotoneaster
Pyrus niitakayamensis (Hayata) M.F.Fay & Christenh., obscurus Rehder & E.H.Wilson, Pl. Wilson. (Sargent)
comb. nov. Basionym: Photinia niitakayamensis 1: 161. 1912.
Hayata, J. Coll. Sci. Imp. Univ. Tokyo 30(Art. 1): 103. Pyrus obtusa (Wall. ex Lindl.) M.F.Fay & Christenh.,
1911. comb. nov. Basionym: Cotoneaster obtusus Wall. ex
Pyrus nitens (Rehder & E.H.Wilson) M.F.Fay & Lindl., Edwards’s Bot. Reg. 15: t. 1229. 1829.
Christenh., comb. nov. Basionym: Cotoneaster Pyrus obtusifolia (Pers.) M.F.Fay & Christenh., comb.
nitens Rehder & E.H.Wilson, Pl. Wilson. (Sargent) 1: nov. Basionym: Crataegus obtusifolia Pers., Syn. Pl.
156. 1912. 2: 37. 1806.
Pyrus nitidifolia (C.Marquand) M.F.Fay & Christenh., Pyrus obtusisepala (Gand.) M.F.Fay & Christenh.,
comb. nov. Basionym: Cotoneaster nitidifolius comb. nov. Basionym: Cotoneaster obtusisepalus
C.Marquand, Hooker’s Icon. Pl. 32(2): pl. 3145. Gand., Fl. Lyon.: 87. 1875.
1930. Replaced synonym: Cotoneaster glomerulatus Pyrus ochracea (Hand.-Mazz.) M.F.Fay & Christenh.,
W.W.Sm., Not. Roy. Bot. Gard. Edinburgh 10: 21. comb. nov. Basionym: Eriobotrya ochracea Hand.-
1917, not: Pyrus glomerulata (Koehne) Bean. Mazz., Symb. Sin. 7: 476. 1933.
Pyrus nitidus (Jacq.) M.F.Fay & Christenh., comb. Pyrus ogisui (J.Fryer & B.Hylmö) M.F.Fay & Christenh.,
nov. Basionym: Cotoneaster nitidus Jacq., J. Soc. comb. nov. Basionym: Cotoneaster ogisui J.Fryer &
Imp. Centr. Hort. 3: 516. 1857. Replaced synonym: B.Hylmö, Cotoneasters Compreh. Guide : 316. 2009.
Cotoneaster rotundifolius Wall. ex Lindl., Edwards’s
Pyrus oleaster M.F.Fay & Christenh., nom. nov.
Bot. Reg. 15: t. 1229. 1829, not: Pyrus rotundifolia
Replaced synonym: Cotoneaster shansiensis J.Fryer
Moench.
& B.Hylmö, New Plantsman 5: 135. 1998, not:
Pyrus nohelii (J.Fryer & B.Hylmö) M.F.Fay & Christenh., Pyrus shansiensis Nakai. Etymology: This species
comb. nov. Basionym: Cotoneaster nohelii J.Fryer & superficially resembles an oleaster, Elaeagnus
B.Hylmö, Cotoneasters Compreh. Guide : 315. 2009. (Elaeagnaceae).
Pyrus norvegica M.F.Fay & Christenh., nom. nov. Pyrus oligantha (Pojark.) M.F.Fay & Christenh., comb.
Replaced synonym: Sorbus lancifolia Hedl., Skr. nov. Basionym: Cotoneaster oliganthus Pojark., Not.
Vidensk.-Selsk. Kristiania. Mat.- Naturvidensk. Kl. Syst. Inst. Bot. Acad. Sci. USSR 8: 141. 1940.
1911(6): 166. 1912, not: Pyrus lancifolia (Rehder)
Pyrus olivacea (McAll.) M.F.Fay & Christenh., comb.
L.H.Bailey. Etymology: This species is found in
nov. Basionym: Sorbus olivacea McAll., Gen. Sorbus:
Norway, hence the name.
232. 2005.
Pyrus notabilis (G.Klotz) M.F.Fay & Christenh., comb.
Pyrus ombrophila (Hand.-Mazz.) M.F.Fay & Christenh.,
nov. Basionym: Cotoneaster notabilis G.Klotz, Wiss.
comb. nov. Basionym: Malus ombrophila Hand.-
Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss.
Mazz., Anz. Akad. Wiss. Wien, Math.-Naturwiss. Kl.
Reihe 21: 972. 1972.
63: 8. 1926.
Pyrus nummularia (Fisch. & C.A.Mey.) M.F.Fay &
Pyrus omissa (J.Fryer & B.Hylmö) M.F.Fay & Christenh.,
comb. nov. Basionym: Cotoneaster omissus J.Fryer &
nummularius Fisch. & C.A.Mey., Ind. Semin. Hort.
B.Hylmö, Cotoneasters Compreh. Guide: 316. 2009.
Bot. Petrop. 2: 34. 1835.
Pyrus orbicularis (Schltdl.) M.F.Fay & Christenh., comb.
Pyrus nummularioides (Pojark.) M.F.Fay & Christenh.,
nov. Basionym: Cotoneaster orbicularis Schltdl.,
comb. nov. Basionym: Cotoneaster nummularioides
Linnaea 27: 544. 1854.
Pojark., Not. Syst. Inst. Bot. Acad. Sci. USSR 16: 128.
1954. Pyrus ormastonii (G.Klotz) M.F.Fay & Christenh., comb.
nov. Basionym: Cotoneaster ormastonii G.Klotz,
Pyrus oblanceolata (Rehder & E.H.Wilson) M.F.Fay
Wiss. Z. Martin-Luther-Univ. Halle-Wittenberg,
& Christenh., comb. nov. Basionym: Stranvaesia
Math.-Naturwiss. Reihe 15: 850. 1866.
nussia var. oblanceolata Rehder & E.H.Wilson, Pl.
Wilson. 1: 193. 1913. Pyrus osteomeles M.F.Fay & Christenh., nom.
nov. Replaced synonym: Osteomeles schwerinii
Pyrus obliqua (Stapf) M.F.Fay & Christenh., comb. nov.
C.K.Schneid., Ill. Handb. Laubholzk. 1: 763. 1906,
Basionym: Photinia obliqua Stapf, Bot. Mag. 149:
as ‘schwerinae’, not: Pyrus schwerinii (C.K.Schneid.)
sub t. 9008. 1924.
M.F.Fay & Christenh. Etymology: Named for the
Pyrus obovata (W.W.Sm.) M.F.Fay & Christenh., comb. genus Osteomeles, now included in Pyrus.
nov. Basionym: Eriobotrya obovata W.W.Sm., Notes
Pyrus ottoschwartzii (G.Klotz) M.F.Fay & Christenh.,
comb. nov. Basionym: Cotoneaster ottoschwartzii

G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.- 2005.

Naturwiss. Reihe 24: 402. 1975. Pyrus parvifructa (McAll.) M.F.Fay & Christenh., comb.
Pyrus ovata (Pojark.) M.F.Fay & Christenh., comb. nov. nov. Basionym: Sorbus parvifructa McAll., Gen.
Basionym: Cotoneaster ovatus Pojark., Not. Syst. Sorbus: 231. 2005.
Inst. Bot. Acad. Sci. USSR 16: 111. 1954. Pyrus parviloba (T.C.G.Rich) M.F.Fay & Christenh.,
Pyrus palcensis (C.K.Schneid.) M.F.Fay & Christenh., comb. nov. Basionym: Sorbus parviloba T.C.G.Rich,
comb. nov. Basionym: Hesperomeles palcensis Watsonia 27: 306. 2009.
C.K.Schneid., Bot. Jahrb. Syst. 42: 86. 1908. Pyrus pauca (M.Lepší & P.Lepší) M.F.Fay & Christenh.,
Pyrus pallasiana (Juz.) M.F.Fay & Christenh., comb. comb. nov. Basionym: Sorbus pauca M.Lepší &
nov. Basionym: Malus pallasiana Juz., Fl. URSS 9: P.Lepší, Preslia 85: 69. 2013.
370. 1939. Pyrus paucinerva (Merr.) M.F.Fay & Christenh., comb.
Pyrus pallescens (Rehder) M.F.Fay & Christenh., comb. nov. Basionym: Sorbus paucinerva Merr., Brittonia 4:
nov. Basionym: Sorbus pallescens Rehder, Pl. Wilson. 75. 1941.
(Sargent) 2: 266. 1915. Pyrus paxiana (Jáv.) M.F.Fay & Christenh., comb. nov.
Pyrus pangiensis (G.Klotz) M.F.Fay & Christenh., Basionym: Sorbus paxiana Jáv., Magyar Bot. Lapok
comb. nov. Basionym: Cotoneaster pangiensis 25: 89. 1927.
G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.- Pyrus peduncularis (Boiss.) M.F.Fay & Christenh.,
Naturwiss. Reihe 19: 342. 1970. comb. nov. Basionym: Cotoneaster peduncularis
Pyrus pannona (Franch.) M.F.Fay & Christenh., comb. Boiss., Diagn. Pl. Orient. 3: 8. 1943.
nov. Basionym: Cotoneaster pannonus Franch., Pl. Pyrus pekarovae (Májovský & Bernátová) M.F.Fay &
Delavay. 3: 223. 1890. Christenh., comb. nov. Basionym: Sorbus pekarovae
Pyrus pannonica (Kárpáti) M.F.Fay & Christenh., comb. Májovský & Bernátová, Biologia (Bratislava) 51: 25.
nov. Basionym: Sorbus pannonica Kárpáti, Borbásia 1996.
Nova 25: 10. 1944. Pyrus pelsoensis (C.Németh) M.F.Fay & Christenh.,
Pyrus paradisii M.F.Fay & Christenh., nom. nov. comb. nov. Basionym: Sorbus pelsoensis C.Németh,
Replaced synonym: Malus orientalis Uglitzk., Trudov Stud. Bot. Hung. 46: 50. 2015.
Sev.-Kavk. Inst. Spets. Tekhn. Kul’tur 1: 18. 1932, Pyrus pendula M.F.Fay & Christenh., nom. nov.
not: Pyrus orientalis Pall. Etymology: Named for the Replaced synonym: Cotoneaster glaucophyllus
‘apple of paradise’. Franch. var. vestitus W.W.Sm., Not. Roy. Bot. Gard.,
Pyrus paradoxa (G.Klotz) M.F.Fay & Christenh., Edinburgh 10: 21. 1917, not: Pyrus vestita Wall.
comb. nov. Basionym: Cotoneaster paradoxus ex G.Don. Etymology: This species has pendent
G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.- branches.
Naturwiss. Reihe 27: 24. 1978. Pyrus perlonga (Meierott) M.F.Fay & Christenh., comb.
Pyrus paris M.F.Fay & Christenh., nom. nov. Replaced nov. Basionym: Sorbus perlonga Meierott, Ber.
synonym: Malus dasyphylla Borkh., Theor. Prakt. Bayer. Bot. Ges. Sonderband: 187. 2005.
Handb. Forstbot. 2: 1269. 1803, not: Pyrus dasyphylla Pyrus perpusilla (C.K.Schneid.) M.F.Fay & Christenh.,
Bluff & Fingerh. Etymology: Named for Πάρις, Prince comb. nov. Basionym: Cotoneaster horizontalis
of Troy. In Greek mythology (Judgment of Paris), Eris Decne var. perpusillus C.K.Schneid., Ill. Handb.
was not invited at the marriage of Peleus and Thetis. Laubholzk. 1: 745. 1906.
She was not amused, so she gave Paris a golden Pyrus persica (Hedl.) M.F.Fay & Christenh., comb.
apple inscribed with the word καλλίστῃ (kallistēi, nov. Basionym: Sorbus persica Hedl., Kongl. Svenska
“for the fairest”) to take to the wedding and give to Vetensk.-Akad. Handl. 35: 70. 1901.
the most beautiful of all goddesses, which did not
Pyrus petiolata (Hook.f.) M.F.Fay & Christenh., comb.
go down well.
nov. Basionym: Eriobotrya petiolata Hook.f., Fl. Brit.
Pyrus parkeri (G.Klotz) M.F.Fay & Christenh., comb. India 2: 370. 1878.
nov. Basionym: Cotoneaster parkeri G.Klotz, Wiss. Z.
Pyrus philippinensis (S.Vidal) M.F.Fay & Christenh.,
Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe
comb. nov. Basionym: Eriobotrya philippinensis
15: 850. 1966, as hybrid.
S.Vidal, Revis. Pl. Vasc. Filip.: 123. 1886.
Pyrus parnassica (Boiss. & Heldr.) M.F.Fay & Christenh.,
Pyrus photinia M.F.Fay & Christenh., nom. nov.
comb. nov. Basionym: Cotoneaster parnassicus
Replaced synonym: Photinia salicifolia C.Presl., Abh.
Boiss. & Heldr., Diagn. Pl. Orient. 2: 48. 1856.
Königl. Böhm. Ges. Wiss. ser. 5, 6: 564. 1851, not:
Pyrus parva (McAll.) M.F.Fay & Christenh., comb. nov. Pyrus salicifolia Pall., nor: Pyrus arbutifolia (L.) L.f.
Basionym: Sorbus parva McAll., Gen. Sorbus: 234. Etymology: This species is named for the former

115
genus Photinia, now included in Pyrus. (Figure 12E) Tadzhikskoi SSR 4: 537. 1975.
Pyrus pilosicalyx (T.T.Yu) M.F.Fay & Christenh., comb. Pyrus porrigentiformis (E.F.Warb.) M.F.Fay &
nov. Basionym: Photinia pilosicalyx T.T.Yu, Acta Christenh., comb. nov. Basionym: Sorbus
Phytotax. Sin. 8: 231. 1963. porrigentiformis E.F.Warb., Watsonia 4: 45. 1957.
Pyrus piranga M.F.Fay & Christenh., nom. nov. Pyrus portae-bohemicae (M.Lepší, P.Lepší, P.Vít
Replaced synonym: Cotoneaster cardinalis J.Fryer & & K.Boublík) M.F.Fay & Christenh., comb. nov.
B.Hylmö, Cotoneasters Compreh. Guide: 309. 2009, Basionym: Sorbus portae-bohemicae M.Lepší,
not: Pyrus cardinalis Poit. & Turpin. Etymology: The P.Lepší, P.Vít & K.Boublík, Preslia 81: 72. 2009.
fruit of this species is as red as a tanager (genus Pyrus potaninii (Pojark.) M.F.Fay & Christenh., comb.
Piranga), hence the name. nov. Basionym: Cotoneaster potaninii Pojark., Not.
Pyrus platyphylla (Merr.) M.F.Fay & Christenh., Syst. Inst. Bot. Acad. Sci. USSR 21: 202. 1961.
comb. nov. Basionym: Eriobotrya platyphylla Merr., Pyrus poteriifolia (Hand.-Mazz.) M.F.Fay & Christenh.,
Brittonia 4: 80. 1941. comb. nov. Basionym: Sorbus poteriifolia Hand.-
Pyrus pluriflora (G.Klotz) M.F.Fay & Christenh., comb. Mazz., Anz. Akad. Wiss. Wien, Math.-Naturwiss. Kl.
nov. Basionym: Cotoneaster pluriflorus G.Klotz, 62: 223. 1925.
Wiss. Z. Martin-Luther-Univ. Halle-Wittenberg, Pyrus pourthiaea M.F.Fay & Christenh., nom. nov.
Math.-Naturwiss. Reihe 12: 781. 1963. Replaced synonym: Pourthiaea lucida Decne., Nouv.
Pyrus pluto M.F.Fay & Christenh., nom. nov. Replaced Arch. Mus. Par. Ser. 1, 10: 148. 1874, not: Pyrus
synonym: Stranvaesia microphylla Vidal, Notul. Syst. lucida (Schltdl.) M.F.Fay & Christenh. Etymology: This
(Paris) 13: 300. 1949, not: Pyrus microphylla (Wall. species is named for the former genus Pourthiaea,
ex Lindl.) M.F.Fay & Christenh. Etymology: This now included in Pyrus.
species is named for the dwarf planet Pluto. Pyrus ×pozdnjakovii (Pojark.) M.F.Fay & Christenh.,
Pyrus podocarpifolia (T.T.Yu) M.F.Fay & Christenh., comb. nov. Basionym: ×Sorbocotoneaster
comb. nov. Basionym: Photinia podocarpifolia T.T.Yu, pozdnjakovii Pojark., Bot. Mater. Gerb. Bot. Inst.
Acta Phytotax. Sin. 8: 230. 1963. Komarova Akad. Nauk S.S.S.R. 15: 93. 1953. (= P.
Pyrus poilanei (J.E.Vidal) M.F.Fay & Christenh., sibirica × P. melanocarpa)
comb. nov. Basionym: Eriobotrya poilanei J.E.Vidal, Pyrus prinoides (Rehder & E.H.Wilson) M.F.Fay &
Adansonia sér. 2, 5: 557. 1965. Christenh., comb. nov. Basionym: Eriobotrya
Pyrus polgariana (C.Németh) M.F.Fay & Christenh., prinoides Rehder & E.H.Wilson, Pl. Wilson. (Sargent)
comb. nov. Basionym: Sorbus polgariana C.Németh, 1(2): 194. 1912.
Acta Bot. Hung. 54: 132. 2012. Pyrus prionophylla (Franch.) M.F.Fay & Christenh.,
Pyrus poluninii (G.Klotz) M.F.Fay & Christenh., comb. comb. nov. Basionym: Eriobotrya prionophylla
nov. Basionym: Cotoneaster poluninii G.Klotz, Wiss. Franch., Pl. Delevay. 225. 1890.
Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Pyrus pritzellii (C.K.Schneid.) M.F.Fay & Christenh.,
Reihe 27: 21. 1978. comb. nov. Basionym: Cotoneaster rugosus E.Pritz.
Pyrus polyanthema (E.L.Wolf) M.F.Fay & Christenh., ex Diels var. pritzellii C.K.Schneid., Ill. Handb.
comb. nov. Basionym: Cotoneaster polyanthemus Laubholzk. 1: 758. 1906. Replaced synonym:
E.L.Wolf, Mitt. Deutsch. Dendrol. Gesell. 34: 325. Cotoneaster rugosus E.Pritz. ex Diels var. henryanus
1924. C.K.Schneid., Ill. Handb. Laubholzk. 1: 758. 1906,
not: Pyrus henryana Poit. & Turpin
Pyrus pomum M.F.Fay & Christenh., nom. nov.
Replaced synonym: Pyrus malus L. var. sylvestris L., Pyrus ×proctoriana (T.C.G.Rich) M.F.Fay & Christenh.,
Sp. Pl. 1: 479. 1753, not: Pyrus sylvestris Moench. comb. nov. Basionym: Sorbus ×proctoriana
Etymology: From Latin pomum, apple or fruit. T.C.G.Rich, Watsonia 27: 370. 2009.
(Figure 13A) Pyrus procumbens (G.Klotz) M.F.Fay & Christenh.,
Pyrus pontica (Zaik.) M.F.Fay & Christenh., comb. nov. comb. nov. Basionym: Cotoneaster procumbens
Basionym: Sorbus pontica Zaik., Bot. Zhurn. (Moscow G.Klotz, Wiss. Z. Martin-Luther-Univ. Halle-
& Leningrad) 67: 1293. 1982. Wittenberg, Math.-Naturwiss. Reihe 6: 982. 1957.
Pyrus pontis-satanae (M.Lepší & P.Lepší) M.F.Fay & Pyrus prostrata (Baker) M.F.Fay & Christenh., comb.
Christenh., comb. nov. Basionym: Sorbus pontis- nov. Basionym: Cotoneaster prostratus Baker,
satanae M.Lepší & P.Lepší, Preslia 87: 137. 2015, as Saunder’s Ref. Bot. 1: 53. 1869.
‘pontis-satani’. Pyrus pruinosa (G.Klotz) M.F.Fay & Christenh., comb.
Pyrus popovii (Pojark.) M.F.Fay & Christenh., comb. nov. Basionym: Cotoneaster pruinosus G.Klotz, Wiss.
nov. Basionym: Cotoneaster popovii Pojark., Fl. Z. Martin-Luther-Univ. Halle-Wittenberg, Math.-

Naturwiss. Reihe 15: 533. 1966. Replaced synonym: Cotoneaster obovatus Wall. ex
Pyrus przewalskii (Pojark.) M.F.Fay & Christenh., comb. Dunn, Bull. Misc. Inform. Kew 1921(3): 119. 1921,
nov. Basionym: Cotoneaster przewalskii Pojark., Not. not: Pyrus obovata (W.W.Sm.) M.F.Fay & Christenh.
Syst. Inst. Bot. Acad. Sci. USSR 21: 196. 1961. Etymology: This is a species found in the Punjab,
India, hence the name.
Pyrus pseudoambigua (J.Fryer & B.Hylmö) M.F.Fay
& Christenh., comb. nov. Basionym: Cotoneaster Pyrus purpurascens (J.Fryer & B.Hylmö) M.F.Fay &
pseudoambiguus J.Fryer & B.Hylmö, Watsonia 21: Christenh., comb. nov. Basionym: Cotoneaster
337. 1997. purpurascens J.Fryer & B.Hylmö, New Plantsman 8:
228. 2001.
Pyrus pseudobakonyensis (Kárpáti) M.F.Fay &
Christenh., comb. nov. Basionym: Sorbus Pyrus pyrenaica (Gand.) M.F.Fay & Christenh., comb.
pseudobakonyensis Kárpáti, Hung. Acta Biol. 1: 117. nov. Basionym: Cotoneaster pyrenaicus Gand., Dec.
1949. Pl. Nov. 1: 8. 1875.
Pyrus pseudofennica (E.F.Warb.) M.F.Fay & Christenh., Pyrus pyrrhula M.F.Fay & Christenh., nom. nov.
comb. nov. Basionym: Sorbus pseudofennica Replaced synonym: Aria mougeotii (Soy.-Will.
E.F.Warb., Watsonia 4: 43. 1957. & Godr.) Fourr. var. austriaca Beck, Fl. Nieder-
Österreich 2(1): 714. 1892., not: Pyrus austriaca
Pyrus pseudohupehensis (McAll.) M.F.Fay & Christenh.,
A.Kern. Etymology: From Greek πυρρός, meaning
comb. nov. Basionym: Sorbus pseudohupehensis
‘flame coloured’, ‘the colour of fire’ or simply ‘red’,
McAll., Gen. Sorbus : 231. 2005.
referring to the fruits.
Pyrus pseudolatifolia (Boros) M.F.Fay & Christenh.,
Pyrus qingshuiana M.F.Fay & Christenh., nom. nov.
comb. nov. Basionym: Sorbus pseudolatifolia Boros,
Replaced synonym: Cotoneaster chingshuiensis
Mitt. Kgl. Ungar. Gartenb.-Lehranst. 3: 51. 1937.
Kun C.Chang & Chih C.Wang, Taiwania 56: 126.
Pyrus pseudomeinichii (Ashley Robertson) M.F.Fay 2011, not: Pyrus chingshuiensis (T.Shimizu) M.F.Fay
& Christenh., comb. nov. Basionym: Sorbus & Christenh. Etymology: This species is found in
pseudomeinichii Ashley Robertson, Watsonia 26: 9. Quinshui, China.
2006.
Pyrus qungbixiensis (J.Fryer & B.Hylmö) M.F.Fay &
Pyrus pseudoobscura (J.Fryer & B.Hylmö) M.F.Fay Christenh., comb. nov. Basionym: Cotoneaster
& Christenh., comb. nov. Basionym: Cotoneaster qungbixiensis J.Fryer & B.Hylmö, J. Bot. Res. Inst.
pseudoobscurus J.Fryer & B.Hylmö, Cotoneasters Texas 2: 53. 2008.
Compreh. Guide: 316. 2009.
Pyrus raboutensis (Flinck, J.Fryer, Garraud, B.Hylmö
Pyrus pseudosemiincisa (Boros) M.F.Fay & Christenh., & J.Zeller) M.F.Fay & Christenh., comb. nov.
comb. nov. Basionym: Sorbus pseudosemiincisa Basionym: Cotoneaster raboutensis Flinck, J.Fryer,
Boros, Mitt. Kgl. Ungar. Gartenb.-Lehranst. 3: 53. Garraud, B.Hylmö & J.Zeller, Bull. Mens. Soc. Linn.
1937. Lyon 67: 276. 1998.
Pyrus pseudothuringiaca (Düll) M.F.Fay & Christenh., Pyrus racemiflora (Desf.) M.F.Fay & Christenh., comb.
comb. nov. Basionym: Sorbus pseudothuringiaca nov. Basionym: Mespilus racemiflora Desf., Tabl.
Düll in N.Mey., Ber. Bayer. Bot. Ges. Sonderband: 93. École Bot., ed. 3 (Cat. Pl. Horti Paris.) : 409. 1829.
2005.
Pyrus radicans (Dammer ex C.K.Schneid.) M.F.Fay
Pyrus pseudovertensis (Boros) M.F.Fay & Christenh., & Christenh., comb. nov. Basionym: Cotoneaster
comb. nov. Basionym: Sorbus pseudovertensis dammeri C.K.Schneid. var. radicans Dammer ex
Boros, Mitt. Kgl. Ungar. Gartenb.-Lehranst. 3: 53. C.K.Schneid., Ill. Handb Laubholzh. 1: 761. 1906.
1937.
Pyrus randaieneis (Hayata) M.F.Fay & Christenh.,
Pyrus pseudovilmorinii (McAll.) M.F.Fay & Christenh., comb. nov. Basionym: Pyrus aucuparia (L.) Gaertn.
comb. nov. Basionym: Sorbus pseudovilmorinii var. randaieneis Hayata, J. Coll. Sci., Imp. Univ. Tokyo
McAll., Gen. Sorbus : 235. 2005. 30: 98. 1911.
Pyrus pteridophylla (Hand.-Mazz.) M.F.Fay & Pyrus rannensis (B.Hylmö & J.Fryer) M.F.Fay &
Christenh., comb. nov. Basionym: Sorbus Christenh., comb. nov. Basionym: Cotoneaster
pteridophylla Hand.-Mazz., Symb. Sin. 7: 470. 1933. rannensis B.Hylmö & J.Fryer, Acta Bot. Fenn. 162:
Pyrus puellarum (Meierott) M.F.Fay & Christenh., 183. 1999.
comb. nov. Basionym: Sorbus puellarum Meierott, Pyrus ratisbonensis (N.Mey.) M.F.Fay & Christenh.,
Ber. Bayer. Bot. Ges. Sonderband : 190. 2005. comb. nov. Basionym: Sorbus ratisbonensis N.Mey.,
Pyrus pulchra (N.Mey.) M.F.Fay & Christenh., comb. Ber. Bayer. Bot. Ges. Sonderband : 158. 2005.
nov. Basionym: Sorbus pulchra N.Mey., Ber. Bayer. Pyrus raunii M.F.Fay & Christenh., nom. nov. Replaced
Bot. Ges. Sonderband : 106. 2005. synonym: Sorbus prattii Koehne, Pl. Wilson. (Sargent)
Pyrus punjabica M.F.Fay & Christenh., nom. nov. 1: 468. 1913, not: Pyrus prattii Hemsl. Etymology:

117
Named for the Finnish thundergod Rauni or Ukko, nipponensis).

who is often associated with a rowan tree (Pyrus Pyrus rinzenii (Rushforth) M.F.Fay & Christenh., comb.
aucuparia, Rosaceae). nov. Basionym: Sorbus rinzenii Rushforth, Plantsman
Pyrus raupingensis (K.C.Kuan) M.F.Fay & Christenh., n.s. 13: 114. 1991.
comb. nov. Basionym: Photinia raupingensis Pyrus ×robertsonii (T.C.G.Rich) M.F.Fay & Christenh.,
K.C.Kuan, Acta Phytotax. Sin. 8: 228. 1963. comb. nov. Basionym: Sorbus ×robertsonii
Pyrus redliana (Kárpáti) M.F.Fay & Christenh., comb. T.C.G.Rich, Watsonia 27: 370. 2009. (note: Pyrus aria
nov. Basionym: Sorbus redliana Kárpáti, Hung. Acta × Pyrus eminens).
Biol. 1: 118. 1949. Pyrus roborowskii (Pojark.) M.F.Fay & Christenh.,
Pyrus reducta (Diels) M.F.Fay & Christenh., comb. nov. comb. nov. Basionym: Cotoneaster roborowskii
Basionym: Sorbus reducta Diels, Notes Roy. Bot. Pojark., Not. Syst. Inst. Bot. Acad. Sci. USSR 21: 190.
Gard. Edinburgh 5: 272. 1912. 1961.
Pyrus reflexa (Carr.) M.F.Fay & Christenh., comb. nov. Pyrus ×robusta (Carrière) M.F.Fay & Christenh., comb.
Basionym: Cotoneaster reflexus Carr., Rev. Hort. 342: nov. Basionym: Malus microcarpa [unranked]
520. 1871. robusta Carrière, Étud. Gén. Pomm. Microcarp.: 76:
Pyrus reflexipetala (Koehne) M.F.Fay & Christenh., 1883. (note: Pyrus baccata × Pyrus prunifolia).
comb. nov. Basionym: Sorbus reflexipetala Koehne, Pyrus rockii (Rehder) M.F.Fay & Christenh., comb. nov.
Mitt. Deutsch. Dendrol. Ges. 15: 58. 1906. Basionym: Malus rockii Rehder, J. Arnold Arbor. 14:
Pyrus rehderi (Pojark.) M.F.Fay & Christenh., comb. 206. 1933.
nov. Basionym: Cotoneaster rehderi Pojark., Not. Pyrus rokujodaisanensis (Hayata) M.F.Fay &
Syst. Inst. Bot. Acad. Sci. USSR 17: 184. 1955. Christenh., comb. nov. Basionym: Cotoneaster
Pyrus reticulata (Rehder & E.H.Wilson) M.F.Fay & rokujodaisanensis Hayata, Icon. Pl. Formosa 5: 63.
Christenh., comb. nov. Basionym: Cotoneaster 1915.
reticulatus Rehder & E.H.Wilson, Pl. Wilson. Pyrus roopiana (Bordz.) M.F.Fay & Christenh., comb.
(Sargent) 1: 160. 1912. nov. Basionym: Sorbus roopiana Bordz., Izv. Kievsk.
Pyrus rhaphiolepis M.F.Fay & Christenh., nom. nov. Bot. Sada 12–13: 131. 1931.
Replaced synonym: Crataegus indica L., Sp. Pl. 1: Pyrus rosea (Edgew.) M.F.Fay & Christenh., comb. nov.
477. 1753, not: Pyrus indica Colebr. Etymology: Basionym: Cotoneaster roseus Edgew., Trans. Linn.
This species is named for the genus Rhaphiolepis, Soc. London 20: 46. 1846.
now included in Pyrus. Note: A new name is Pyrus rosiflora (Kun C.Chang & F.Y.Lu) M.F.Fay &
proposed here based on the Linnaean type. There Christenh., comb. nov. Basionym: Cotoneaster
are numerous alternative names, but it is currently rosiflorus Kun C.Chang & F.Y.Lu, Bot. Stud. (Taipei)
uncertain to what extent they refer to this species, 52: 214. 2011.
or to forms, varieties or cultivars of it. A thorough Pyrus royleana (Dippel) M.F.Fay & Christenh., comb.
revision of the nomenclature and taxonomy of nov. Basionym: Cotoneaster racemiflorus (Desf.)
this species and its varieties, forms and hybrids is Booth ex Bosse var. royleanus Dippel) J.Fryer &
needed. (Figure 13B) B.Hylmö, Handb. Laubholzk. 3: 415. 1893.
Pyrus rhodanthera (Kovanda) M.F.Fay & Christenh., Pyrus rubens (W.W.Sm.) M.F.Fay & Christenh., comb.
comb. nov. Basionym: Sorbus rhodanthera Kovanda, nov. Basionym: Cotoneaster rubens W.W.Sm., Not.
Verh. Zool.-Bot. Ges. Österreich 133: 321. 1996. Roy. Bot. Gard. Edinburgh 10: 24. 1917.
Pyrus rhombiformis (C.Németh) M.F.Fay & Christenh., Pyrus rubescens (McAll.) M.F.Fay & Christenh., comb.
comb. nov. Basionym: Sorbus rhombiformis nov. Basionym: Sorbus rubescens McAll., Gen.
C.Németh, Stud. Bot. Hung. 47: 305. 2016. Sorbus : 235. 2005.
Pyrus rhytidophylla (Rehder & E.H.Wilson) M.F.Fay Pyrus rubra M.F.Fay & Christenh., nom. nov. Replaced
& Christenh., comb. nov. Basionym: Cotoneaster synonym: Cotoneaster salwinensis G.Klotz, Wiss. Z.
rhytidophyllus Rehder & E.H.Wilson, Pl. Wislon. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe
(Sargent) 1: 175. 1912. 21: 1003. 1972, not: Pyrus salwinensis (Hand.-Mazz)
Pyrus richii (L.Houston) M.F.Fay & Christenh., comb. M.F.Fay & Christenh.. Etymology: This species has
nov. Basionym: Sorbus richii L.Houston, New J. Bot. red flowers followed by orange-red fruit, hence the
4: 5. 2014. epithet rubra, meaning red.
Pyrus ×rikuchuensis (Koidz.) M.F.Fay & Christenh., Pyrus rufopilosa (C.K.Schneid.) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus rikuchuensis Makino, comb. nov. Basionym: Sorbus rufopilosa
J. Jap. Bot. 6: 7. 1929. (note: Pyrus commixta × Pyrus C.K.Schneid., Bull. Herb. Boissier Ser. 2, 6: 317. 1906.
Pyrus rugosa (E.Pritzel) M.F.Fay & Christenh., comb.

A B
C D
E F
Figure 13: Angiosperms (Eudicots). A, Pyrus pomum M.F.Fay & Christenh.; B, Pyrus rhaphiolepis M.F.Fay &
Christenh.; C, Pyrus sargentii (Rehder) M.F.Fay & Christenh.; D, Pyrus speciosa (Sweet) M.F.Fay & Christenh.; E,
Pyrus umbellularia M.F.Fay & Christenh.; and F, Pyrus villosa (Thunb.) M.F.Fay & Christenh.. All images © Maarten
Christenhusz.

119
nov. Basionym: Cotoneaster rugosus E.Pritzel ex 13C)

Diels, Bot. Jahrb. Syst. 29: 385. 1900. Pyrus saturnus M.F.Fay & Christenh., nom. nov.
Pyrus rupicoloides (L.Houston & T.C.G.Rich) M.F.Fay Replaced synonym: Crataegus cuspidata Spach, Hist.
& Christenh., comb. nov. Basionym: Sorbus Nat. Vég. 2: 106. 1834, not: Pyrus cuspidata Bertol.
rupicoloides L.Houston & T.C.G.Rich, Watsonia 27: Etymology: Named for the planet Saturn.
291. 2009. Pyrus saxicola (T.C.G.Rich) M.F.Fay & Christenh., comb.
Pyrus rushforthii (McAll.) M.F.Fay & Christenh., comb. nov. Basionym: Sorbus saxicola T.C.G.Rich, Watsonia
nov. Basionym: Sorbus rushforthii McAll., Gen. 27: 307. 2009.
Sorbus: 232. 2005. Pyrus scandinavica (B.Hylmö) M.F.Fay & Christenh.,
Pyrus sinohimalaica M.F.Fay & Christenh., nom. nov. comb. nov. Basionym: Cotoneaster scandinavicus
Replaced synonym: Sorbus spongbergii Rushforth, B.Hylmö, Svensk Bot. Tidskr. 87: 317. 1993, not:
Int. Dendrol. Soc. Year Book 2009: 96. 2010, not: Pyrus scandinavica hort. ex Lavallée, nom. inval.,
Pyrus spongbergii (J.Fryer & B.Hylmö) M.F.Fay & nom. nud.
Christenh. Etymology: This species is found in the Pyrus scannelliana (T.C.G.Rich) M.F.Fay & Christenh.,
Chinese Himalayas, hence the name. comb. nov. Basionym: Sorbus scannelliana
Pyrus rutilans (McAll.) M.F.Fay & Christenh., comb. T.C.G.Rich, Watsonia 27: 213. 2009.
nov. Basionym: Sorbus rutilans McAll., Gen. Sorbus: Pyrus scepusiensis (Kovanda) M.F.Fay & Christenh.,
235. 2005. comb. nov. Basionym: Sorbus scepusiensis Kovanda,
Pyrus sabrina (Flinck & B.Hylmö) M.F.Fay & Christenh., Willdenowia 16: 119. 1986.
comb. nov. Basionym: Cotoneaster sabrina Hadden Pyrus schantungensis (G.Klotz) M.F.Fay & Christenh.,
ex Krüssm., Laubgehölze: 117. 1951, pro hybr. comb. nov. Basionym: Cotoneaster schantungensis
Synonym: Cotoneaster splendens Flinck & B.Hylmö, G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-
Bot. Not. (Lund) 117: 124. 1964, not: Pyrus splendens Naturwiss. Reihe 21: 1018. 1972.
Poit. & Turpin Pyrus schemachensis (Zinserl.) M.F.Fay & Christenh.,
Pyrus sachalinensis (Juz.) M.F.Fay & Christenh., comb. comb. nov. Basionym: Sorbus schemachensis
nov. Basionym: Malus sachalinensis Juz., Fl. URSS 9: Zinserl., Fl. URSS 9: 392. 1939.
372. 1939. Pyrus ×schinzii (Düll) M.F.Fay & Christenh., comb. nov.
Pyrus salatinii (Bernátová & Májovský) M.F.Fay & Basionym: Sorbus ×schinzii Düll, Ber. Bayer. Bot. Ges.
Christenh., comb. nov. Basionym: Sorbus salatinii 34: 61. 1961.
Bernátová & Májovský, Biologia (Bratislava) 58: 790. Pyrus schizomeles M.F.Fay & Christenh., nom. nov.
2003, as ‘salatini’. Replaced synonym: Cotoneaster latifolius J.Fryer &
Pyrus salwinensis (Hand.-Mazz.) M.F.Fay & Christenh., B.Hylmö, New Plantsman 8: 228. 2001, not: Pyrus
comb. nov. Basionym: Eriobotrya salwinensis Hand.- ×latifolia Lindl. ex Steud. Etymology: This species
Mazz., Symb. Sin. Pt. 7: 475. 1933. has pomes that split open when mature, hence the
Pyrus sandakphuensis (G.Klotz) M.F.Fay & Christenh., name.
comb. nov. Basionym: Cotoneaster sandakphuensis Pyrus schlechtendalii (G.Klotz) M.F.Fay & Christenh.,
G.Klotz, Bull. Bot. Surv. India 5: 213. 1963. comb. nov. Basionym: Cotoneaster schlechtendalii
Pyrus sapientiae M.F.Fay & Christenh., nom. nov. G.Klotz, Wiss. Z. Martin-Luther-Univ. Halle-
Replaced synonym: Malus montana Uglitzk., Trudov Wittenberg, Math.-Naturwiss. Reihe 12: 776. 1963.
Sev.-Kavk. Inst. Spets. Tekhn. Kul’tur 1(3): 18. 1932, Pyrus schneideriana (Rehder & E.H.Wilson) M.F.Fay
not: Pyrus montana Poit. & Turpin. Etymology: & Christenh., comb. nov. Basionym: Photinia
Named for the ‘apple of wisdom’. schneideriana Rehder & E.H.Wilson, Pl. Wilson.
Pyrus sargentaster M.F.Fay & Christenh., nom. nov. (Sargent) 1: 188. 1912.
Replaced synonym: Cotoneaster salicifolius Franch., Pyrus schnizleiniana (N.Mey.) M.F.Fay & Christenh.,
Nouv. Arch. Mus. Hist. Hat., sér. 2, 8: 225. 1886, not: comb. nov. Basionym: Sorbus schnizleiniana N.Mey.,
Pyrus salicifolia Pall. = Cotoneaster sargentii G.Klotz, Ber. Bayer. Bot. Ges. Sonderband: 146. 2005.
not Pyrus sargentii (Rehder) M.F.Fay & Christenh. Pyrus schubertii (G.Klotz) M.F.Fay & Christenh., comb.
Etymology: Named for Charles Sprague Sargent and nov. Basionym: Cotoneaster schubertii G.Klotz, Wiss.
aster, a star. Z. Martin-Luther-Univ. Halle-Wittenberg, Math.-
Pyrus sargentii (Rehder) M.F.Fay & Christenh., comb. Naturwiss. Reihe 12: 759. 1963.
nov. Basionym: Malus sargentii Rehder, Trees & Pyrus schuwerkiorum (N.Mey.) M.F.Fay & Christenh.,
Shrubs [Sargent] 1: 71. 1903. Note: The name Pyrus comb. nov. Basionym: Sorbus schuwerkiorum
sargentii hort. has been in use, but it was never N.Mey., Ber. Bayer. Bot. Ges. Sonderband : 162.
formally published as far as we can trace. (Figure

2005. Z. Martin-Luther-Univ. Halle-Wittenberg, Math.-

Pyrus schwarziana (N.Mey.) M.F.Fay & Christenh., Naturwiss. Reihe 12: 776. 1963.
comb. nov. Basionym: Sorbus schwarziana N.Mey., Pyrus shimlana M.F.Fay & Christenh., nom. nov.
Ber. Bayer. Bot. Ges. Sonderband : 110. 2005. Replaced synonym: Cotoneaster wattii G.Klotz,
Pyrus schwerinii (C.K.Schneid.) M.F.Fay & Christenh., Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-
comb. nov. Basionym: Micromeles schwerinii Naturwiss. Reihe 19: 342. 1970, not: Pyrus wattii
C.K.Schneid., Ill. Handb. Laubholzk. 1: 702. 1906, as (Koehne) Bennet. Etymology: Named for the Shimla
‘schwerini’. Mountains in Himachal Pradesh, India, where this
species occurs.
Pyrus scopulina (Greene) M.F.Fay & Christenh., comb.
nov. Basionym: Sorbus scopulina Greene, Pittonia 4: Pyrus shirinensis (E.Hadac & Chrtek) M.F.Fay &
130. 1900. Christenh., comb. nov. Basionym: Sorbus shirinensis
E.Hadac & Chrtek, Candollea 35: 325. 1980.
Pyrus seguinii (H.Lév.) M.F.Fay & Christenh., comb.
nov. Basionym: Symplocos seguinii H.Lév., Repert. Pyrus sichuanensis (G.Klotz) M.F.Fay & Christenh.,
Spec. Nov. Regni Veg. 10: 431. 1912. comb. nov. Basionym: Cotoneaster sichuanensis
Pyrus sellii (T.C.G.Rich) M.F.Fay & Christenh., comb.
Naturwiss. Reihe 21: 1013. 1972.
nov. Basionym: Sorbus sellii T.C.G.Rich, New J. Bot.
4: 7. 2014. Pyrus sikangensis (Flinck & B.Hylmö) M.F.Fay &
Pyrus semiincisa (Borbás) M.F.Fay & Christenh., comb.
sikangensis Flinck & B.Hylmö, Bot. Not. (Lund) 115:
nov. Basionym: Sorbus aria L. f. semiincisa Borbás,
376. 1962.
Term. Tud. Közl. 11: 34. 1879.
Pyrus silvestrii (Pamp.) M.F.Fay & Christenh., comb.
Pyrus sennikoviana M.F.Fay & Christenh., nom. nov.
nov. Basionym: Cotoneaster silvestrii Pamp., Nuov.
Replaced synonym: Cotoneaster ambiguus Rehder
Giorn. Bot. Ital. 17: 288. 1910.
& E.H.Wilson, Pl. Wilson. (Sargent) 1: 159. 1912,
not: Pyrus ×ambigua (Decne.) M.F.Fay & Christenh. Pyrus simonkaiana (Kárpáti) M.F.Fay & Christenh.,
Etymology: Named in honour of Alexander Sennikov, comb. nov. Basionym: Sorbus simonkaiana Kárpáti,
for his contributions to the taxonomy of Rosaceae in Agrártud. Egyet. Kert- Szölögazdaságtud. Karának
general and Cotoneaster in particular. Évk. 1: 38. 1950.
Pyrus serpentae M.F.Fay & Christenh., nom. nov. Pyrus simonsii (Baker) M.F.Fay & Christenh., comb.
Replaced synonym: Eriobotrya glabrescens J.E.Vidal, nov. Basionym: Cotoneaster simonsii Baker,
Adansonia sér. 2, 5: 554. 1965, not: Pyrus glabrescens Saunder’s Ref. Bot. 1: t. 55. 1869.
Cardot. Etymology: This species is named for the Pyrus slovenica (Kovanda) M.F.Fay & Christenh., comb.
snake that presented the apple of paradise to Eve in nov. Basionym: Sorbus slovenica Kovanda, Acta Univ.
the Book of Genesis. Carol., Biol. 1961: 73. 1961.
Pyrus serrata (J.E.Vidal) M.F.Fay & Christenh., comb. Pyrus smithii (G.Klotz) M.F.Fay & Christenh., comb.
nov. Basionym: Eriobotrya serrata J.E.Vidal, nov. Basionym: Cotoneaster smithii G.Klotz, Mitt.
Adansonia sér. 2, 5: 558. 1965. Deutsch. Dendrol. Ges. 82: 77. 1996.
Pyrus serratifolia (Desf.) M.F.Fay & Christenh., comb. Pyrus soczaviana (Pojark.) M.F.Fay & Christenh., comb.
nov. Basionym: Crataegus serratifolia Desf., Tabl. nov. Basionym: Cotoneaster soczavianus Pojark.,
École Bot. (ed. 3): 288. 1829. Not. Syst. Inst. Bot. Acad. Sci. USSR 17: 179. 1955.
Pyrus seyboldiana (S.Hammel & Haynold) M.F.Fay Pyrus sodomacea M.F.Fay & Christenh., nom. nov.
& Christenh., comb. nov. Basionym: Sorbus Replaced synonym: Rhaphiolepis ferruginea
seyboldiana S.Hammel & Haynold, Jahresh. Ges. F.P.Metcalf, Lingnan Sci. J. 18: 509. 1939, not:
Naturk. Württemberg 171: 52. 2015. Pyrus ferruginea Hook.f. Etymology: Named for the
Pyrus shannanensis (J.Fryer & B.Hylmö) M.F.Fay & mythological ‘Apple of Sodom’.
Christenh., comb. nov. Basionym: Cotoneaster Pyrus sognensis (Hedl. ex Sennikov, Hjertson &
shannanensis J.Fryer & B.Hylmö, New Plantsman 8: Salvesen) M.F.Fay & Christenh., comb. nov.
237. 2001. Basionym: Sorbus sognensis Hedl. ex Sennikov,
Pyrus sharmae (M.F.Watson, V.Manandhar Hjertson & Salvesen, Ann. Bot. Fenn. 53: 7. 2016.
& Rushforth) M.F.Fay & Christenh., comb. Pyrus ×sorbaronia M.F.Fay & Christenh., nom. nov.
nov. Basionym: Sorbus sharmae M.F.Watson, Replaced synonym: Mespilus ×sorbifolia hort. ex
V.Manandhar & Rushforth, Int. Dendrol. Soc. Year Poir., Encyc. [Lamarck] Suppl. 4: 72. 1816. (note:
Book 2009: 79. 2010. Pyrus americana × Pyrus melanocarpa), not: Pyrus
Pyrus sherriffii (G.Klotz) M.F.Fay & Christenh., comb. sorbifolia Bosc ex P.Watson. Etymology: This is
nov. Basionym: Cotoneaster sherriffii G.Klotz, Wiss. a hybrid between a former Sorbus and a former

121
Aronia species. Watsonia 27: 215. 2009.

Pyrus sordida (G.Klotz) M.F.Fay & Christenh., comb. Pyrus suavis (Pojark.) M.F.Fay & Christenh., comb. nov.
nov. Basionym: Cotoneaster sordidus G.Klotz, Mitt. Basionym: Cotoneaster suavis Pojark., Not. Syst. Inst.
Deutsch. Dendrol. Ges. 82: 72. 1996. Bot. Acad. Sci. USSR 16: 118. 1954.
Pyrus soulieana (G.Klotz) M.F.Fay & Christenh., Pyrus subacuta (Pojark.) M.F.Fay & Christenh., comb.
comb. nov. Basionym: Cotoneaster soulieanus nov. Basionym: Cotoneaster subacutus Pojark., Not.
G.Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.- Syst. Inst. Bot. Acad. Sci. USSR 21: 165. 1961.
Naturwiss. Reihe 21: 1014. 1972. Pyrus subadpressa (T.T.Yu) M.F.Fay & Christenh., comb.
Pyrus speciosa (Sweet) M.F.Fay & Christenh., comb. nov. Basionym: Cotoneaster subadpressus T.T.Yu,
nov. Basionym: Cydonia speciosa Sweet, Hort. Acta Phytotax. Sin. 8: 219. 1963.
Suburb. Lond.: 113. 1818. (Figure 13D) Pyrus subarranensis (Hyl. ex Sennikov, Hjertson
Pyrus spectans (L.Houston) M.F.Fay & Christenh., & Salvesen) M.F.Fay & Christenh., comb. nov.
comb. nov. Basionym: Sorbus spectans L.Houston, Basionym: Sorbus subarranensis Hyl. ex Sennikov,
New J. Bot. 4: 2. 2014. Hjertson & Salvesen, Ann. Bot. Fenn. 53: 9. 2016.
Pyrus spiraeaster M.F.Fay & Christenh., nom. nov. Pyrus subcuneata (Wilmott) M.F.Fay & Christenh.,
Replaced synonym: Cotoneaster racemiflorus comb. nov. Basionym: Sorbus subcuneata Wilmott,
(Desf.) Nooth ex Bosse var. microcarpus Rehder & Proc. Linn. Soc. Lond. 146: 76. 1934.
E.H.Wilson, Pl. Wilson. (Sargent) 1: 169. 1912, not: Pyrus subdanubialis (Soó) M.F.Fay & Christenh., comb.
Pyrus microcarpa (Pursh) DC. Etymology: From nov. Basionym: Sorbus aria L. f. subdanubialis Soó,
Greek σπιραιά, spirits, and αστέρ, star. Tisia 2: 222. 1937.
Pyrus ×splendida (Hedl.) M.F.Fay & Christenh., comb. Pyrus submultiflora (Popov) M.F.Fay & Christenh.,
nov. Basionym: Sorbus ×splendida Hedl., Kongl. comb. nov. Basionym: Cotoneaster submultiflorus
Svenska Vetensk.-Akad. Handl. 35: 43. 1901. Popov, Bull. Soc. Imp. Naturalistes Moscou 44: 126.
Pyrus spongbergii (J.Fryer & B.Hylmö) M.F.Fay & 1935.
Christenh., comb. nov. Basionym: Cotoneaster Pyrus subochracea (T.T.Yu & L.T.Lu) M.F.Fay &
spongbergii J.Fryer & B.Hylmö, Cotoneasters Christenh., comb. nov. Basionym: Sorbus
Compreh. Guide : 316. 2009. subochracea T.T.Yu & L.T.Lu, Acta Phytotax. Sin. 18:
Pyrus staceana M.F.Fay & Christenh., nom. nov. 494. 1980.
Replaced synonym: Cotoneaster melanocarpus Pyrus subpinnata (Hedl.) M.F.Fay & Christenh., comb.
(Ledeb.) Lodd., G.Lodd. & W.Lodd. ex M.Roem. nov. Basionym: Sorbus subpinnata Hedl., Nyt Mag.
var. commixtus C.K.Schneid., Ill. Handb. Laubholzk. Naturvidensk. 49: 198. 1911.
1: 752. 1906, not: Pyrus commixta (Hedl.) Pyrus subrotunda (K.Koch) M.F.Fay & Christenh.,
Cardot. Etymology: Named for Clive Stace, for his comb. nov. Basionym: Osteomeles subrotunda
contributions to Rosaceae taxonomy. K.Koch, Ann. Mus. Bot. Lugduno-Batavi 1: 250. 1864.
Pyrus staintonii (G.Klotz) M.F.Fay & Christenh., comb. Pyrus subsimilis (Hedl.) M.F.Fay & Christenh., comb.
nov. Basionym: Cotoneaster staintonii G.Klotz, Wiss. nov. Basionym: Sorbus subsimilis Hedl., Nyt Mag.
Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Naturvidensk. 52: 257. 1914.
Reihe 21: 971. 1972.
Pyrus subuniflora (Kitam.) M.F.Fay & Christenh., comb.
Pyrus stankovii (Juz.) M.F.Fay & Christenh., comb. nov. nov. Basionym: Cotoneaster racemiflorus (Desf.)
Basionym: Sorbus stankovii Juz., Bot. Mater. Gerb. K.Koch var. subuniflorus Kitam, Results Kyoto Univ.
Bot. Inst. Komarova Akad. Nauk SSSR 12: 7. 1950. Sci. Exp. Karakoram & Hindukush, 1955: 173. 1960.
Pyrus stenophylla (Hand.-Mazz.) M.F.Fay & Christenh., Pyrus sujoyi (Ghora) M.F.Fay & Christenh., comb. nov.
comb. nov. Basionym: Photinia stenophylla Hand.- Basionym: Sorbus sujoyi Ghora, Bull. Bot. Surv. India
Mazz., Symb. Sin. Pt. 7: 480. 1933. 49: 199. 2007.
Pyrus sterniana (Turrill) M.F.Fay & Christenh., comb. Pyrus ×superba (Frahm) M.F.Fay & Christenh., comb.
nov. Basionym: Cotoneaster franchetii Bois var. nov. Basionym: Cydonia maulei T.Moore var.
sternianus Turrill, Bot. Mag. 167: pl. 130. 1950, as superba Frahm, Gartenwelt 2: 214. 1898. (note:
‘sterniana’. Pyrus speciosa × Pyrus japonica)
Pyrus stipularis (Craib) M.F.Fay & Christenh., comb. Pyrus svenhedinii (J.Fryer & B.Hylmö) M.F.Fay &
nov. Basionym: Eriobotrya stipularis Craib, Bull. Christenh., comb. nov. Basionym: Cotoneaster
Misc. Inform. Kew 1929: 109. 1929. svenhedinii J.Fryer & B.Hylmö, Cotoneasters
Pyrus stirtoniana (T.G.C.Rich) M.F.Fay & Christenh., Compreh. Guide : 316. 2009.
comb. nov. Basionym: Sorbus stirtoniana T.G.C.Rich, Pyrus ×sverigensis M.F.Fay & Christenh., nom. nov.
Basionym: Cotoneaster ×suecicus G.Klotz, Beitr.

Phytotax. 10: 47. 1982. (note: P. conspicua × P. Basionym: Sorbus tenuis McAll., Gen. Sorbus: 233.
dammeri), not: Pyrus suecica Roehl. ex Steud. 2005.
Etymology: Named for Sverige, the Swedish name Pyrus teodori (Liljef.) M.F.Fay & Christenh., comb. nov.
for Sweden, where this hybrid was found. Basionym: Sorbus teodori Liljef., Acta Horti Berg. 16:
Pyrus symondsii (Standish ex T.Moore) M.F.Fay & 283. 1953.
Christenh., comb. nov. Basionym: Cotoneaster Pyrus tergestina (H.Lindb.) M.F.Fay & Christenh.,
symondsii Standish ex T.Moore, Proc. Roy. Hort. comb. nov. Basionym: Pyrus aria (L.) Ehrh. subsp.
Soc. London 1: 298. 1861. Replaced synonym: tergestina H.Lindb., Öfvers. Finska Vetensk.-Soc.
Cotoneaster marginatus (Loudon) Schltdl., Linnaea Förh. 48: 45. 1906.
27: 541. 1856, as ‘marginata’, not: Pyrus marginata Pyrus thaiszii (Soó) M.F.Fay & Christenh., comb. nov.
Kit. Basionym: Sorbus aria (L.) Crantz thaiszii Soó, Tisia
Pyrus takhtajanii (Gabrieljan) M.F.Fay & Christenh., 2: 223. 1937.
comb. nov. Basionym: Sorbus takhtajanii Gabrieljan, Pyrus thayensis (M.Lepší & P.Lepší) M.F.Fay &
Biol. Zhurn. Armenii 22: 39. 1969. Christenh., comb. nov. Basionym: Sorbus thayensis
Pyrus talgarica (M.Popov) M.F.Fay & Christenh., comb. M.Lepší & P.Lepší, Preslia 87: 148. 2015.
nov. Basionym: Cotoneaster talgaricus M.Popov, Pyrus thimphuensis (J.Fryer & B.Hylmö) M.F.Fay &
Bull. Soc. Imp. Naturalistes Moscou 44: 126. 1935. Christenh., comb. nov. Basionym: Cotoneaster
Pyrus tanpaensis (J.Fryer & B.Hylmö) M.F.Fay & thimphuensis J.Fryer & B.Hylmö, New Plantsman 8:
Christenh., comb. nov. Basionym: Cotoneaster 237. 2001.
tanpaensis J.Fryer & B.Hylmö, Cotoneasters Pyrus thunbergii M.F.Fay & Christenh., nom. nov.
Compreh. Guide: 317. 2009. Replaced synonym: Crataegus glabra Thunb., Syst.
Pyrus taoensis (G.Klotz) M.F.Fay & Christenh., comb. Veg. (ed. 14.): 465. 1784, not: Pyrus glabra Boiss.
nov. Basionym: Cotoneaster taoensis G.Klotz, Wiss. Etymology: This species is named for Carl Peter
Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Thunberg, who first described it.
Reihe 21: 1010. 1972. Pyrus thymifolia (Baker) M.F.Fay & Christenh., comb.
Pyrus taofuensis (J.Fryer & B.Hylmö) M.F.Fay & nov. Basionym: Cotoneaster thymifolius Baker,
Christenh., comb. nov. Basionym: Cotoneaster Saunder’s Ref. Bot. 1: 50. 1869.
taofuensis J.Fryer & B.Hylmö, Cotoneasters Pyrus tiantangensis (X.M.Liu & C.L.Wang) M.F.Fay
Compreh. Guide : 317. 2009. & Christenh., comb. nov. Basionym: Sorbus
Pyrus tapashana (C.K.Schneid.) M.F.Fay & Christenh., tiantangensis X.M.Liu & C.L.Wang, Bull. Bot. Res.,
comb. nov. Basionym: Sorbus tapashana Harbin 29: 131. 2009.
C.K.Schneid., Bull. Herb. Boissier Ser. 2, 6: 313. 1906. Pyrus tibetica (G.Klotz) M.F.Fay & Christenh., comb.
Pyrus taurica (Zinserl.) M.F.Fay & Christenh., comb. nov. Basionym: Cotoneaster tibeticus G.Klotz, Wiss.
nov. Basionym: Sorbus taurica Zinserl. in Komarov, Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss.
Fl. USSR 9: 497. 1939. Reihe 17: 334. 1968, not: Pyrus thibetica Cardot
Pyrus tauricola (Zaik. ex Sennikov) M.F.Fay & Pyrus tiliifolia (H.Zare, Amini & Assadi) M.F.Fay &
Christenh., comb. nov. Basionym: Sorbus tauricola Christenh., comb. nov. Basionym: Sorbus tiliifolia
Zaik. ex Sennikov, Willdenowia 43: 39. 2013. H.Zare, Amini & Assadi, Iran. J. Bot. 10: 148. 2004.
Pyrus taylorii (T.T.Yu) M.F.Fay & Christenh., comb. nov. Pyrus tjuliniae (Pojark. ex Peshkova) M.F.Fay &
Basionym: Cotoneaster taylorii T.T.Yu, Bull. Brit. Mus. Christenh., comb. nov. Basionym: Cotoneaster
(Nat. Hist.), Bot. 1: 129. 1954. tjuliniae Pojark. ex Peshkova, Novosti Sist. Vyssh.
Pyrus teijiashanensis (J.Fryer & B.Hylmö) M.F.Fay Rast. 15: 232. 1979.
& Christenh., comb. nov. Basionym: Cotoneaster Pyrus tobanii (C.Németh) M.F.Fay & Christenh.,
teijiashanensis J.Fryer & B.Hylmö, Cotoneasters comb. nov. Basionym: Sorbus tobanii C.Németh, Fl.
Compreh. Guide : 317. 2009. Pannonica 5: 177. 2007, as ‘tobani’.
Pyrus tengyuehensis (W.W.Sm.) M.F.Fay & Christenh., Pyrus tomentella (Pojark.) M.F.Fay & Christenh., comb.
comb. nov. Basionym: Eriobotrya tengyuehensis nov. Basionym: Cotoneaster tomentellus Pojark.,
W.W.Sm., Notes Roy. Bot. Gard. Edinburgh 10: 30. Not. Syst. Inst. Bot. Acad. Sci. USSR 21: 200. 1961.
1917. Pyrus transcaucasica (Pojark.) M.F.Fay & Christenh.,
Pyrus tenuipes (Rehder & E.H.Wilson) M.F.Fay & comb. nov. Basionym: Cotoneaster transcaucasicus
Christenh., comb. nov. Basionym: Cotoneaster Pojark., Bot. Mater. Gerb. Bot. Inst. Komarova Akad.
tenuipes Rehder & E.H.Wilson, Pl. Wilson. (Sargent) Nauk S.S.S.R. 21: 162. 1961.
1: 171. 1912. Pyrus transens (G.Klotz) M.F.Fay & Christenh., comb.
Pyrus tenuis (McAll.) M.F.Fay & Christenh., comb. nov. nov. Basionym: Cotoneaster transens G.Kloz, Wiss.

123
Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Christenh., comb. nov. Basionym: Sorbus ulmifolia

Reihe 17: 337. 1968. Kárpáti ex Németh, Acta Bot. Hung. 52: 394. 2010.
Pyrus tripyrena (J.Fryer & B.Hylmö) M.F.Fay & Pyrus umbellifera M.F.Fay & Christenh., nom. nov.
Christenh., comb. nov. Basionym: Cotoneaster Replaced synonym: Crataegus umbellata Desf., Cat.
tripyrenus J.Fryer & B.Hylmö, Cotoneasters Compreh. Pl. Hort. Reg. Paris.: 408. 1829, not: Pyrus umbellata
Guide : 317. 2009. Dum.Cours. Etymology: This species has umbellate
Pyrus tristaniicarpa (Kurz) M.F.Fay & Christenh., comb. inflorescences.
nov. Basionym: Dichotomanthes tristaniicarpa Kurz, Pyrus umbellularia M.F.Fay & Christenh., nom. nov.
J. Bot. 11: 195. 1873, as ‘tristaniaecarpa’. Replaced synonym: Laurus umbellata Thunb., Syst.
Pyrus tsaii (Rehder) M.F.Fay & Christenh., comb. nov. Veg. (ed. 14): 384. 1784, not: Pyrus umbellata
Basionym: Photinia tsaii Rehder, J. Arnold Arbor. 19: Dum.Cours. Etymology: This species has umbellate
274. 1938. inflorescences and like Umbellularia resembles a
Lauraceae. (Figure 13E)
Pyrus tsarongensis (J.Fryer & B.Hylmö) M.F.Fay &
Christenh., comb. nov. Basionym: Cotoneaster Pyrus undulata (J.Fryer & B.Hylmö) M.F.Fay &
tsarongensis J.Fryer & B.Hylmö, Cotoneasters Christenh., comb. nov. Basionym: Cotoneaster
Compreh. Guide : 317. 2009. undulatus J.Fryer & B.Hylmö, Cotoneasters Compreh.
Guide : 317. 2009.
Pyrus tsinlingensis (C.L.Tang) M.F.Fay & Christenh.,
comb. nov. Basionym: Sorbus tsinlingensis C.L.Tang, Pyrus uniflora (Bunge) M.F.Fay & Christenh., comb.
Fl. Tsinling. 1: 608. 1974. nov. Basionym: Cotoneaster uniflorus Bunge in
Ledeb., Fl. Altaica 2: 220. 1830.
Pyrus tumetica (Pojark.) M.F.Fay & Christenh., comb.
nov. Basionym: Cotoneaster tumeticus Pojark., Not. Pyrus uralensis (B.Hylmö & J.Fryer) M.F.Fay &
Syst. Inst. Bot. Acad. Sci. USSR 21: 204. 1961. Christenh., comb. nov. Basionym: Cotoneaster
uralensis B.Hylmö & J.Fryer, Acta Bot. Fennica 162:
Pyrus turbinata (Craib) M.F.Fay & Christenh., comb.
181. 1999.
nov. Basionym: Cotoneaster turbinatus Craib,
Curtis’s Bot. Mag. 140: t. 8546. 1914. Pyrus uranus M.F.Fay & Christenh., nom. nov. Replaced
synonym: Photinia serrulata Lindl. var. prunifolia
Pyrus turcica (Zinserl.) M.F.Fay & Christenh., comb.
Hook. & Arn., Bot. Beechey Voy. 4: 185. 1833, not:
nov. Basionym: Sorbus turcica Zinserl., Fl. URSS 9:
Pyrus prunifolia Willd. Etymology: Named for the
399. 1939.
planet Uranus.
Pyrus turdus M.F.Fay & Christenh., nom. nov. Replaced
Pyrus uva-ursi (Lindl.) M.F.Fay & Christenh., comb.
synonym: Cotoneaster mairei H.Lév., Bull. Geogr. Bot.
nov. Basionym: Cotoneaster microphyllus Wall. ex
25: 45. 1915, not: Pyrus mairei H.Lév. Etymology:
Lindl. var. uva-ursi Lindl., Edwards’s Bot. Reg. 14: t.
Named for thrushes (genus Turdus), birds that are
1187. 1828.
keen on its fruit in autumn.
Pyrus uzbezica (Grevtsova ex J.Fryer & B.Hylmö)
Pyrus turkmenorum (Juz. & Popov) M.F.Fay
M.F.Fay & Christenh., comb. nov. Basionym:
& Christenh., comb. nov. Basionym: Malus
Cotoneaster uzbezicus Grevtsova ex J.Fryer &
turkmenorum Juz. & Popov, Fl. URSS 9: 364. 1939.
B.Hylmö, Cotoneasters Compreh. Guide : 78. 2009.
Pyrus tushanensis (T.T.Yu) M.F.Fay & Christenh., comb.
Pyrus vaccinioides M.F.Fay & Christenh., nom. nov.
nov. Basionym: Photinia tushanensis T.T.Yu, Acta
Replaced synonym: Vaccinium corymbiferum
Phytotax. Sin. 8: 229. 1963.
Miq., Fl. Ned. Ind., Eerste Bijv. 1: 588. 1861, not:
Pyrus tytthocarpa (Pojark.) M.F.Fay & Christenh., Pyrus corymbifera Nakai. Etymology: This species
comb. nov. Basionym: Cotoneaster tytthocarpus somewhat resembles a Vaccinium (Ericaceae).
Pojark., Not. Syst. Inst. Bot. Acad. Sci. USSR 21: 181.
Pyrus vajdae (Boros) M.F.Fay & Christenh., comb. nov.
1961.
Basionym: Sorbus vajdae Boros, Agrártud. Egyet.
Pyrus udvardyana (Somlyay & Sennikov) M.F.Fay Kert-Szölögazdaságtud. Karának Közlem. 13: 154.
& Christenh., comb. nov. Basionym: Sorbus 1949.
udvardyana Somlyay & Sennikov, Phytotaxa 164:
Pyrus vallerubusensis (C.Németh) M.F.Fay &
268. 2014.
Christenh., comb. nov. Basionym: Sorbus
Pyrus ujhelyii (Somlyay & Sennikov) M.F.Fay & vallerubusensis C.Németh, Kitaibelia 14: 93. 2009.
Christenh., comb. nov. Basionym: Sorbus ujhelyii
Pyrus vallusensis (C.Németh) M.F.Fay & Christenh.,
Somlyay & Sennikov, Ann. Bot. Fenn. 53: 363. 2016.
comb. nov. Basionym: Sorbus vallusensis C.Németh,
Pyrus ulleungensis (Chin S.Chang) M.F.Fay & Christenh., Stud. Bot. Hung. 47: 310. 2016.
comb. nov. Basionym: Sorbus ulleungensis Chin
Pyrus vandelaarii (J.Fryer & B.Hylmö) M.F.Fay &
S.Chang, Harvard Pap. Bot. 19: 252. 2014.
Pyrus ulmifolia (Kárpáti ex Németh) M.F.Fay &

vandelaarii J.Fryer & B.Hylmö, New Plantsman 8: Barabits, Tilia 13: 17. 2007.
238. 2001. Pyrus vexans (E.F.Warb.) M.F.Fay & Christenh., comb.
Pyrus variabilis M.F.Fay & Christenh., nom. nov. nov. Basionym: Sorbus vexans E.F.Warb., Watsonia
Replaced synonym: Cotoneaster tengyuehensis 4: 46. 1957.
J.Fryer & B.Hylmö, Watsonia 21: 338. 1997, not: Pyrus viburnoides M.F.Fay & Christenh., nom.
Pyrus tengyuehensis (W.W.Sm.) M.F.Fay & Christenh. nov. Replaced synonym: Cotoneaster glabratus
Etymology: This species is variable, hence the name. Rehder & E.H.Wilson, Pl. Wilson (Sargent) 1: 171.
Pyrus ×veitchii (Trabut) M.F.Fay & Christenh., comb. 1912, not: Pyrus glabrata (Rehder) L.H.Bailey.
nov. Basionym: Cydonia ×veitchii Trab., J. Heredity 7: Etymology: named for the resemblance to Viburnum
416. 1916. (note: Pyrus cydonia × Pyrus communis) (Adoxaceae)
Pyrus velebilii M.F.Fay & Christenh., nom. nov. Pyrus ×vilmoriniana (C.Weber) M.F.Fay & Christenh.,
Replaced synonym: Sorbus omissa Velebil, Preslia comb. nov. Basionym: Chaenomeles ×vilmoriniana
84: 377. 2012, not: Pyrus omissa (J.Fryer & B.Hylmö) C.Weber, Arnoldia 23: 64. 1963. (note: Pyrus
M.F.Fay & Christenh. Etymology: This species is speciosa × Pyrus cathayensis).
named for Jiří Velebil, who first described this Pyrus villosa (Thunb.) M.F.Fay & Christenh., comb.
species. nov. Basionym: Crataegus villosa Thunb., Crataegus
Pyrus velebitica (Kárpáti) M.F.Fay & Christenh., comb. villosa Thunb., Syst. Veg. (ed. 14): 465. 1784. (Figure
nov. Basionym: Sorbus velebitica Kárpáti in Németh, 13F)
Acta Bot. Hung. 52: 395. 2010. Pyrus villosula (Rehder & E.H.Wilson) M.F.Fay &
Pyrus velutina (Albov) M.F.Fay & Christenh., comb. Christenh., comb. nov. Basionym: Cotoneaster
nov. Basionym: Sorbus aria (L.) Crantz var. velutina acutifolius Turcz. var. villosulus Rehder & E.H.Wilson,
Albov, Prodr. Fl. Colchic.: 71. 1895. Pl. Wilson. (Sargent) 1: 158. 1912.
Pyrus venus M.F.Fay & Christenh., nom. nov. Replaced Pyrus vilmorinii (C.K.Schneid.) M.F.Fay & Christenh.,
synonym: Micromeles khasiana Decne., Nouv. Arch. comb. nov. Basionym: Sorbus vilmorinii C.K.Schneid.,
Mus. Hist. Nat. 10: 169. 1874, not: Pyrus khasiana Bull. Herb. Boissier, sér. 2, 6: 317. 1906.
Hook.f. Etymology: Named for the planet Venus. Pyrus vitis-idaea M.F.Fay & Christenh., nom. nov.
Pyrus vermilliona M.F.Fay & Christenh., nom. nov. Basionym: Cotoneaster browiczii J.Fryer & B.Hylmö,
Replaced synonym: Cotoneaster cinnabarinus Juz., Cotoneasters Compreh. Guide: 308. 2009, not:
Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk Pyrus browiczii Mulk. Etymology: Named for the
S.S.S.R. 13: 32. 1950, not: Pyrus cinnabarina Poit. superficial similarity to lingon, Vaccinium vitis-idaea
& Turpin. Etymology: Named for the vermillion L. (Ericaceae).
coloured fruits; vermillion is a pigment originally Pyrus vivax M.F.Fay & Christenh., nom. nov. Replaced
made from cinnabar. synonym: Cotoneaster cuspidatus C.Marquand ex
Pyrus vernae (C.K.Schneid.) M.F.Fay & Christenh., J.Fryer & B.Hylmö, New Plantsman 5: 138. 1998,
comb. nov. Basionym: Cotoneaster vernae not: Pyrus cuspidata Bertol. Etymology: From Latin,
C.K.Schneid., Bot. Gaz. 64: 71. 1917. vivacis, alive.
Pyrus vertesica M.F.Fay & Christenh., nom. nov. Pyrus wallichiana (G.Klotz) M.F.Fay & Christenh.,
Replaced synonym: Sorbus acutiserrata C.Németh, comb. nov. Basionym: Cotoneaster wallichianus
Kitaibelia 14: 90. 2009, as ‘acutiserratus’, not: Pyrus G.Klotz, Wiss. Z. Martin-Luther-Univ. Halle-
acutiserrata Gladkova. Etymology: Named for the Wittenberg, Math.-Naturwiss. Reihe 15: 541. 1966.
Vertes Mountains in Hungary where this species is Pyrus wanbooyensis (J.Fryer & B.Hylmö) M.F.Fay
found. & Christenh., comb. nov. Basionym: Cotoneaster
Pyrus verokotschyi (J.Fryer & B.Hylmö) M.F.Fay & wanbooyensis J.Fryer & B.Hylmö, Cotoneasters
Christenh., comb. nov. Basionym: Cotoneaster Compreh. Guide : 318. 2009.
verokotschyi J.Fryer & B.Hylmö, Cotoneasters Pyrus wardii (W.W.Sm.) M.F.Fay & Christenh., comb.
Compreh. Guide : 318. 2009. nov. Basionym: Cotoneaster wardii W.W.Sm., Not.
Pyrus verruculosa (Diels) M.F.Fay & Christenh., comb. Roy. Bot. Gard. Edinburgh 10: 25. 1917.
nov. Basionym: Cotoneaster verruculosus Diels, Not. Pyrus washanensis (J.Fryer & B.Hylmö) M.F.Fay &
Bot. Gard. Edinburgh 5: 272. 1912. Christenh., comb. nov. Basionym: Cotoneaster
Pyrus vertesensis (Boros) M.F.Fay & Christenh., comb. washanensis J.Fryer & B.Hylmö, Cotoneasters
nov. Basionym: Sorbus vertesensis Boros, Mitt. Kgl. Compreh. Guide : 318. 2009.
Ungar. Gartenb.-Lehranst. 3: 52. 1937. Pyrus ×watereri (Exell) M.F.Fay & Christenh., comb.
Pyrus veszpremensis (Barabits) M.F.Fay & Christenh., nov. Basionym: Cotoneaster ×watereri Exell, Gard.
comb. nov. Basionym: Sorbus veszpremensis

125
Chron. 3: 83. 1928. M.Mizush. ex T.Shimizu, Fl. Nagano Pref.: 1508. 1997.
Pyrus weberbaueri (C.K.Schneid.) M.F.Fay & Christenh., (note: Pyrus sambucifolia × Pyrus matsumurana).
comb. nov. Basionym: Hesperomeles weberbaueri Pyrus yuana (Spongberg) M.F.Fay & Christenh., comb.
C.K.Schneid., Bot. Jahrb. Syst. 42: 85. 1908. nov. Basionym: Sorbus yuana Spongberg, J. Arnold
Pyrus whiteana (T.C.G.Rich & L.Houston) M.F.Fay & Arbor. 67: 257. 1986.
Christenh., comb. nov. Basionym: Sorbus whiteana Pyrus yui (J.Fryer & B.Hylmö) M.F.Fay & Christenh.,
T.C.G.Rich & L.Houston, Watsonia 26: 2. 2006. comb. nov. Basionym: Cotoneaster yui J.Fryer &
Pyrus wilfordii (Koehne) M.F.Fay & Christenh., comb. B.Hylmö, New Plantsman 8: 237. 2001.
nov. Basionym: Sorbus wilfordii Koehne, Repert. Pyrus yulingkongensis (J.Fryer & B.Hylmö) M.F.Fay
Spec. Nov. Regni Veg. 10: 503. 1912. & Christenh., comb. nov. Basionym: Cotoneaster
Pyrus williamtelliana M.F.Fay & Christenh., nom. nov. yulingkongensis J.Fryer & B.Hylmö, Cotoneasters
Replaced synonym: Eriobotrya fragrans Champ. ex Compreh. Guide : 319. 2009.
Benth., Hooker’s J. Bot. Kew Gard. Misc. 4: 80. 1852, Pyrus zabelii (C.K.Schneid.) M.F.Fay & Christenh.,
not: Pyrus fragrans E.T.Geddes. Etymology: Named comb. nov. Basionym: Cotoneaster zabelii
for William Tell, a folk hero of Switzerland, who shot C.K.Schneid., Illustr. Handb. Laubholzk. 1: 749. 1906.
an apple off the head of his son with a crossbow and Pyrus zayulensis (G.Klotz) M.F.Fay & Christenh., comb.
saved both their lives. nov. Basionym: Cotoneaster zayulensis G.Klotz, Wiss.
Pyrus wilmottiana (E.F.Warb.) M.F.Fay & Christenh., Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss.
comb. nov. Basionym: Sorbus wilmottiana E.F. Reihe 17: 334. 1968.
Warb., Watsonia 6: 296. 1967. Pyrus zayunesis (T.T.Yu & L.T.Lu) M.F.Fay & Christenh.,
Pyrus wilsonii (Nakai) M.F.Fay & Christenh., comb. comb. nov. Basionym: Sorbus zayunesis T.T.Yu &
nov. Basionym: Cotoneaster wilsonii Nakai, Bot. L.T.Lu, Acta Phytotax. Sin. 18: 494. 1980.
Mag. (Tokyo) 32: 104. 1918. Pyrus zeravschanica (Pojark.) M.F.Fay & Christenh.,
Pyrus woronowii (Zinserl.) M.F.Fay & Christenh., comb. comb. nov. Basionym: Cotoneaster zeravschanicus
nov. Basionym: Sorbus woronowii Zinserl., Fl. URSS Pojark., Not. Syst. Inst. Bot. Acad. Sci. USSR 17: 208.
9: 395. 1939. Replaced synonym: Sorbus caucasica 1955.
Zinserl., Bot. Mater. Gerb. Glavn. Bot. Sada R.S.F.S.R. Pyrus zertovae (Kárpáti) M.F.Fay & Christenh., comb.
4: 142. 1923, not: Pyrus caucasica Fed. nov. Basionym: Sorbus zertovae Kárpáti, Bot.
Pyrus wrightiana (Maxim.) M.F.Fay & Christenh., Közlem. 52: 137. 1966, as ’žertovae’.
comb. nov. Basionym: Photinia wrightiana Maxim., Pyrus zeto M.F.Fay & Christenh., nom. nov. Replaced
Bull. Acad. Imp. Sci. St-Pétersbourg 32: 486. 1888. synonym: Photinia microcarpa Standl., Publ.
Pyrus wuzhishanensis (W.B.Liao, R.H.Miao & Q.Fan) Carnegie Inst. Wash. 461: 57. 1935, not: Pyrus
M.F.Fay & Christenh., comb. nov. Basionym: microcarpa (Pursh) DC. Etymology: From Greek
Rhaphiolepis wuzhishanensis W.B.Liao, R.H.Miao & Ζήτω, meaning ‘viva!’, wishing a long life.
Q.Fan, Novon 17: 429. 2007. Pyrus zhejiangensis (P.L.Chiu) M.F.Fay & Christenh.,
Pyrus xizangensis M.F.Fay & Christenh., nom. nov. comb. nov. Basionym: Photinia zhejiangensis
Replaced synonym: Chaenomeles thibetica T.T.Yu, P.L.Chiu, Acta Phytotax. Sin. 18: 97. 1980.
Acta Phytotax. Sin. 8: 234. 1963, not: Pyrus Pyrus zhijiangensis (T.C.Ku) M.F.Fay & Christenh.,
thibetica Cardot. Etymology: Named for Xizang, an comb. nov. Basionym: Photinia zhijiangensis T.C.Ku,
autonomous region in China known as Tibet. Acta Phytotax. Sin. 31: 192. 1993.
Pyrus yakutica (J.Fryer & B.Hylmö) M.F.Fay & Pyrus zolyomii (Soó) M.F.Fay & Christenh., comb. nov.
Christenh., comb. nov. Basionym: Cotoneaster Basionym: Sorbus aria (L.) Crantz zolyomii Soó, Tisia
yakuticus J.Fryer & B.Hylmö, Cotoneasters Compreh. 2: 220. 1937.
Guide : 318. 2009. Pyrus zuzanae (Bernátová & Májovský) M.F.Fay &
Pyrus yalungensis (J.Fryer & B.Hylmö) M.F.Fay & Christenh., comb. nov. Basionym: Sorbus zuzanae
Christenh., comb. nov. Basionym: Cotoneaster Bernátová & Májovský, Biologia (Bratislava) 58: 784.
yalungensis J.Fryer & B.Hylmö, Cotoneasters 2003.
Pyrus yinchangensis (J.Fryer & B.Hylmö) M.F.Fay & 149. Cannabaceae. Reference: Yang et al. (2013).
Note: Trema Lour. is paraphyletic with respect to
yinchangensis J.Fryer & B.Hylmö, Cotoneasters
Parasponia Miq., which is hereby transferred.
Trema andersonii (Planch.) Byng & Christenh., comb.
Pyrus ×yokouchii (M.Mizush. ex T.Shimizu) M.F.Fay &
nov. Basionym: Parasponia andersonii Planch. in
Christenh., comb. nov. Basionym: Sorbus ×yokouchii

DC., Prodr. 17: 195. 1873. 161. 1868.

Trema aspera (Blume) Byng & Christenh., comb. nov. Psorospermum laurentii (De Wild.) Byng & Christenh.,
Basionym: Parasponia aspera Blume, Mus. Bot. 2: comb. nov. Basionym: Vismia laurentii De Wild.,
66. 1856. Ann. Mus. Congo Belge, Bot. sér. 5, 2: 311. 1908.
Trema eurhyncha (Miq.) Byng & Christenh., comb. nov. Psorospermum orientale (Engl.) Byng & Christenh.,
Basionym: Parasponia eurhyncha Miq., Fl. Ned. Ind., comb. nov. Basionym: Vismia orientalis Engl., Nat.
Eerste Bijv. 3: 411. 1861. Pflanzenfam., ed. 2. 21: 186. 1925.
Trema melastomatifolia (J.J.Sm.) Byng & Christenh., Psorospermum periclitatum Byng & Christenh., nom.
comb. nov. Basionym: Parasponia melastomatifolia nov. Replaced synonym: Vismia pauciflora Milne-
J.J.Sm., Nova Guinea 8: 891. 1912. Redh., Kew Bull. 8: 437. 1953, not: Psorospermum
Trema parviflora (Miq.) Byng & Christenh., comb. nov. pauciflorum Baker. Etymology: This species is listed
Basionym: Parasponia parviflora Miq., Pl. Jungh. 1: as Endangered by IUCN due to habitat loss in its
69. 1851. native Tanzania. The epithet is Latin, meaning
endangered.
Trema paucinervia (Merr. & L.M.Perry) Byng &
Christenh., comb. nov. Basionym: Parasponia Psorospermum torrei (Mendes) Byng & Christenh.,
paucinervia Merr. & L.M.Perry, J. Arnold Arbor. 20: comb. nov. Basionym: Vismia torrei Mendes, Bol.
324. 1939. Soc. Brot. sér. 2, 43: 335. 1969.
Trema rigida (Merr. & L.M.Perry) Byng & Christenh.,
comb. nov. Basionym: Parasponia rigida Merr. & 189. Putranjivaceae. Reference: Wurdack et al. (2004).
L.M.Perry, J. Arnold Arbor. 22: 254. 1941. Drypetes pleioneura (Radcl.-Sm.) Christenh. & Byng,
Trema rugosa (Blume) Byng & Christenh., comb. nov. comb. nov. Basionym: Sibangea pleioneura Radcl.-
Basionym: Parasponia rugosa Blume, Mus. Bot. 2: Sm., Kew Bull. 32: 481. 1978.
66. 1856.
Trema similis (Blume) Byng & Christenh., comb. nov. 200. Violaceae. Reference: Wahlert et al. (2014).
Basionym: Parasponia similis Blume, Mus. Bot. 2: 66. Note: Violaceae is a complicated family with several
1856. genera not monophyletic. Particularly problematic
Trema simulans (Merr. & L.M.Perry) Byng & Christenh., are Hybanthus Jacq. and Rinorea Aubl., that are
comb. nov. Basionym: Parasponia simulans Merr. & widely polyphyletic. To solve at least some of the
L.M.Perry, J. Arnold Arbor. 22: 255. 1941. issues the genus Corynostylis Mart. is expanded
to include Agatea A.Gray, Hekkingia H.E.Ballard
154. Myricaceae & Munzinger is included in Paypayrola Aubl. and
Leonia Ruiz & Pav. is expanded to emcompass the
Note: Myrica L. is treated in the traditional, broad sense
entire ‘Clade 3’ of Wahlert et al. (2014), including
here, including Morella Lour.
Amphirrhox Spreng., Gloeospermum Triana &
Myrica chevalieri (Verd. & Polhill) Christenh. & Byng, Planch., Mayanaea Lundell, Orthion Standl. &
comb. nov. Basionym: Morella chevalieri Parra-Os., Steyerm. and some species of Hybanthus and
Brittonia 52: 321. 2001. Rinorea. This does not solve all the issues, but is a
Myrica chimanimaniana (Verd. & Polhill) Christenh. step toward monophyletic genera in Violaceae.
& Byng, comb. nov. Basionym: Morella Corynostylis lecointei (Munzinger) Byng & Christenh.,
chimanimaniana Verdc. & Polhill, Fl. Zambes. 9: 259. comb. nov. Basionym: Agatea lecointei Munzinger,
2006. Bot. J. Linn. Soc. 137: 93. 2001.
Corynostylis longipedicellata (Baker f.) Byng &
186. Hypericaceae. References: Ruhfel et al. (2013), Christenh., comb. nov. Basionym: Agation
Sun et al. (2016). longipedicellatum Baker f., J. Linn. Soc., Bot. 45: 270.
Note: Psorospermum Spach is expanded to include all 1921. (Figure 14A)
African and Malagasy species of Vismieae except Corynostylis macrobotrys (K.Schum. & Lauterb.)
Vismia rubescens Oliv. which is associated with Byng & Christenh., comb. nov. Basionym: Agatea
Harungana Lam. and is sister to American Vismia macrobotrys K.Schum. & Lauterb., Fl. Schutzgeb.
Vand. Südsee: 453. 1900.
Harungana rubescens (Oliv.) Byng & Christenh., comb. Corynostylis pancheri (Brongn.) Byng & Christenh.,
nov. Basionym: Vismia rubescens Oliv., Fl. Trop. Afr. comb. nov. Basionym: Agation pancheri Brongn.,
1: 160. 1868. Bull. Soc. Bot. France 8: 80. 1861.
Psorospermum affine (Oliv.) Byng & Christenh., comb. Corynostylis rufotomentosa (Baker f.) Byng &
nov. Basionym: Vismia affinis Oliv., Fl. Trop. Afr. 1: Christenh., comb. nov. Basionym: Agation

127
rufotomentosum Baker f., J. Linn. Soc., Bot. 45: 269. Phytologia 43: 467. 1979.
1921. Leonia martii (Eichler) Byng & Christenh., comb. nov.
Corynostylis schlechteri (Melch.) Byng & Christenh., Basionym: Gloeospermum martii Eichler, Fl. Bras.
comb. nov. Basionym: Agatea schlechteri Melch. (Martius) 13: 389. 1871.
in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 21: 361. Leonia pauciflora (Hekking) Byng & Christenh.,
1925. comb. nov. Basionym: Gloeospermum pauciflorum
Corynostylis veillonii (Munzinger) Byng & Christenh., Hekking, Proc. Kon. Ned. Akad. Wetensch. C 89: 399.
comb. nov. Basionym: Agatea veillonii Munzinger, 1986.
Bot. J. Linn. Soc. 137: 91. 2001. Leonia pilosa (Melch.) Byng & Christenh., comb. nov.
Corynostylis violaris (A.Gray) Byng & Christenh., comb. Basionym: Gloeospermum pilosum Melch., Notizbl.
nov. Basionym: Agatea violaris A.Gray, Proc. Amer. Bot. Gart. Berlin-Dahlem 8: 623. 1923.
Acad. Arts & Sci. 2: 324. 1852. Leonia portobelensis (A.Robyns) Byng & Christenh.,
Leonia andina (Tul.) Byng & Christenh., comb. nov. comb. nov. Basionym: Gloeospermum portobelense
Basionym: Alsodeia andina Tul., Ann. Sci. Nat., Bot. A.Robyns, Ann. Missouri Bot. Gard. 53: 111. 1966.
sér. 3. 7: 366. 1847. Leonia sclerophylla (Cuatrec.) Byng & Christenh.,
Leonia blakeana (Standl.) Byng & Christenh., comb. comb. nov. Basionym: Gloeospermum sclerophyllum
nov. Basionym: Rinorea blakeana Standl., Publ. Field Cuatrec., Fieldiana, Bot. 27: 97. 1950.
Mus. Nat. Hist., Bot. Ser. 22: 349. 1940. Leonia sphaerocarpa (Triana & Planch.) Byng &
Leonia borealis (C.V.Morton) Byng & Christenh., comb. Christenh., comb. nov. Basionym: Gloeospermum
nov. Basionym: Gloeospermum boreale C.V.Morton, sphaerocarpum Triana & Planch., Ann. Sci. Nat., Bot.
Publ. Field Mus. Nat. Hist., Bot. Ser. 9: 309. 1940. sér. 4, 17: 129. 1862.
Leonia crassicarpa (W.Palacios) Byng & Christenh., Leonia sprucei (Eichler) Byng & Christenh., comb. nov.
comb. nov. Basionym: Gloeospermum crassicarpum Basionym: Gloeospermum sprucei Eichler, Fl. Bras.
W.Palacios, Caldasia 34: 77. 2012. (Martius) 13: 390. 1871.
Leonia caudata (Lundell) Byng & Christenh., comb. Leonia surinamensis (Eichler) Byng & Christenh.,
nov. Basionym: Orthion caudatum Lundell, Wrightia comb. nov. Basionym: Amphirrhox surinamensis
4: 38. 1968. Eichler, Fl. Bras. (Martius) 13: 377. 1871.
Leonia dichotoma (Rusby) Byng & Christenh., comb. Leonia ulei (Melch.) Byng & Christenh., comb. nov.
nov. Basionym: Rinorea dichotoma Rusby, Descr. S. Basionym: Gloeospermum ulei Melch., Notizbl. Bot.
Amer. Pl.: 61. 1920. Gart. Berlin-Dahlem 9: 164. 1924.
Leonia diversipetala (L.O.Williams) Byng & Paypayrola bordenavei (H.E.Ballard & Munzinger)
Christenh., comb. nov. Basionym: Gloeospermum Byng & Christenh., comb. nov. Basionym: Hekkingia
diversipetalum L.O.Williams, Ceiba 4: 273. 1955. bordenavei H.E.Ballard & Munzinger, Syst. Bot. 28:
Leonia eneidensis (Hekking) Byng & Christenh., comb. 345. 2003.
nov. Basionym: Gloeospermum eneidense Hekking,
Phytologia 43: 463. 1979. 202. Passifloraceae. References: Tokuoka (2012), Sun et
Leonia equatoriensis (Hekking) Byng & Christenh., al. (2016).
comb. nov. Basionym: Gloeospermum equatoriense Barteria laevigata (Sol. ex R.Br.) Byng & Christenh.,
Hekking, Phytologia 43: 464. 1979. comb. nov. Basionym: Smeathmannia laevigata Sol.
Leonia falcata (Hekking) Byng & Christenh., comb. ex R.Br., Trans. Linn. Soc. London 13: 221. 1821.
nov. Basionym: Gloeospermum falcatum Hekking, Barteria pubescens (Sol. ex R.Br.) Byng & Christenh.,
Phytologia 43: 465. 1979. comb. nov. Basionym: Smeathmannia pubescens
Leonia ferrugineosticta (A.Robyns) Byng & Sol. ex R.Br., Trans. Linn. Soc. London 13: 221. 1821.
Christenh., comb. nov. Basionym: Gloeospermum
ferrugineostictum A.Robyns, Ann. Missouri Bot. 207. Euphorbiaceae.
Gard. 53: 111. 1966.
Note: The following variety is best elevated to species
Leonia gossypium (Tul.) Byng & Christenh., comb. nov. level.
Basionym: Alsodeia gossypium Tul., Ann. Sci. Nat.,
Euphorbia pubescenticocca (Christenh.) Christenh.,
Bot. sér. 3, 7: 367. 1847.
comb. & stat. nov. Basionym: Euphorbia hypericifolia
Leonia grandifolia (Hekking) Byng & Christenh., L. var. pubescenticocca Christenh., Harvard Pap.
comb. nov. Basionym: Gloeospermum grandifolium
Hekking, Phytologia 43: 461. 1979.
Leonia longifolia (Hekking) Byng & Christenh., comb.
nov. Basionym: Gloeospermum longifolium Hekking,

Botany 7: 6. 2002. Singapore 12: 116. 1949, as ‘angustifolia’.

Syzygium barotsense (F.White) Byng & Christenh.,
208. Linaceae. Reference: McDill & Simpson (2011). comb. & stat. nov. Basionym: Syzygium guineense
subsp. barotsense F.White, Forest Fl. N. Rhodes.:
Note: Hugonia L. is not monophyletic and therefore
455. 1962.
species of Durandea Delarbre and Roucheria Planch.
are hereby transferred. Syzygium bleeseri (O.Schwarz) Byng & Christenh.,
comb. nov. Basionym: Eugenia bleeseri O.Schwarz,
Hugonia angustifolia (Stapf) Byng & Christenh., comb.
Repert. Spec. Nov. Regni Veg. 24: 90. 1927.
nov. Basionym: Durandea angustifolia Stapf, Bull.
Misc. Inform. Kew 1908: 11. 1908. Syzygium cadetii Byng & Christenh., nom. nov.
Replaced synonym: Syzygium cymosum var.
Hugonia viscosa (Stapf) Byng & Christenh., comb. nov.
montanum J.Guého & A.J.Scott, Kew Bull. 34: 490
Basionym: Durandea viscosa Stapf, Hooker’s Icon. Pl.
(1980). Etymology: Named after Thérésian Cadet
29: sub t. 2822. 1906.
who collected the type specimen of this species.
Hugonia griffithiana (Planch.) Byng & Christenh., (Figures 14C–D)
comb. nov. Basionym: Roucheria griffithiana Planch.,
Syzygium corneri Byng & Christenh., nom. nov.
London J. Bot. 6: 143. 1847.
Replaced synonym: Syzygium oblanceolatum subsp.
Hugonia magnifolia (Stapf) Byng & Christenh., comb. kinabaluense P.S.Ashton, Gard. Bull. Singapore 61:
nov. Basionym: Durandea magnifolia Stapf, Hooker’s 12. 2009, not: Syzygium kinabaluense (Stapf) Merr.
Icon. Pl. 29: t. 2822. 1906. & L.M.Perry. Etymology: Named for Edred J.H.
Hugonia vitiensis (Stapf) Byng & Christenh., comb. Corner, for his contributions to Malaysian botany.
nov. Basionym: Durandea vitiensis Stapf, Hooker’s Syzygium fraserense Byng & Christenh., nom. nov.
Icon. Pl. 29: sub t. 2822. 1906 Replaced synonym: Eugenia rugosa var. cordata
M.R.Hend., Gard. Bull. Singapore 12: 246. 1949, not:
218. Myrtaceae. References: Byng (2015), SYZWG Syzygium cordatum Hochst. Etymology: The type of
(2016), Byng et al. (in prep.). this species was collected at the mountain resort of
Note: Following extensive herbarium work, Fraser’s Hill in Pahang, an area with high biodiversity.
supplemented with molecular data, the following Syzygium garcinifolioides (M.R.Hend.) Byng &
taxa need elevating to species rank to provide a Christenh., comb. & stat. nov. Basionym: Eugenia
more consistent classification. burkilliana var. garcinifolioides M.R.Hend., Gard.
Eugenia drummondii Byng & Christenh., nom. nov. Bull. Singapore 12: 118. 1949.
Replaced synonym: Eugenia capensis subsp. Syzygium gununganum Byng & Christenh., nom. nov.
gracilipes F.White, Kirkia 10: 403. 1977, not: Eugenia Replaced synonym: Eugenia claviflora var. montana
gracilipes Kiaersk., nor: Eugenia gracilipes A.Gray. M.R. Hend., Gard. Bull. Singapore 12: 260. 1949, not:
Etymology: Named in honour of Robert Baily Syzygium montanum Thwaites & Hook.f. Etymology:
Drummond who collected the type specimen and From Malay gunung, meaning mountain.
was a former Keeper of the Herbarium at the Royal Syzygium holttumii (Ridl.) Byng & Christenh., comb.
Botanic Gardens, Kew. nov. Basionym: Eugenia holttumii Ridl., J. Bot. 62:
Eugenia verdcourtii Byng, nom. nov. Replaced 296. 1924.
synonym: Eugenia capensis subsp. multiflora Verdc., Syzygium jugorum (Craib) Byng & Christenh., comb.
Kew Bull. 54: 43. 1999, not: Eugenia multiflora nov. Basionym: Eugenia jugorum Craib, Bull. Misc.
Lam. Etymology: Named for Bernard Verdcourt, an Inform. Kew 1930: 168. 1930.
authority on East African plants. He first recognised Syzygium littorosum Byng, nom. nov. Replaced
and described this taxon. synonym: Syzygium guineense var. littorale Keay,
Syzygium afromontanum (F.White) Byng, comb. & Kew Bull. 8: 289, 1953, not: Syzygium littorale
stat. nov. Basionym: Syzygium guineense subsp. (Blume) Amshoff. Etymology: From Latin littus,
afromontana F.White, Forest Fl. N. Rhodes.: 455. shore, meaning shore-loving. (Figures 14E–F)
1962. (Figure 14B) Syzygium niassense Byng & J.E.Burrows sp. nov.
Syzygium altecastaneum (P.S.Ashton) Byng & (Figure 15). This is a riverine species with terete
Christenh., comb. & stat. nov. Basionym: Syzygium branchlets, and leaf apices that are usually acute.
castaneum subsp. altecastaneum P.S.Ashton, Kew The hypanthium is light green to magenta above,
Bull. 61: 116. 2006. sepals are magenta and fruits are oblong. It is similar
Syzygium angustifolium (M.R.Hend.) Byng & to Syzygium guineense (Willd.) DC. but differs in
Christenh., comb. & stat. nov. Basionym: Eugenia its longer petioles and oblong fruit that are white
kiahii var. angustifolium M.R.Hend., Gard. Bull. initially but turn pale pink when mature. Tree to
18 m; bark smooth, pale grey to whitish mottled;

129
A B
C D
Figure 14: Angiosperms (Eudicots). A, Corynostylis longipedicellata (Baker f.) Byng & Christenh.; B, Syzygium
afromontanum (F.White) Byng; C-D, habit and flowers of Syzygium cadetii Byng & Christenh.; E-F, Syzygium littorosum
Byng; and G, Syzygium pratense Byng. Images A, © Maarten Christenhusz; B-G, © James Byng.

A B
E F
Figure 15: Angiosperms (Eudicots). Syzygium niassense Byng & J.E.Burrows sp. nov. A, habit; B, buttressed stem;
C-D, inflorescences; E, white fruit; and F, maturing pale pink fruit. All images © John Burrows.

131
branchlets terete, greyish brown, peeling. Leaves Bull. 61: 123. 2006.
drying dark green to dark olive green above, lighter Syzygium paucinervium (Chantar. & J.Parn.) Byng &
below; coriaceous, 7.0‒13.0 × 2.5‒5.0 cm, elliptic, Christenh., comb. & stat. nov. Basionym: Syzygium
lanceolate-elliptic or oblong-elliptic, base cuneate, hemsleyanum subsp. paucinervium Chantar. &
apex acute to acuminate, margin slightly revolute; J.Parn., Kew Bull. 48: 599. 1993.
10‒18 secondary vein pairs, these are 5–7 mm Syzygium phaeophyllum (Merr. & L.M.Perry) Byng &
apart, prominent on both sides, tertiary veins many, Christenh., comb. & stat. nov. Basionym: Syzygium
prominent; inner intramarginal vein ca. 1 mm from leucoxylon var. phaeophyllum Merr. & L.M.Perry,
the leaf margin, outer intramarginal obscure, < 0.3 Mem. Amer. Acad. Arts, n.s., 18: 194. 1939.
mm from the leaf margin; petioles 10‒17 mm long.
Syzygium pratense Byng, nom. nov. Replaced synonym:
Inflorescences terminal, up to 7.5 cm long, axes
Syzygium guineense var. macrocarpum Engl. in
terete, flowers up to 30, in clusters of 3; bracts and
H.G.A.Engler & C.G.O.Drude, Veg. Erde 3(2): 738.
bracteoles deltoid, ca. 0.75 mm long, caducous.
1921, not: Syzygium macrocarpum Bahadur &
Flower buds ca. 6 × 3 mm; pseudostalks ca. 2 mm
R.C.Gaur. Etymology: Refers to the grasslands where
long; hypanthium 4.0–6.0 × 5.0–5.5 mm, pyriform,
this species often occurs. (Figure 14G)
light green below to pinkish at the top; sepals
0.5–1.0 × 2.0–3.0 mm, round, magenta; petals ca. Syzygium quadrisepalum (P.S.Ashton) Byng &
3–4 × 3–4 mm, orbicular, creamy white, calyptrate; Christenh., comb. & stat. nov. Basionym: Syzygium
stamens 5‒7 mm long, white; anthers ca. 0.5 mm; barringtonioides var. quadrisepalum P.S.Ashton, Kew
styles 6‒7 mm long. Fruits oblong, 2.0–2.5 × ca. Bull. 61: 113. 2006.
1.5 cm, white at first, ripening to pale pink. Type: Syzygium shimbaense (Verdc.) Byng & Christenh.,
MOZAMBIQUE, Sofala, Cheringoma Plateau, ca. 40 comb. & stat. nov. Basionym: Syzygium cordatum
km from the Dondo-Inhaminga road on the Chiniziua subsp. shimbaense Verdc., Fl. Trop. E. Afr., Myrtac.
road, next to the crossing over the Chiniziua River, 93 : 74. 2001.
m, fl., 8 October 2007, J.E. Burrows & S.M. Burrows Syzygium smetsianum Byng & Christenh., nom. nov.
10156 (holotype BNRH). Distribution: Known only Replaced synonym: Eugenia claviflora var. riparia
from Mozambique, but likely to occur in southern M.R.Hend., Gard. Bull. Singapore 12: 257. 1949,
Tanzania. Ecology: Riverbanks, riverine, coastal not: Syzygium riparium (Diels) Merr. & L.M.Perry.
and miombo woodlands; 30–404 m elevations. Etymology: Named in honour of Erik Smets, Director
Etymology: Named after the Niassa Game Reserve of Naturalis Biodiversity Center, for his contributions
where it occurs. Additional specimens examined: to botany.
MOZAMBIQUE. Cabo Delgado, Pemba to Mahate Syzygium sublaetum (Craib) Byng & Christenh., comb.
(Quissanga), sterile, 9 October 2005, J.E. Burrows nov. Basionym: Eugenia sublaeta Craib, Bull. Misc.
& S.M. Burrows 9159 (BNRH); ca. 10 km south Inform. Kew 1930: 169. 1930.
of Palma on road to Mocimboa de Praia, fl., 2
Syzygium tavaiense (P.S.Ashton) Byng & Christenh.,
December 2008, J.E. Burrows & S.M. Burrows 10923
comb. & stat. nov. Basionym: Syzygium claviflorum
(BNRH); Rio Mute, ca. 10 km S of Palma on road to
subsp. tavaiense P.S.Ashton, Gard. Bull. Singapore
Mocimboa da Praia, fr., 21 March 2009, J.E. Burrows
61: 8. 2009.
& S.M. Burrows 10923 (BNRH); Nangade district,
Camp of Hunters Mozambique concession, on Syzygium urophyllum (Welw. ex Hiern) Byng &
banks of Rovuma River, sterile, 8 September 2009, Christenh., comb. nov. Basionym: Eugenia urophylla
J.E. Burrows & S.M. Burrows 10923 (BNRH). Niassa, Welw. ex Hiern, Cat. Afr. Pl. 1: 360. 1898.
base of track up Serra Mecula from west side, Niassa
Game Reserve, fr., 16 March 2009, J.E. Burrows & 222. Penaeaceae. Reference: Schönenberger & Conti
S.M. Burrows 11220 (BNRH). Zambezia, between (2003).
Caia and Nicuadala, sterile, 2 October 2005, J.E. Note: Morphologically the genera of subfamily
Burrows & S.M. Burrows 9088 (BNRH). Penaeoideae are similar but have nevertheless been
Syzygium parnellii Byng & Christenh., nom. nov. divided into seven genera based on trivial characters.
Replaced synonym: Syzygium boisianum subsp. Molecular data did not support these genera. We
longifolium Chantar. & J.Parn., Kew Bull. 48: 593. thus expand the circumscription of Penaea L. to all
1993, not: Syzygium longifolium (Brongn. & Gris) members of Penaeoideae and elevate subspecies of
J.W.Dawson. Etymology: Named after John Parnell Penaea cneorum Meisn. to species level.
who studied this genus in Thailand and Indochina, Penaea geneiophora Byng & Christenh., nom. nov.
and who first recognised this as a new taxon. Replaced synonym: Stylapterus barbatus A.Juss.,
Syzygium patentinervium (P.S.Ashton) Byng & Ann. Sci. Nat., Bot. sér. 3, 6: 23, t. 1. 1846, not: Penaea
Christenh., comb. & stat. nov. Basionym: Syzygium barbata Endl. Etymology: From Greek γενειοφόρος,
garciniifolium var. patentinervium P.S.Ashton, Kew

meaning bearded. comb. nov. Basionym: Triceros cochinchinensis Lour.,

Penaea gigantea (R.Dahlgren) Byng & Christenh., Fl. Cochinch. 1: 184. 1790
comb. & stat. nov. Basionym: Penaea cneorum Staphylea formosana (Nakai) Byng & Christenh.,
subsp. gigantea R.Dahlgren, Opera Bot. 29: 40. 1971. comb. nov. Basionym: Turpinia formosana Nakai, J.
Penaea lanceolata (R.Dahlgren) Byng & Christenh., Arnold Arbor. 5: 80. 1924.
comb. & stat. nov. Basionym: Penaea cneorum Staphylea indochinensis (Merr.) Byng & Christenh.,
subsp. lanceolata R.Dahlgren, Opera Bot. 29: 50. comb. nov. Basionym: Turpinia indochinensis Merr.,
1971. J. Arnold Arbor. 19: 43. 1938.
Penaea micrantha (R.Dahlgren) Byng & Christenh., Staphylea insignis (Tul.) Byng & Christenh., comb. nov.
comb. nov. Basionym: Stylapterus micranthus Basionym: Turpinia insignis Tul., Ann. Sci. Nat., Bot.
R.Dahlgren, Opera Bot. 15: 18. 1967. sér. 3, 7: 296. 1847.
Penaea microphylla (Rourke) Byng & Christenh., comb. Staphylea macrosperma (C.C.Huang) Byng &
nov. Basionym: Brachysiphon microphyllus Rourke, Christenh., comb. nov. Basionym: Turpinia
Nordic J. Bot. 15: 63. 1995. macrosperma C.C.Huang, Fl. Yunnan. 2: 358. 1979.
Penaea mundii (Sond.) Byng & Christenh., comb. nov. Staphylea megaphylla (Tul.) Byng & Christenh., comb.
Basionym: Brachysiphon mundii Sond., Linnaea 23: nov. Basionym: Turpinia megaphylla Tul., Ann. Sci.
102. 1850. Nat., Bot. sér. 3, 6: 361. 1846.
Penaea petraea (W.F.Barker) Byng & Christenh., comb. Staphylea ovalifolia (Elmer) Byng & Christenh., comb.
nov. Basionym: Brachysiphon petraeus W.F.Barker, J. nov. Basionym: Turpinia ovalifolia Elmer, Leafl.
S. African Bot. 29: 167. 1963. Philipp. Bot. 2: 490. 1908.
Penaea retzioides (Sond.) Byng & Christenh., comb. Staphylea pomifera (Roxb.) Byng & Christenh., comb.
nov. Basionym: Endonema retzioides Sond., Linnaea nov. Basionym: Dalrympelea pomifera Roxb., Pl.
23: 103. 1850. Coromandel 3: 76. 1819.
Penaea roxburghii Byng & Christenh., nom. nov. Staphylea robusta (Craib) Byng & Christenh., comb.
Replaced synonym: Brachysiphon ericifolius A.Juss., nov. Basionym: Turpinia robusta Craib, Bull. Misc.
Ann. Sci. Nat., Bot. sér. 3, 6: 24. 1846, not: Penaea Inform. Kew 1926: 361. 1926.
ericifolia Gilg. Etymology: Named for J. Roxburgh, Staphylea simplicifolia (Merr.) Byng & Christenh.,
collector of the type specimen. comb. nov. Basionym: Turpinia simplicifolia Merr.,
Penaea rupestris (Sond.) Byng & Christenh., comb. Philipp. J. Sci. 27: 34. 1925.
nov. Basionym: Brachysiphon rupestris Sond., Staphylea subsessilifolia (C.Y.Wu) Byng & Christenh.,
Linnaea 23: 101. 1850. comb. nov. Basionym: Turpinia subsessilifolia C.Y.Wu,
Penaea ruscifolia (R.Dahlgren) Byng & Christenh., Fl. Yunnan. 2: 355. 1979, nom inval. = Turpinia
comb. & stat. nov. Basionym: Penaea cneorum subsessilifolia C.Y.Wu, Cat. Type Sp. Herb. China: 625.
subsp. ruscifolia R.Dahlgren, Opera Bot. 29: 48. 1994.
1971. Staphylea ternata (Nakai) Byng & Christenh., comb.
Penaea speciosa (W.F.Barker) Byng & Christenh., nov. Basionym: Turpinia ternata Nakai, J. Arnold
comb. nov. Basionym: Brachysiphon speciosus Arbor. 5: 78. 1924.
Sond., Linnaea 23: 103. 1850. Staphylea tricornuta (Lundell) Byng & Christenh.,
Penaea sulcata (R.Dahlgren) Byng & Christenh., comb. comb. nov. Basionym: Turpinia tricornuta Lundell,
nov. Basionym: Stylapterus sulcatus R.Dahlgren, Bull. Torrey Bot. Club 66: 598. 1939.
Opera Bot. 15: 26. 1967. Staphylea zanthoxyloides Byng & Christenh., nom.
nov. Replaced synonym: Zanthoxylum montanum
226. Staphyleaceae. Reference: Harris et al. (2017). Blume, not: Staphylea montana Roxb. ex Wight &
Arn., Bijdr. Fl. Ned. Ind. 5: 248. 1825. Etymology:
Note: Euscaphis Siebold & Zucc. and Turpinia Vent.
This species superficially resembles a Zanthoxylum
are embedded in Staphylea L. and the species are
(Rutaceae), hence the chosen name.
hereby transferred.
Staphylea affinis (Merr. & L.M.Perry) Byng &
Christenh., comb. nov. Basionym: Turpinia affinis 238. Burseraceae. References: Weeks et al. (2014),
Merr. & L.M.Perry, J. Arnold Arbor. 22: 550. 1941. Muellner-Riehl et al. (2016).
Staphylea arguta (Seem.) Byng & Christenh., comb. Note: Crepidospermum Hook.f. and Tetragastris Gaertn.
nov. Basionym: Turpinia arguta Seem., Bot. Voy. are embedded in Protium and any new combinations
Herald 10: 371. 1857. required are made here. African Dacryodes Vahl
do not form a clade with typical species from the
Staphylea cochinchinensis (Lour.) Byng & Christenh.,
Americas. Santiria Blume is also split with the sole

133
African species sister to African Dacryodes. No type (Rose) I.M.Johnst. Etymology: This species resembles
species was designated for Santiria, but Blume a species of Picramnia (Picramniaceae).
described it from Asia only and hence the genus Protium prancei (Daly.) Byng & Christenh., comb. nov.
should pertain to an Asian concept. We reinstate Basionym: Crepidospermum prancei Daly, Brittonia
Pachylobus G.Don for the African Dacryodes clade 39: 56. 1987.
and transfer the African Santiria species (for which Protium rhoifolium (Benth.) Byng & Christenh., comb.
combinations in Pachylobus already exist) into it. nov. Basionym: Hedwigia rhoifolia Benth., Hooker’s
Pachylobus bampsiana (Pierlot) Byng & Christenh., J. Bot. Kew Gard. Misc. 4: 17. 1852.
comb. nov. Basionym: Dacryodes bampsiana Pierlot, Protium surinamense Byng & Christenh., nom. nov.
Bull. Jard. Bot. Natl. Belg. 65: 366. 1996. Replaced synonym: Hedwigia hostmannii Engl. in
Pachylobus camerunensis (Onana) Byng & Christenh., A.DC. & C.DC., Monogr. Phan. 4: 97. 1883, not:
comb. nov. Basionym: Dacryodes camerunensis Protium hostmannii Engl. Etymology: This species is
Onana, Kew Bull. 61: 579. 2007. only known from Suriname.
Pachylobus ebatom (Aubrév. & Pellegr.) Byng & Protium varians (Little) Byng & Christenh., comb. nov.
Christenh., comb. nov. Basionym: Dacryodes Basionym: Tetragastris varians Little, J. Wash. Acad.
ebatom Aubrév. & Pellegr., Fl. Gabon 3: 89. 1962. Sci. 38: 96. 6. 1948.
Pachylobus igaganga (Aubrév. & Pellegr.) Byng &
Christenh., comb. nov. Basionym: Dacryodes 244. Cytinaceae
igaganga Aubrév. & Pellegr., Fl. Gabon 3: 86. 1962.
Note: We do not think the parasitic species found in
Pachylobus normandii (Aubrév. & Pellegr.) Byng Colombia in 2012 morphologically differs enough
& Christenh., comb. nov. Basionym: Dacryodes from the Central American Bdallophytum Eichler
normandii Aubrév. & Pellegr. Fl. Gabon 3: 72. 1962. to warrant generic status. The genus Sanguisuga
Pachylobus trapnellii (Onana) Byng & Christenh., Fern.Alonso & H.Cuadros, in which it was placed, is
comb. nov. Basionym: Dacryodes trapnellii Onana, therefore transferred.
Kew Bull. 58: 219. 2003. Bdallophytum caesareum (Fern.Alonso & H.Cuadros)
Pachylobus villiersiana (Onana) Byng & Christenh., Byng & Christenh., comb. nov. Basionym:
comb. nov. Basionym: Dacryodes villiersiana Onana, Sanguisuga caesarea Fern.Alonso & H.Cuadros,
Kew Bull. 63: 395. 2008. Caldasia 34: 293. 2012.
Protium atlanticum (Daly) Byng & Christenh., comb.
nov. Basionym: Crepidospermum atlanticum Daly, 247. Malvaceae. Reference: Whitlock & Hale (2011).
Kew Bull. 57: 471. 2002.
Note: Ayenia L. is embedded in Byttneria Loefl. and the
Protium breviacuminatum (Swart) Byng & Christenh., two genera are here merged with the small sister
comb. nov. Basionym: Tetragastris breviacuminata genus Rayleya Cristóbal. Ayenia has nomenclatural
Swart, Recueil Trav. Bot. Néerl. 39: 206. 1942. priority.
Protium cuneifolium (Cuatrec.) Byng & Christenh., Ayenia abutiloides (A.St.-Hil.) Christenh. & Byng,
comb. nov. Basionym: Hemicrepidospermum comb. nov. Basionym: Byttneria abutiloides A.St.-Hil.
cuneifolium Cuatrec., Webbia 12: 417. 1957. & Naudin, Ann. Sci. Nat., Bot. sér. 2, 18: 31. 1842.
Protium excelsior Byng & Christenh., nom. nov. Ayenia aculeata (Jacq.) Christenh. & Byng, comb. nov.
Replaced synonym: Icica altissima Aubl., Hist. Pl. Basionym: Chaetaea aculeata Jacq., Enum. Syst. Pl.:
Guiane 1: 342. 1775, not: Protium altissimum 17. 1760.
Marchand. Etymology: As this is a tall species, it
Ayenia affinis (Pohl) Christenh. & Byng, comb. nov.
‘reaches to the heavens’, hence ‘excelsior’.
Basionym: Byttneria affinis Pohl, Pl. Bras. Icon.
Protium goudotianum (Tul.) Byng & Christenh., comb. Descr. 2: 73. 1830.
nov. Basionym: Icica goudotiana Tul., Ann. Sci. Nat.,
Ayenia ambongensis (Arènes) Christenh. & Byng,
Bot. sér. 3, 6: 372. 1846.
comb. nov. Basionym: Byttneria ambongensis
Protium multijugum (Swart) Byng & Christenh., comb. Arènes, Mém. Inst. Sci. Madagascar, Sér. B, Biol. Vég.
nov. Basionym: Crepidospermum multijugum Swart, 7: 95. 1956.
Recueil Trav. Bot. Néerl. 39: 205. 1942.
Ayenia ancistrodonta (Mildbr.) Christenh. & Byng,
Protium osseum (Gaertn.) Byng & Christenh., comb. comb. nov. Basionym: Byttneria ancistrodonta
nov. Basionym: Tetragastris ossea Gaertn., Fruct. Mildbr, Notizbl. Bot. Gart. Berlin-Dahlem 11: 142.
Sem. Pl. 2: 130. 1791. 1931.
Protium picramnioides Byng & Christenh., nom. nov. Ayenia andamanensis (Kurz) Christenh. & Byng, comb.
Replaced synonym: Hedwigia panamensis Engl., Bot. nov. Basionym: Byttneria andamanensis Kurz, J.
Jahrb. Syst. 1: 42. 1880, not: Protium panamense

Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 40: 47. 1871. Sci. Nat., Bot. sér. 6, 11: 155. 1881, not: Ayenia
Ayenia aristeguietae (Cristóbal) Christenh. & Byng, cordifolia Sessé ex DC. Etymology: Named for the
comb. nov. Basionym: Byttneria aristeguietae genus Byttneria, now a synonym of Ayenia.
Cristóbal, Bonplandia (Corrientes) 4: 93. 1976. Ayenia caripensis (Cristóbal) Christenh. & Byng, comb.
Ayenia aspera (Colebr. ex Roxb.) Christenh. & Byng, nov. Basionym: Byttneria caripensis Cristóbal,
comb. nov. Basionym: Byttneria aspera Colebr. ex Bonplandia (Corrientes) 6: 125. 1989.
Roxb., Fl. Ind., ed. Carey & Wall. 2: 383. 1824. Ayenia catalpifolia (Jacq.) Christenh. & Byng, comb.
Ayenia asplundii (Cristóbal) Christenh. & Byng, comb. nov. Basionym: Byttneria catalpifolia Jacq., Pl. Rar.
nov. Basionym: Byttneria asplundii Cristóbal, Hort. Schoenbr. 1: 21. 1797.
Bonplandia (Corrientes) 4: 114. 1976. Ayenia celtoides (A.St.-Hil.) Christenh. & Byng, comb.
Ayenia asterotricha (Mildbr.) Christenh. & Byng, comb. nov. Basionym: Byttneria celtoides A.St.-Hil., Fl. Bras.
nov. Basionym: Byttneria asterotricha Mildbr., Merid. (quarto ed.) 1: 41. 1825.
Notizbl. Bot. Gart. Berlin-Dahlem 11: 143. 1931. Ayenia cordata (Lam.) Christenh. & Byng, comb. nov.
Ayenia arenesii Christenh. & Byng, nom. nov. Replaced Basionym: Byttneria cordata Lam., Encycl. 1(2): 523.
synonym: Byttneria hirta Arènes, Mém. Inst. Sci. 1785.
Madagascar, Sér. B, Biol. Vég. 7: 103. 1956, not: Ayenia coriacea (Britton) Christenh. & Byng, comb.
Ayenia hirta A.St.Hil. & Naudin. Etymology: Named nov. Basionym: Byttneria coriacea Britton, Bull.
for Jean Arènes, who first described this species. Torrey Bot. Club 16: 156. 1889.
Ayenia atrata (Bullock) Christenh. & Byng, comb. Ayenia corylifolia (Humb. & Bonpl. ex Roem. & Schult.)
nov. Basionym: Byttneria atrata Bullock, Bull. Misc. Christenh. & Byng, comb. nov. Basionym: Byttneria
Inform. Kew 1936: 1. 1936. corylifolia Humb. & Bonpl. ex Roem. & Schult., Syst.
Ayenia attenuatifolia (Cristóbal) Christenh. & Byng, Veg., ed. 15 bis 5: 470. 1819.
comb. nov. Basionym: Byttneria attenuatifolia Ayenia crenulata (Wall. ex Mast.) Christenh. & Byng,
Cristóbal, Bonplandia (Corrientes) 4: 389. 1976. comb. nov. Basionym: Byttneria crenulata Wall. ex
Ayenia aurantiaca (Mildbr.) Christenh. & Byng, comb. Mast., Fl. Brit. India 1(2): 376. 1874.
nov. Basionym: Byttneria aurantiaca Mildbr., Notizbl. Ayenia cristobaliana (Dorr) Christenh. & Byng, comb.
Bot. Gart. Berlin-Dahlem 11: 141. 1931. nov. Basionym: Byttneria cristobaliana Dorr, Kew
Ayenia australis (A.St.-Hil.) Christenh. & Byng, comb. Bull. 54: 991. 2000.
nov. Basionym: Byttneria australis A.St.-Hil., Fl. Bras. Ayenia crotonoides Christenh. & Byng, nom. nov.
Merid. (quarto ed.) 1: 145. 1825. Replaced synonym: Byttneria fruticosa K.Schum. ex
Ayenia bahiensis (Cristóbal) Christenh. & Byng, Engl., Abh. Königl. Akad. Wiss. Berlin 1894: 33. 1894,
comb. nov. Basionym: Rayleya bahiensis Cristóbal, not: Ayenia fruticosa Rose. Etymology: Named for its
Bonplandia (Corrientes) 5: 45. 1981. superficial resemblance to Croton (Euphorbiaceae).
Ayenia baronii (Arènes) Christenh. & Byng, comb. nov. Ayenia dahomensis (N.Hallé) Christenh. & Byng, comb.
Basionym: Byttneria baronii Arènes, Mém. Inst. Sci. nov. Basionym: Byttneria dahomensis N.Hallé,
Madagascar, Sér. B, Biol. Vég. 7: 86. 1956. Adansonia sér. 2, 2: 287. 1962.
Ayenia benensis (Britton) Christenh. & Byng, comb. Ayenia decaryana (Arènes) Christenh. & Byng, comb.
nov. Basionym: Byttneria benensis Britton, Bull. nov. Basionym: Byttneria decaryana Arènes, Mém.
Torrey Bot. Club 16: 155. 1889. Inst. Sci. Madagascar, Sér. B, Biol. Vég. 7: 94. 1956.
Ayenia bernieri (Hochr.) Christenh. & Byng, comb. Ayenia divaricata (Benth.) Christenh. & Byng, comb.
nov. Basionym: Byttneria bernieri Hochr., Annuaire nov. Basionym: Byttneria divaricata Benth., J. Bot.
Conserv. Jard. Bot. Genève 11 & 12: 4. 1907. (Hooker) 4: 124. 1841.
Ayenia besalampensis (Arènes) Christenh. & Byng, Ayenia echinata (Wall. ex Kurz) Christenh. & Byng,
comb. nov. Basionym: Byttneria besalampensis comb. nov. Basionym: Byttneria echinata Wall. ex
Arènes, Mém. Inst. Sci. Madagascar, Sér. B, Biol. Vég. Kurz, Forest Fl. Burma 1: 151. 1877.
7: 100. 1956 Ayenia elliptica (Pohl) Christenh. & Byng, comb. nov.
Ayenia beyrichiana (K.Schum.) Christenh. & Byng, Basionym: Byttneria elliptica Pohl, Pl. Bras. Icon.
comb. nov. Basionym: Byttneria beyrichiana Descr. 2: 71. 1830.
K.Schum., Fl. Bras. (Martius) 12: 96. 1886. Ayenia ellipticifolia (Arènes) Christenh. & Byng, comb.
Ayenia biloba (Baill.) Christenh. & Byng, comb. nov. nov. Basionym: Byttneria ellipticifolia Arènes, Mém.
Basionym: Byttneria biloba Baill., Adansonia 10: 178. Inst. Sci. Madagascar, Sér. B, Biol. Vég. 7: 101. 1956
1872. Ayenia engleri Christenh. & Byng, nom. nov. Replaced
Ayenia byttnerioides Christenh. & Byng, nom. nov. synonym: Byttneria glabra K.Schum. & Engl., Bot.
Replaced synonym: Byttneria cordifolia Sagot, Ann. Jahrb. Syst. 39: 592. 1907, not: Ayenia glabra

135
S.Watson. Etymology: Named for Heinrich G.A. comb. nov. Basionym: Byttneria implacabilis
Engler, who first described this species together with Cristóbal, Bonplandia (Corrientes) 4: 190. 1976.
Karl Schumann. Ayenia indica Christenh. & Byng, nom. nov. Replaced
Ayenia fernandesii (Cristóbal) Christenh. & Byng, synonym: Byttneria pilosa Roxb., Fl. Ind., ed. 1820, 2:
comb. nov. Basionym: Byttneria fernandesii 381. 1824, not: Ayenia pilosa Cristóbal. Etymology:
Cristóbal, Bonplandia (Corrientes) 4: 99. 1976. This species occurs in India, hence the name.
Ayenia filipes (Mart. ex K.Schum.) Christenh. & Byng, Ayenia integrifolia (Lace) Christenh. & Byng, comb.
comb. nov. Basionym: Byttneria filipes Mart. ex nov. Basionym: Byttneria integrifolia Lace, Bull.
K.Schum., Fl. Bras. (Martius) 12: 95. 1886. Misc. Inform. Kew : 396. 1915.
Ayenia flexuosa (Killip) Christenh. & Byng, comb. nov. Ayenia irwinii (Cristóbal) Christenh. & Byng, comb. nov.
Basionym: Byttneria flexuosa Killip, J. Wash. Acad. Basionym: Byttneria irwinii Cristóbal, Bonplandia
Sci. 24: 50. 1934. (Corrientes) 4: 285. 1976.
Ayenia fluvialis (Fryxell & Guadarr.) Christenh. & Byng, Ayenia ivorensis (N.Hallé) Christenh. & Byng, comb.
comb. nov. Basionym: Byttneria fluvialis Fryxell & nov. Basionym: Byttneria ivorensis N.Hallé,
Guadarr., Brittonia 53: 59. 2001. Adansonia sér. 2, 2: 287. 1962. Note: This species is
Ayenia fulva (Poepp.) Christenh. & Byng, comb. nov. extinct (Assi, 1998).
Basionym: Byttneria fulva Poepp., Nov. Gen. Sp. Pl. Ayenia jaculifolia (Pohl) Christenh. & Byng, comb. nov.
(Poeppig & Endlicher) 3: 73. 1845. Basionym: Byttneria jaculifolia Pohl, Pl. Bras. Icon.
Ayenia gayana (A.St.-Hil.) Christenh. & Byng, comb. Descr. 2: 77. 1830.
nov. Basionym: Byttneria gayana A.St.-Hil., Fl. Bras. Ayenia jaramilloana (Dorr) Christenh. & Byng,
Merid. (quarto ed.) 1: 145. 1825. comb. nov. Basionym: Byttneria jaramilloana Dorr,
Ayenia genistella (Triana & Planch.) Christenh. & Byng, BioLlania, Ed. Espec. 6: 305. 1997.
comb. nov. Basionym: Byttneria genistella Triana & Ayenia lancifolia (A.St.-Hil. & Naudin) Christenh. &
Planch., Ann. Sci. Nat., Bot. sér. 4, 17: 333. 1862. Byng, comb. nov. Basionym: Byttneria lancifolia
Ayenia glazioui (Hochr.) Christenh. & Byng, comb. A.St.-Hil. & Naudin, Ann. Sci. Nat., Bot. sér. 2, 18: 32.
nov. Basionym: Byttneria glazioui Hochr., Annuaire 1842.
Conserv. Jard. Bot. Genève 11–12: 7. 1907. Ayenia lappa Christenh. & Byng, nom. nov. Replaced
Ayenia gracilipes (Decne. ex Baill.) Christenh. & Byng, synonym: Byttneria ovata Lam., Encycl. 1(2): 522.
comb. nov. Basionym: Byttneria gracilipes Decne. ex 1785, not: Ayenia ovata Hemsl. Etymology: From
Baill., Adansonia 2: 167. 1861. Latin lappa, a bur.
Ayenia grandifolia (DC.) Christenh. & Byng, comb. nov. Ayenia lasiophylla (Cristóbal) Christenh. & Byng, comb.
Basionym: Byttneria grandiflora DC., Prodr. [A. P. de nov. Basionym: Byttneria lasiophylla Cristóbal,
Candolle] 1: 486. 1824. Bonplandia (Corrientes) 4: 299. 1976.
Ayenia guineensis (Keay & Milne-Redh.) Christenh. & Ayenia latipetala (Arènes) Christenh. & Byng, comb.
Byng, comb. nov. Basionym: Byttneria guineensis nov. Basionym: Byttneria latipetala Arènes, Mém.
Keay & Milne-Redh., Kew Bull. 9: 263. 1954. Inst. Sci. Madagascar, Sér. B, Biol. Vég. 7: 107. 1956.
Ayenia hatschbachii (Cristóbal) Christenh. & Byng, Ayenia lobata (Baill.) Christenh. & Byng, comb. nov.
comb. nov. Basionym: Byttneria hatschbachii Basionym: Byttneria lobata Baill., Bull. Mens. Soc.
Cristóbal, Bonplandia (Corrientes) 4: 307. 1976. Linn. Paris 1: 503. 1885.
Ayenia heteromorpha (Arènes) Christenh. & Byng, Ayenia lopezmirandae (Cristóbal) Christenh. & Byng,
comb. nov. Basionym: Byttneria heteromorpha comb. nov. Basionym: Byttneria lopezmirandae
Arènes, Mém. Inst. Sci. Madagascar, Sér. B, Biol. Vég. Cristóbal, Bonplandia (Corrientes) 4: 159. 1976, as
7: 85. 1956. ‘lopez-mirandae’.
Ayenia heterophylla (Hook.) Christenh. & Byng, comb. Ayenia loxensis (Cristóbal) Christenh. & Byng, comb.
nov. Basionym: Byttneria heterophylla Hook., Bot. nov. Basionym: Byttneria loxensis Cristóbal,
Misc. 1: 291. 1830. Bonplandia (Corrientes) 4: 102. 1976.
Ayenia hirsuta (Ruiz & Pav.) Christenh. & Byng, comb. Ayenia lucida (Arènes) Christenh. & Byng, comb. nov.
nov. Basionym: Byttneria hirsuta Ruiz & Pav., Fl. Basionym: Byttneria lucida Arènes, Mém. Inst. Sci.
Peruv. 3: 10. 1802. Madagascar, Sér. B, Biol. Vég. 7: 108. 1956.
Ayenia humbertiana (Arènes) Christenh. & Byng, Ayenia macrantha (Arènes) Christenh. & Byng, comb.
comb. nov. Basionym: Byttneria humbertiana nov. Basionym: Byttneria macrantha Arènes, Mém.
Arènes, Mém. Inst. Sci. Madagascar, Sér. B, Biol. Vég. Inst. Sci. Madagascar, Sér. B, Biol. Vég. 7: 89. 1956.
7: 95. 1956. Ayenia melantha (Mart. ex K.Schum.) Christenh. &
Ayenia implicabilis (Cristóbal) Christenh. & Byng, Byng, comb. nov. Basionym: Byttneria melantha

Mart. ex K.Schum., Fl. Bras. (Martius) 12(3): 98. Bonplandia (Corrientes) 4: 276. 1976.
1886. Ayenia pauxilla Christenh. & Byng, nom. nov. Replaced
Ayenia melastomifolia (A.St.-Hil.) Christenh. & Byng, synonym: Byttneria parviflora Benth., Pl. Hartw.
comb. nov. Basionym: Byttneria melastomifolia 114. 1843, not: Ayenia parviflora Sessé & Moc.
A.St.-Hil., Fl. Bras. Merid. (quarto ed.) 1(4): 144. Etymology: From Latin pauxillum, little, because it
1825, as ‘melastomaefolia’. has small flowers.
Ayenia melleri (Baker) Christenh. & Byng, comb. nov. Ayenia pedersenii (Cristóbal) Christenh. & Byng, comb.
Basionym: Byttneria melleri Baker, J. Linn. Soc., Bot. nov. Basionym: Byttneria pedersenii Cristóbal,
22: 451. 1886. Bonplandia (Corrientes) 4: 314. 1976.
Ayenia microphylla (Jacq.) Christenh. & Byng, comb. Ayenia perrieri (Hochr.) Christenh. & Byng, comb. nov.
nov. Basionym: Byttneria microphylla Jacq., Hort. Basionym: Byttneria perrieri Hochr., Candollea 3:
Bot. Vindob. 1: 10. 1770. 145. 1926.
Ayenia minytricha (Cristóbal) Christenh. & Byng, comb. Ayenia pescapraeifolia (Britton) Christenh. & Byng,
nov. Basionym: Byttneria minytricha Cristóbal, comb. nov. Basionym: Byttneria pescrapaeifolia
Bonplandia (Corrientes) 4: 204. 1976. Britton, Bull. Torrey Bot. Club 16: 155. 1889.
Ayenia morii (L.C.Barnett & Dorr) Christenh. & Byng, Ayenia petiolata (Cristóbal) Christenh. & Byng, comb.
comb. nov. Basionym: Byttneria morii L.C.Barnett & nov. Basionym: Byttneria petiolata Cristóbal,
Dorr, Brittonia 42: 271. 1990. Bonplandia (Corrientes) 4: 279. 1976.
Ayenia nitidula (Baker) Christenh. & Byng, comb. nov. Ayenia piresii (Cristóbal) Christenh. & Byng, comb. nov.
Basionym: Byttneria nitidula Baker, J. Linn. Soc., Bot. Basionym: Byttneria piresii Cristóbal, Bonplandia
25: 304. 1889. (Corrientes) 4: 380. 1976.
Ayenia nossibeensis (Arènes) Christenh. & Byng, comb. Ayenia pohlii Christenh. & Byng, nom. nov. Replaced
nov. Basionym: Byttneria nossibeensis Arènes, Mém. synonym: Byttneria dentata Pohl, Pl. Bras. Icon.
Inst. Sci. Madagascar, Sér. B, Biol. Vég. 7: 109. 1956. Descr. 2: 79. 1830, not: Ayenia dentata Brandegee.
Ayenia obcordata (Arènes) Christenh. & Byng, comb. Etymology: Named for Johann B.E. Pohl, who first
nov. Basionym: Byttneria obcordata Arènes, Mém. described this species.
Inst. Sci. Madagascar, Sér. B, Biol. Vég. 7: 109. 1956. Ayenia ramosissima (Pohl) Christenh. & Byng, comb.
Ayenia obliqua (Benth.) Christenh. & Byng, comb. nov. nov. Basionym: Byttneria ramosissima Pohl, Pl. Bras.
Basionym: Byttneria obliqua Benth., J. Bot. (Hooker) Icon. Descr. 2: 75. 1830.
4: 124. 1842. Ayenia rhamnifolia (Benth.) Christenh. & Byng, comb.
Ayenia oblongata (Pohl) Christenh. & Byng, comb. nov. Basionym: Byttneria rhamnifolia Benth.,
nov. Basionym: Byttneria oblongata Pohl, Pl. Bras. Hooker’s J. Bot. Kew Gard. Misc. 3: 164. 1851.
Icon. Descr. 2: 74. 1830. Ayenia rojasii (Cristóbal) Christenh. & Byng, comb. nov.
Ayenia oblongifolia (Arènes) Christenh. & Byng, comb. Basionym: Byttneria rojasii Cristóbal, Bonplandia
nov. Basionym: Byttneria oblongifolia Arènes, Mém. (Corrientes) 4: 296. 1976.
Inst. Sci. Madagascar, Sér. B, Biol. Vég. 7: 102. 1956. Ayenia rubriflora (Arènes) Christenh. & Byng, comb.
Ayenia obtusata (Benth.) Christenh. & Byng, comb. nov. Basionym: Byttneria rubriflora Arènes, Mém.
nov. Basionym: Byttneria obtustata Benth. ex Inst. Sci. Madagascar, Sér. B, Biol. Vég. 7: 107. 1956.
Hochr., Annuaire Conserv. Jard. Bot. Genève 11–12: Ayenia sagasteguii (Cristóbal) Christenh. & Byng,
5. 1907. comb. nov. Basionym: Byttneria sagasteguii
Ayenia oligantha (Arènes) Christenh. & Byng, comb. Cristóbal, Novon 13: 302. 2003.
nov. Basionym: Byttneria oligantha Arènes, Occas. Ayenia sagittifolia (A.St.-Hil.) Christenh. & Byng, comb.
Pap. Bernice Pauahi Bishop Mus. 14: 313. 1939. nov. Basionym: Byttneria sagittifolia A.St.-Hil., Fl.
Ayenia oranensis (Cristóbal) Christenh. & Byng, comb. Bras. Merid. (quarto ed.) 1(4): 142. 1825.
nov. Basionym: Byttneria oranensis Cristóbal, Ayenia sambiranensis (Arènes) Christenh. & Byng,
Bonplandia (Corrientes) 4: 365. 1976. comb. nov. Basionym: Byttneria sambiranensis
Ayenia ostenii (Cristóbal) Christenh. & Byng, comb. Arènes, Mém. Inst. Sci. Madagascar, Sér. B, Biol. Vég.
nov. Basionym: Byttneria ostenii Cristóbal, 7: 104. 1956.
Bonplandia (Corrientes) 4: 226. 1976. Ayenia scabra (L.) Christenh. & Byng, comb. nov.
Ayenia ovatifolia (Arènes) Christenh. & Byng, comb. Basionym: Byttneria scabra L., Syst. Nat., ed. 10, 2:
nov. Basionym: Byttneria ovatifolia Arènes, Mém. 939. 1759.
Inst. Sci. Madagascar, Sér. B, Biol. Vég. 7: 105. 1956. Ayenia scapellata (Pohl) Christenh. & Byng, comb. nov.
Ayenia palustris (Cristóbal) Christenh. & Byng, comb. Basionym: Byttneria scapellata Pohl, Pl. Bras. Icon.
nov. Basionym: Byttneria palustris Cristóbal,

137
Descr. 2: 78. 1830. we decided to unite them.

Ayenia schumannii (Cristóbal) Christenh. & Byng, Cochlospermum gonzalezii (Sprague & L.Riley) Byng
comb. nov. Basionym: Byttneria schumannii & Christenh., comb. nov. Basionym: Amoreuxia
Cristóbal, Bonplandia (Corrientes) 4: 208. 1976. gonzalezii Sprague & L.Riley, Bull. Misc. Inform. Kew
Ayenia schunkei (Cristóbal) Christenh. & Byng, comb. 1922: 102. 1922.
nov. Basionym: Byttneria schunkei Cristóbal, Bol. Cochlospermum malvifolium (A.Gray) Byng &
Soc. Argent. Bot. 24: 125. 1985. Christenh., comb. nov. Basionym: Amoreuxia
Ayenia sparrei (Cristóbal) Christenh. & Byng, comb. malvifolia A.Gray, Smithsonian Contr. Knowl. 3(5):
nov. Basionym: Byttneria sparrei Cristóbal, 29. 1852.
Bonplandia (Corrientes) 4: 146. 1976. Cochlospermum palmatifidum (Moc. & Sessé ex
Ayenia stenophylla (Cristóbal) Christenh. & Byng, DC.) Byng & Christenh., comb. nov. Basionym:
comb. nov. Basionym: Byttneria stenophylla Amoreuxia palmatifida Moc. & Sessé ex DC., Prodr.
Cristóbal, Bonplandia (Corrientes) 4: 261. 1976. 2: 638. 1825.
Ayenia subsessilis (Cristóbal) Christenh. & Byng, comb. Cochlospermum wrightii (A.Gray) Byng & Christenh.,
nov. Basionym: Byttneria subsessilis Cristóbal, comb. nov. Basionym: Amoreuxia wrightii A.Gray,
Bonplandia (Corrientes) 4: 271. 1976. Smithsonian Contr. Knowl. 5: 26. 1853.
Ayenia triadenia (Cristóbal) Christenh. & Byng, comb.
nov. Basionym: Byttneria triadenia Cristóbal, 251. Cistaceae. References: Civeyrel et al. (2011), Sun
Bonplandia (Corrientes) 4: 194. 1976. et al. (2016).
Ayenia tucumanensis (Cristóbal) Christenh. & Byng, Note: Neither Halimium Spach and Cistus L. form a
comb. nov. Basionym: Byttneria tucumanensis monophyletic group in numerous studies (e.g.
Cristóbal, Bonplandia (Corrientes) 4: 211. 1976. Civeyrel et al., 2011, Sun et al., 2016). Although,
Ayenia uaupensis (Spruce ex K.Schum.) Christenh. & traditionally treated separately based on flower
Byng, comb. nov. Basionym: Byttneria uaupensis colour and locule number the two genera share
Spruce ex K.Schum., Fl. Bras. (Martius) 12: 93. 1886. numerous morphological characters and even share
the same chromosome number (2n=18), which is
Ayenia urosepala (Mildbr.) Christenh. & Byng, comb.
different from all other genera in the family. They
nov. Basionym: Byttneria urosepala Mildbr., Notizbl.
also hybridise in the wild and in cultivation making
Bot. Gart. Berlin-Dahlem 11: 144. 1931.
delimitation even more difficult. We propose to
Ayenia urticifolia (K.Schum.) Christenh. & Byng, comb. place Halimium in Cistus to create a morphological
nov. Basionym: Byttneria urticifolia K.Schum., Fl. coherent and monophyletic group.
Bras. (Martius) 12: 98. 1886.
Cistus commutatus (Pau) Byng & Christenh., comb.
Ayenia vargasii (Cristóbal) Christenh. & Byng, comb. nov. Basionym: Halimium commutatum Pau, Bol.
nov. Basionym: Byttneria vargasii Cristóbal, Soc. Aragonesa Ci. Nat. 3: 263. 1904. (Figure 16A)
Bonplandia (Corrientes) 4: 151. 1976.
Cistus lasiocalycinus (Boiss. & Reut.) Byng & Christenh.,
Ayenia vitifolia (Baill.) Christenh. & Byng, comb. nov. comb. nov. Basionym: Helianthemum lasiocalycinum
Basionym: Byttneria vitifolia Baill., Adansonia 10: Boiss. & Reut., Diagn. Pl. Orient. ser. 2, 1: 50. 1854.
177. 1872.
Cistus macrocalycinus (Pau) Byng & Christenh., comb.
Ayenia voulily (Baill.) Christenh. & Byng, comb. nov. nov. Basionym: Helianthemum atriplicifolium Willd.
Basionym: Byttneria voulily Baill., Bull. Mens. Soc. var. macrocalycinum Pau, Iter Marocc. 1928: no. var.
Linn. Paris 1: 502. 1885. macrocalycinum Pau, Iter Marocc. 1928: no. 279.
Ayenia weberbaueri (Mildbr.) Christenh. & Byng, 1929.
comb. nov. Basionym: Byttneria weberbaueri
Mildbr., Notizbl. Bot. Gart. Berlin-Dahlem 11: 145.
268. Capparaceae. Reference: Tamboli (2018).
1931.
Note: The circumscription of the genus Capparis L. has
Commersonia rhamnoides (Seem.) Christenh. & Byng,
been in flux in recent times. Molecular data has
comb. nov. Basionym: Pimia rhamnoides Seem.,
shown that there are basically three clades, one New
Bonplandia 10: 366. 1862.
World clade and an Old World clade, with a number
of African Capparaceae genera (Boscia Lam., Cadaba
250. Bixaceae. Reference: Johnson-Fulton & Watson Forssk., Maerua Forssk., Ritchiea R.Br. ex G.Don and
(2017). Thilachium Lour.) sister to the Old World Capparis
Note: Amoreuxia Moq. was found to make that includes Australian Apophyllum F.Muell.,
Cochlospermum Kunth paraphyletic. As the genera which is here transferred. Dhofaria A.G.Mill. was
are not very divergent and morphologically similar, not sampled, but is morphologically similar to

Apophyllum and is hence also placed in Capparis. Cornejo & Iltis, J. Bot. Res. Inst. Texas 2: 76. 2008.
The circumscription of American Capparis species Morisonia bonifaziana (Carnejo & Iltis) Christenh. &
has been contentious and even though a high degree Byng, comb. nov. Basionym: Capparis bonifaziana
of splitting can be applied, many of these genera will Cornejo & Iltis, Novon 15: 397. 2005.
still not be monophyletic and not all taxa have been Morisonia brasiliana (Banks ex DC.) Christenh. & Byng,
transferred from Capparis. It is thus best to treat all comb. nov. Basionym: Capparis brasiliana Banks ex
New World Capparis as a single genus, the name DC., Prodr. 1: 249. 1824.
Morisonia L. having nomenclatural priority. Sister
Morisonia brasiliana (DC.) Christenh. & Byng, comb.
to all other Capparaceae is a clade composed of
nov. Basionym: Capparis brasiliana DC., Prodr. 1:
Crateva L. with Euadenia Oliv. embedded in it, the
249. 1824.
latter of which is also hereby transferred.
Morisonia breynia (L.) Christenh. & Byng, comb. nov.
Capparis anomala (F.Muell.) Christenh. & Byng, comb.
Basionym: Capparis breynia L., Syst. Pl. ed. 10: 1071.
nov. Basionym: Apophyllum anomalum F.Muell.,
1759. Replaced synonym: Breynia indica L., Sp. Pl. 1:
Hooker’s J. Bot. Kew Gard. Misc. 9: 307. 1857. (Figure
503. 1753, nom. rej.
16C)
Morisonia calciphila (Standl. & Steyerm.) Christenh.
Capparis macleishii (A.G.Mill.) Christenh. & Byng,
& Byng, comb. nov. Basionym: Capparis calciphila
comb. nov. Basionym: Dhofaria macleishii A.G.Mill.,
Standl. & Steyerm., Field. Mus. Bot. 23: 158. 1944.
Notes Roy. Bot. Gard., Edinburgh 45: 55. 1988.
Morisonia coimbrana (Cornejo & Iltis) Christenh. &
Crateva brevipetala (Exell) Christenh. & Byng, comb.
Byng, comb. nov. Basionym: Capparis coimbrana
nov. Basionym: Euadenia brevipetala Exell, J. Bot. 73
Cornejo & Iltis, Brittonia 57: 155. 2005.
(Suppl.): 12. 1935.
Morisonia crotonoides (Kunth) Christenh. & Byng,
Crateva eminens (Hook.f.) Christenh. & Byng, comb.
comb. nov. Basionym: Capparis crotonoides Kunth,
nov. Basionym: Euadenia eminens Hook.f., Bot. Mag.
Nov. Gen. Sp. Pl. (quarto ed.) 5: 95. 1821.
107: t. 6578. 1881. (Figure 16B)
Morisonia cuatrecasana (Dugand) Christenh. & Byng,
Crateva kirkii (Oliv.) Christenh. & Byng, comb. nov.
comb. nov. Basionym: Capparis cuatrecasana
Basionym: Euadenia kirkii Oliv., Fl. Trop. Afr. 1: 91.
Dugand, Caldasia 1: 41. 1941.
1868.
Morisonia cynophallophora (L.) Christenh. & Byng,
Crateva monticola (Gilg & Gilg-Ben.) Christenh. &
comb. nov. Basionym: Capparis cynophallophora L.,
Byng, comb. nov. Basionym: Euadenia monticola
Sp. Pl.: 504. 1753.
Gilg & Gilg-Ben., Bot. Jahrb. Syst. 53: 170. 1915.
Replaced synonym: Euadenia trifoliata (Vahl) Oliv.; Morisonia declinata (Vell.) Christenh. & Byng, comb.
syn.: Stroemia trifoliata Schumach. & Thonn., nor: nov. Basionym: Capparis declinata Vell., Fl. Flumin.:
Crateva trifoliata (Roxb.) B.S.Sun 230. 1829.
Morisonia alainiana (Cornejo & Iltis) Christenh. & Morisonia detonsa (Triana & Planch.) Christenh. &
Byng, comb. nov. Basionym: Quadrella alainiana Byng, comb. nov. Basionym: Capparis detonsa
Cornejo & Iltis, J. Bot. Res. Inst. Texas 4: 75. 2010. Triana & Planch., Ann. Sci. Nat., Bot., sér. 4, 17: 80.
1862.
Morisonia amplissima (Lam.) Christenh. & Byng,
comb. nov. Basionym: Capparis amplissima Lam., Morisonia didymobotrys (Ruiz & Pav. ex DC.)
Encycl. 1: 607. 1785. Christenh. & Byng, comb. nov. Basionym: Capparis
didymobotrys Ruiz & Pav. ex DC., Prodr. 1: 253. 1824.
Morisonia angustifolia (Kunth) Christenh. & Byng,
comb. nov. Basionym: Capparis angustifolia Kunth, Morisonia discolor (Donn.Sm.) Christenh. & Byng,
Nov. Gen. Sp. Pl. 5: 46. 1821. comb. nov. Basionym: Capparis discolor Donn.Sm.,
Bot. Gaz. 24: 389. 1897.
Morisonia antonensis (Woodson) Christenh. &
Byng, comb. nov. Basionym: Capparis antonensis Morisonia dolichopoda (Helwig) Christenh. & Byng,
Woodson, Ann. Missouri Bot. Gard. 35: 90. 1948. comb. nov. Basionym: Capparis dolichopoda Helwig,
Ark. Bot. 22A, no. 10: 11. 1929.
Morisonia asperifolia (C.Presl) Christenh. & Byng,
comb. nov. Basionym: Capparis asperifolia C.Presl, Morisonia domingensis (Spreng. ex DC.) Christenh. &
Reliq. Haenk. 2:86. 1835. Byng, comb. nov. Basionym: Capparis domingensis
Spreng. ex DC., Prodr. 1: 253. 1824.
Morisonia atamisquea (Kuntze) Christenh. & Byng,
comb. nov. Basionym: Capparis atamisquea Kuntze, Morisonia dressleri (Cornejo & Iltis) Christenh. & Byng,
Revis. Gen. Pl. 3: 6. 1898. comb. nov. Basionym: Quadrella dressleri Cornejo &
Iltis, J. Bot. Res. Inst. Texas 4: 77. 2010.
Morisonia bahiana (Cornejo & Iltis) Christenh. & Byng,
comb. nov. Basionym: Colicodendron bahianum Morisonia ecuadorica (Iltis) Christenh. & Byng, comb.
nov. Basionym: Capparis ecuadorica Iltis, Selbyana

139
A B
D E
Figure 16: Angiosperms (Eudicots). A, Cistus commutatus (Pau) Byng & Christenh.; B, Crateva eminens (Hook.f.)
Christenh. & Byng; C, Capparis anomala (F.Muell.) Christenh. & Byng; D, Goniolimon salicorniaceum (F.Muell.)
Christenh. & Byng; E, Morisonia frondosa (Jacq.) Christenh. & Byng; F, Sesamum trilobum (Bernh.) Byng & Christenh.;
and G, Morisonia paradoxa (Jacq.) Christenh. & Byng. All images © Maarten Christenhusz.

2: 303. 1978. Pl. 24: 1760.

Morisonia elegans (Mart.) Christenh. & Byng, comb. Morosonia lineata (Dombey ex Pers.) Christenh.
nov. Basionym: Capparis elegans Mart., Flora 22 (1, & Byng, comb. nov. Basionym: Capparis lineata
Beibl. 1): 24. 1839. Dombey ex Pers., Syn. Pl. 2: 60. 1807.
Morisonia ferruginea (L.) Christenh. & Byng, comb. Morisonia longifolia (Mart.) Christenh. & Byng, comb.
nov. Basionym: Capparis ferruginea L., Syst. Nat., ed. nov. Basionym: Colicodendron longifolium Mart.,
10: 1071. 1759. Flora 22 (2, Beibl. 1): 26. 1839.
Morisonia filipes (Donn.Sm.) Christenh. & Byng, comb. Morisonia lundellii (Standl.) Christenh. & Byng, comb.
nov. Basionym: Capparis filipes Donn.Sm., Bot. Gaz. nov. Basionym: Capparis lundellii Standl., Carnegie
23: 2. 1897. Inst. Wash. Publ. 461: 57. 1935.
Morisonia frondosa (Jacq.) Christenh. & Byng, comb. Morisonia macrantha (Standl.) Christenh. & Byng,
nov. Basionym: Capparis frondosa Jacq., Enum. Syst. comb. nov. Basionym: Steriphoma macranthum
Pl.: 24. 1760. (Figure 16E) Standl., Contr. U.S. Natl. Herb. 20: 183. 1919, as
Morisonia gonaivensis (Helwig) Christenh. & Byng, ‘macrantha’.
comb. nov. Basionym: Capparis gonaivensis Helwig, Morisonia macrophylla (Kunth) Christenh. & Byng,
Ark. Bot. 22A: 10. 1929. comb. nov. Basionym: Capparis macrophylla Kunth,
Morisonia grandiflora (Cornejo & Iltis.) Christenh. Nov. Gen. Sp. (quarto ed.) 5: 91. 1821.
& Byng, comb. nov. Basionym: Capparidastrum Morisonia martiana (Cornejo) Christenh. & Byng,
grandiflorum Cornejo & Iltis, Harvard Pap. Bot. 15: comb. nov. Basionym: Colicodendron martianum
155. 2010. Cornejo, Harvard Pap. Bot. 15: 103. 2010.
Morisonia guayaquilensis (Kunth) Christenh. & Byng, Morisonia matogrossensis (Pilg.) Christenh. & Byng,
comb. nov. Basionym: Capparis guayaquilensis comb. nov. Basionym: Capparis matogrossensis
Kunth, Nov. Gen. Sp. Pl. (quarto ed.) 5: 89. 1821. Pilg., Rev. Mus. Paul. Univ. São Paulo 13: 1250. 1922.
Morisonia hagenii (H.Karst.) Christenh. & Byng, comb. Morisonia megalosperma (Cornejo & Iltis) Christenh.
nov. Basionym: Belencita hagenii H.Karst., Berliner & Byng, comb. nov. Basionym: Capparidastrum
Allg. Gartenzeit. 25: 25. 1857. megalospermum Cornejo & Iltis, Harvard Pap. Bot.
Morisonia hastata (Jacq.) Christenh. & Byng, comb. 15: 159. 2010.
nov. Basionym: Capparis hastata Jacq., Enum. Syst. Morisonia mirifica (Standl.) Christenh. & Byng, comb.
Pl.: 23. 1760. nov. Basionym: Capparis mirifica Standl. in Woodson
Morisonia heterophylla (Ruiz & Pav. ex DC.) Christenh. & Schery, Ann. Missouri Bot. Gard. 27: 311. 1940.
& Byng, comb. nov. Basionym: Capparis heterophylla Morisonia mollicella (Standl.) Christenh. & Byng,
Ruiz & Pav. ex DC., Prodr. 1: 250. 1824. comb. nov. Basionym: Capparis mollicella Standl.,
Morisonia heydeana (Donn.Sm.) Christenh. & Byng, Proc. Biol. Soc. Washington 37: 44. 1924.
comb. nov. Basionym: Capparis heydeana Donn. Morisonia morenoi (Cornejo & Iltis) Christenh. & Byng,
Sm., Bot. Gaz. 18: 197. 1893. comb. nov. Basionym: Quadrella morenoi Cornejo &
Morisonia humilis (Jacq.) Christenh. & Byng, comb. Iltis, J. Bot. Res. Inst. Texas 4: 86. 2010.
nov. Basionym: Capparis humilis Hassl., Repert. Morisonia nemorosa (Jacq.) Christenh. & Byng, comb.
Spec. Nov. Regni Veg. 12: 252. 1913. nov. Basionym: Capparis nemorosa Jacq., Enum.
Morisonia incana (Kunth) Christenh. & Byng, comb. Syst. Pl. 24. 1760.
nov. Basionym: Capparis incana Kunth, Nov. Gen. Sp. Morisonia odoratissmia (Jacq.) Christenh. & Byng,
5: 94. 1821. comb. nov. Basionym: Capparis odoratissima Jacq.,
Morisonia isthmensis (Eichler) Christenh. & Byng, Hort. Schoenbr. 1: 57. 1797.
comb. nov. Basionym: Capparis isthmensis Eichler in Morisonia osmantha (Diels) Christenh. & Byng, comb.
Martius, Fl. Bras. 13: 269. 1865. nov. Basionym: Capparis osmantha Diels, Notizbl.
Morisonia jamaicensis (Jacq.) Christenh. & Byng, Bot. Gart. Berlin-Dahlem 14: 332. 1939.
comb. nov. Basionym: Capparis jamaicensis Jacq., Morisonia pachaca (Kunth) Christenh. & Byng, comb.
Enum. Pl. Carib. 23. 1760. nov. Basionym: Capparis pachaca Kunth, Nov. Gen.
Morisonia lindeniana (Cornejo & Iltis) Christenh. & Sp. Pl. (quarto ed.) 5: 93. 1821.
Byng, comb. nov. Basionym: Quadrella lindeniana Morisonia panamensis (Iltis) Christenh. & Byng, comb.
Cornejo & Iltis, J. Bot. Res. Inst. Texas 4: 83. 2010. nov. Basionym: Capparis panamensis Iltis, Ann.
Morisonia linearis (Jacq.) Christenh. & Byng, comb. Missouri Bot. Gard. 68: 681. 1981.
nov. Basionym: Capparis linearis Jacq., Enum. Syst. Morisonia paradoxa (Jacq.) Christenh. & Byng, comb.
nov. Basionym: Capparis paradoxa Jacq., Pl. Hort.

141
Schoenbr. 1: 58. 1797. (Figure 16G) 5: 359. 1934.

Morisonia peruviana (Spruce ex Echler) Christenh. & Morisonia speciosa (Iltis) Christenh. & Byng, comb.
Byng, comb. nov. Basionym: Steriphoma peruvianum nov. Basionym: Capparis speciosa Griseb., Abh.
Spruce ex Eichler in Mart., Fl. Bras. 13: 267. 1865. Königl. Ges. Wiss. Göttingen 24: 18. 1879.
Morisonia petiolaris (Kunth) Christenh. & Byng, comb. Morisonia sprucei (Eichler) Christenh. & Byng, comb.
nov. Basionym: Capparis petiolaris Kunth, Nov. Gen. nov. Basionym: Capparis sprucei Eichler in Mart., Fl.
Sp. Pl. (quarto ed.) 5: 91. 1821. Bras. 13: 281. 1865.
Morisonia pittieri (Standl.) Christenh. & Byng, comb. Morisonia stenopetala (Urb.) Christenh. & Byng,
nov. Basionym: Capparis pittieri Standl., J. Wash. comb. nov. Basionym: Capparis stenosepala Urb.,
Acad. Sci. 17: 253. 1927. Symb. Antill. 5: 529. 1908.
Morisonia polyantha (Triana & Planch.) Christenh. & Morisonia stephania Christenh. & Byng, nom. nov.
Byng, comb. nov. Basionym: Capparis polyantha Replaced synonym: Stephania elliptica DC., Prodr. 1:
Triana & Planch., Ann. Sci. Nat., Bot., sér. 4, 17: 76. 253. 1824, not: Morisonia elliptica Rusby.
1862. Morisonia steyermarkii (Standl.) Christenh. & Byng,
Morisonia pringlei (Briq.) Christenh. & Byng, comb. comb. nov. Basionym: Capparis steyermarkii Standl.,
nov. Basionym: Capparis pringlei Briq., Ann. Conserv. Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 140. 1940.
& Jard. Bot. Genève 17:390. 1914. Morisonia tafallana (Cornejo, Iltis & Cerón) Christenh.
Morisonia pulcherrima (Jacq.) Christenh. & Byng, & Byng, comb. nov. Basionym: Capparidastrum
comb. nov. Basionym: Capparis pulcherrima Jacq., tafallanum Cornejo, Iltis & Cerón, Harvard Pap. Bot.
Enum. Syst. Pl.: 24. 1760. 19: 189. 2014.
Morisonia quina (J.F.Macbr.) Christenh. & Byng, comb. Morisonia tarapotensis (Echler) Christenh. & Byng,
nov. Basionym: Capparis quina J.F.Macbr., Candollea comb. nov. Basionym: Capparis tarapotensis Echler
5: 357. 1934. in Mart., Fl. Bras. 13: 284. 1865.
Morisonia quintanarooensis (Iltis & Cornejo) Morisonia tenuisiliqua (Jacq.) Christenh. & Byng,
Christenh. & Byng, comb. nov. Basionym: Quadrella comb. nov. Basionym: Capparis tenuisiliqua Jacq.,
quintanarooensis Iltis & Cornejo, J. Bot. Res. Inst. Enum. Syst. Pl.: 24. 1760.
Texas 4: 104. 2010. Morisonia tuxtlensis (Cornejo & Iltis) Christenh.
Morisonia quiriguensis (Standl.) Christenh. & Byng, & Byng, comb. nov. Basionym: Capparidastrum
comb. nov. Basionym: Capparis quiriguensis Standl., tuxtlense Cornejo & Iltis, Brittonia 62: 120. 2010.
Proc. Biol. Soc. Washington 37: 52. 1924. Morisonia tweediana (Eichler) Christenh. & Byng,
Morisonia retusa (Griseb.) Christenh. & Byng, comb. comb. nov. Basionym: Capparis tweediana Eichler in
nov. Basionym: Capparis retusa Griseb., Abh. Königl. Mart., Fl. Bras. 13: 273. 1865.
Ges. Wiss. Göttingen 24: 18. 1879. Morisonia urbanii (Eggers) Christenh. & Byng, comb.
Morisonia salicifolia (Griseb.) Christenh. & Byng, nov. Basionym: Steriphoma urbanii Eggers, Bot.
comb. nov. Basionym: Capparis salicifolia Griseb., Zentralbl. 73: 51. 1898.
Abh. Königl. Ges. Wiss. Göttingen 24: 17. 1879. Morisonia valerabella (Iltis, T.Ruiz & G.S.Bunting)
Morisonia scabrida (Kunth) Christenh. & Byng, comb. Christenh. & Byng, comb. nov. Basionym:
nov. Basionym: Capparis scabrida Kunth, Nov. Gen. Colicodendron valerabellum Iltis, T.Ruiz &
Sp. Pl. (quarto ed.) 5: 95. 1821. G.S.Bunting, J. Bot. Res. Inst. Texas 2: 82. 2008.
Morisonia sclerophylla (Iltis) Christenh. & Byng, comb. Morisonia verrucosa (Jacq.) Christenh. & Byng, comb.
nov. Basionym: Capparis sclerophylla Iltis & Cornejo, nov. Basionym: Capparis verrucosa Jacq., Enum.
Novon 15: 429. 2005. Syst. Pl.: 23. 1760.
Morisonia sessilis (Banks ex DC.) Christenh. & Byng, Morisonia yco (Mart.) Christenh. & Byng, comb. nov.
comb. nov. Basionym: Capparis sessilis Banks ex DC., Basionym: Capparis yco Mart., Reise Bras. 2: 599.
Prodr. 1: 249. 1824. 1828.
Morisonia siliquosa (L.) Christenh. & Byng, comb. nov. Morisonia yunckeri (Standl.) Christenh. & Byng, comb.
Basionym: Capparis siliquosa L., Syst. Nat. ed. 10 (2): nov. Basionym: Capparis yunckeri Standl., Publ. Field
1071. 1759. Mus. Nat. Hist., Bot. Ser. 9: 291. 1940.
Morisonia singularis (R.Rankin) Christenh. & Byng,
comb. nov. Basionym: Capparis singularis R.Rankin, 269. Cleomaceae. References: Inda et al. (2008),
Willdenowia 34: 263. 2004. Patchell et al. (2014).
Morisonia sola (J.F.Macbr.) Christenh. & Byng, comb. Note: Cleome L. was traditionally considered a single
nov. Basionym: Capparis sola J.F.Macbr., Candollea large genus, but the generic concept has narrowed

and diverged adopting a large number of small Hist. ser. 3, 4: 362. 1859.
genera that were long recognised as sections or Olax papillosa (J.O.Rangel) Christenh. & Byng, comb.
series (Iltis, 1952). This splitting of an otherwise nov. Basionym: Liriosma papillosa J.O.Rangel, Bol.
monophyletic clade seems to be unnecessarily Mus. Bot. Munic. 14: 2. 1974.
destabilising the classification of this clade and Olax redmondii (Steyerm.) Christenh. & Byng, comb.
hence here we accept a broader concept of Cleome, nov. Basionym: Dulacia redmondii Steyerm.,
keeping in mind the monophyly of the genus. The Phytologia 38: 217. 1978.
family then consists of four monophyletic genera:
Olax singularis (Vell.) Christenh. & Byng, comb. nov.
Cleomella DC., Peritoma DC. (incl. Isomeria Nutt.),
Basionym: Dulacia singularis Vell., Fl. Flumin. 32.
Rorida J.F.Gmel. and Cleome (now including
1829.
Gynandropsis DC., Haptocarpum Ule, Polanisia
A.Juss., Puccionia Chiov., Siliquaria Forssk., Tarenaya Olax tepuiensis (Steyerm.) Christenh. & Byng, comb.
Raf. and others). This requires the fewest number of nov. Basionym: Liriosma tepuiensis Steyerm., Bol.
name changes and is likely to be easily applied by Soc. Venez. Ci. Nat. 32: 328. 1976.
the general user.
Cleome bahiensis (Ule) Christenh. & Byng, comb. nov. 276. Santalaceae. References: Danser (1940, 1955), Der
Basionym: Haptocarpum bahiense Ule, Ber. Deutsch. & Nickrent (2008), Kuijt & Hansen (2014).
Bot. Ges. 26A: 223. 1908. Note: Danser (1940) split Henslowia Blume into four
Cleome puccionia Christenh. & Byng, nom. nov. separate genera (Dendromyza Danser, Cladomyza
Replaced synonym: Puccionia macradenia Chiov., Danser, Hylomyza Danser and Dendrotrophe
Fl. Somala 1: 78. 1929, not: Cleome macradenia Danser) based on fruit morphology. Hylomyza was
Schweinf. subsequently found to be a synonym of Dufrenoya
Chati, but as more material was collected it became
increasingly difficult to recognise the other three
273. Olacaceae. References: Malecot et al. (2004),
genera as distinct from each other. The general trend
Malecot & Nickrent (2008), Su et al. (2015).
in the literature is sinking Cladomyza in Dendromyza,
Note: Olax L. is divided into four sections based on the but synonymising Dendrotrophe appears to be
number of fertile stamens and staminodes: sect. premature at this stage.
Pentandrae Engler (5–6 stamens and 3 staminodes),
Dendromyza acrosclera (Danser) Byng & Christenh.,
sect. Hemiandrae Engler (6 stamens and 6
comb. nov. Basionym: Cladomyza acrosclera Danser,
staminodes), sect. Triandrae Engler (3 stamens and
Nova Guinea ser. 2, 4: 138. 1940.
5 staminodes), and sect. Estaminodiales Engler (6
stamens, staminodes lacking). Neotropical Dulacia Dendromyza acutata (Pilg.) Byng & Christenh., comb.
Vellozo was mainly distinguished from paleotropical nov. Basionym: Henslowia acutata Pilg., Bot. Jahrb.
Olax by distyly and is embedded in Olax. Dulacia Syst. 59: 125. 1924.
have three fertile stamens and six staminodes and Dendromyza angustifolia (Stauffer) Byng & Christenh.,
this is not so different from the great staminal comb. nov. Basionym: Cladomyza angustifolia
variation that is found in Olax. We hereby make the Stauffer., Mitt. Bot. Mus. Univ. Zurich 242: 64. 1969.
necessary combinations. Dendromyza cuneata (Danser) Byng & Christenh.,
Olax candida (Poepp.) Christenh. & Byng, comb. nov. comb. nov. Basionym: Cladomyza cuneata Danser,
Basionym: Liriosma candida Poepp., Nov. Gen. Sp. Pl. Nova Guinea ser. 2, 4: 137. 1940.
(Poeppig & Endlicher) 3: 33, t. 239. 1841. Dendromyza dendromyzoides (Stauffer) Byng &
Olax crassa (Monach.) Christenh. & Byng, comb. nov. Christenh., comb. nov. Basionym: Cladomyza
Basionym: Liriosma crassa Monach., Brittonia 13: dendromyzoides Stauffer, Mitt. Bot. Mus. Univ.
114. 1961. Zurich 242: 62. 1969.
Olax cyanocarpa (Sleumer) Christenh. & Byng, comb. Dendromyza densa (Stauffer) Byng & Christenh.,
nov. Basionym: Dulacia cyanocarpa Sleumer, Fl. comb. nov. Basionym: Cladomyza densa Stauffer,
Neotrop. Monogr. 38: 130. 1984. Mitt. Bot. Mus. Univ. Zurich 242: 63. 1969.
Olax egleri (J.O.Rangel) Christenh. & Byng, comb. nov. Dendromyza dubia (Stauffer) Byng & Christenh., comb.
Basionym: Liriosma egleri J.O.Rangel, Bol. Mus. Bot. nov. Basionym: Cladomyza dubia Stauffer, Mitt. Bot.
Munic. 14: 1. 1974. Mus. Univ. Zurich 242: 65. 1969.
Olax guianensis (Engl.) Christenh. & Byng, comb. Dendromyza erecta (Danser) Byng & Christenh., comb.
nov. Basionym: Liriosma guianensis Engl., Fl. Bras. nov. Basionym: Cladomyza erecta Danser, Nova
(Martius) 12: 25. 1872. Guinea ser. 2, 6: 275. 1955.
Olax inopiflora (Miers) Christenh. & Byng, comb. nov. Dendromyza gracilis (Danser) Byng & Christenh.,
Basionym: Liriosma inopiflora Miers, Ann. Mag. Nat. comb. nov. Basionym: Cladomyza gracilis Danser,

143
Nova Guinea ser. 2, 6: 275. 1955. a synonym of Plumbago.

Dendromyza kaniensis (Pilg.) Byng & Christenh., comb. Plumbago pendula (Kuntze) Christenh. & Byng, comb.
nov. Basionym: Henslowia kaniensis Pilg., Bot. Jahrb. nov. Basionym: Dyerophytum pendulum Kuntze,
Syst. 59: 127. 1924. Revis. Gen. Pl. 2: 394. 1891.
Dendromyza laevis (Pilg.) Byng & Christenh., comb.
nov. Basionym: Henslowia laevis Pilg., Bot. Jahrb. 314. Talinaceae. References: Hershkovitz & Zimmer
Syst. 59: 125. 1924. (1997), Applequist & Wallace (2001), Nyffeler (2007).
Dendromyza latifolia (Danser) Byng & Christenh., Note: Talinella Baill. is found to be inseparable from
comb. nov. Basionym: Cladomyza latifolia Danser, Talinum Adans., so the two genera are hereby
Nova Guinea ser. 2, 4: 141. 1940. merged.
Dendromyza microphylla (Lauterb.) Byng & Christenh., Talinum albiflorum (Appleq.) Christenh. & Byng,
comb. nov. Basionym: Henslowia microphylla comb. nov. Basionym: Talinella albidiflora Appleq.,
Lauterb., Nova Guinea 8: 813. 1912. Adansonia, sér. 3, 27: 52. 2005.
Dendromyza multinervis (Danser) Byng & Christenh., Talinum ankaranense (Appleq.) Christenh. & Byng,
comb. nov. Basionym: Cladomyza multinervis comb. nov. Basionym: Talinella ankaranensis
Danser, Nova Guinea ser. 2, 4: 139. 1940. Appleq., Adansonia, sér. 3., 27: 54. 2005.
Dendromyza nivalis (Ridl.) Byng & Christenh., comb. Talinum boivinianum (Baill.) Christenh. & Byng, comb.
nov. Basionym: Henslowia nivalis Ridl., Trans. Linn. nov. Basionym: Talinella boiviniana Baill., Bull. Mens.
Soc. London, Bot. 9: 146. 1916. Soc. Linn. Paris 1: 569. 1886.
Dendromyza robustior (Danser) Byng & Christenh., Talinum bosseri (Appleq.) Christenh. & Byng, comb.
comb. nov. Basionym: Cladomyza robustior Danser, nov. Basionym: Talinella bosseri Appleq., Adansonia,
Nova Guinea ser. 2, 4: 137. 1940. sér. 3, 27: 58. 2005.
Dendromyza stellata (Stauffer) Byng & Christenh., Talinum dauphinense (Scott-Elliot) Christenh. & Byng,
comb. nov. Basionym: Cladomyza stellata Stauffer, comb. nov. Basionym: Talinella dauphinensis Scott-
Mitt. Bot. Mus. Univ. Zurich 242: 62. 1969. Elliot, J. Linn. Soc., Bot. 29: 4. 1891.
Dendromyza trinervia (Stauffer) Byng & Christenh., Talinum grevei (Danguy) Christenh. & Byng, comb.
comb. nov. Basionym: Cladomyza trinervia Stauffer, nov. Basionym: Talinella grevei Danguy, Notul. Syst.
Mitt. Bot. Mus. Univ. Zurich 242: 65. 1969. (Paris) 3: 159. 1915.
Dendromyza uncinata (Danser) Byng & Christenh., Talinum humbertii (Appleq.) Christenh. & Byng,
comb. nov. Basionym: Cladomyza uncinata Danser, comb. nov. Basionym: Talinella humbertii Appleq.,
Nova Guinea ser. 2, 4: 139. 1940. Adansonia, sér. 3, 27: 68. 2005.
Leptomeria rubescens (Stauffer) Byng & Christenh., Talinum latifolium (Appleq.) Christenh. & Byng,
comb. nov. Basionym: Spirogardnera rubescens comb. nov. Basionym: Talinella latifolia Appleq.,
Stauffer, Mitt. Bot. Mus. Univ. Zurich 240: 307. 1968. Adansonia, sér. 3, 27: 70. 2005.
Talinum microphyllum (Eggli) Christenh. & Byng,
282. Plumbaginaceae. Reference: Lledó et al. (2005) comb. nov. Basionym: Talinella microphylla Eggli,
Note: Muellerolimon Lincz. was found to be embedded Adansonia, sér. 3, 19: 56. 1997.
in Goniolimon Boiss. and is hereby transferred. Talinum pachypodum (Eggli) Christenh. & Byng,
Plumbago L. is expanded with Dyerophytum Kuntze. comb. nov. Basionym: Talinella pachypoda Eggli,
Goniolimon salicorniaceum (F.Muell.) Christenh. & Adansonia, sér. 3, 19: 57. 1997.
Byng, comb. nov. Basionym: Statice salicorniacea Talinum tsitondroinense (Appleq.) Christenh. & Byng,
F.Muell., Fragm. 11: 7. 1878. (Figure 16G) comb. nov. Basionym: Talinella tsitondroinensis
Plumbago africana (Lam.) Christenh. & Byng, comb. Appleq., Adansonia, sér. 3, 27: 75. 2005.
nov. Basionym: Vogelia africana Lam., Icon. Pl. 2: Talinum xerophilum (Appleq.) Christenh. & Byng,
147. 1792. comb. nov. Basionym: Talinella xerophila Appleq.,
Plumbago arabica (Boiss.) Christenh. & Byng, comb. Adansonia, sér. 3 27: 76. 2005.
nov. Basionym: Vogelia arabica Boiss. in DC. Prodr.
12: 696. 1848. 332. Pentaphylacaceae. Reference: Luna & Ochoterena
Plumbago dyerophyta Christenh. & Byng, nom. nov. (2004).
Replaced synonym: Dyerophytum indicum Kuntze, Note: Killipiodendron Kobuski is here moved to Freziera
Revis. Gen. Pl. 2: 394. 1891, not: Plumbago indica L. Willd., where molecular data places it.
Etymology: Named for the genus Dyerophytum, now Freziera colombiana (Kobuski) Christenh. & Byng,
comb. nov. Basionym: Killipiodendron colombianum

Kobuski, J. Arnold Arbor. 23: 231. 1942. comb. & stat. nov. Basionym: Labordia hedyosmifolia
Baill. var. hosakana Sherff, Amer. J. Bot. 25: 584.
1938.
348. Icacinaceae. References: Manchester (1994), Stull
(unpublished data). Geniostoma hymenopodum O.Deg. & Sherff) Byng
& Christenh., comb. nov. Basionym: Labordia
Note: The replacement name Pyrenacantha
hymenopoda O.Deg. & Sherff, Amer. J. Bot. 38: 60.
occidentalis Byng & Utteridge for Polycephalium
1951. Replaced synonym: Labordia sessilis A.Gray,
capitatum (Baill.) Keay is invalid, because an earlier
Proc. Amer. Acad. Arts & Sci. 4: 323. 1859, not:
homonym existed for a fossil taxon. A new name is
Geniostoma sessile Kaneh.
here proposed. Nothapodytes Blume is embedded
in Mappia Jacq. and is hereby transferred. Geniostoma lorencianum (K.R.Wood, W.L.Wagner &
T.Motley) Byng & Christenh., comb. nov. Basionym:
Mappia amamiana (Nagam. & Mak.Kato) Byng & Stull,
Labordia lorenciana K.R.Wood, W.L.Wagner &
comb. nov. Basionym: Nothapodytes amamianus
T.Motley, Syst. Bot. 32: 195. 2007.
Nagam. & Mak.Kato, Acta Phytotax. Geobot. 55: 75.
2004. Geniostoma lydgatei (C.N.Forbes) Byng & Christenh.,
comb. nov. Basionym: Labordia lydgatei C.N.Forbes,
Mappia collina (C.Y.Wu) Byng & Stull., comb. nov.
Occas. Pap. Bernice Pauahi Bishop Mus. 6: 60. 1916,
Basionym: Nothapodytes collina C.Y.Wu, Fl. Yunnan.
as ‘lydgateii’.
1: 169. 1977.
Geniostoma pumilum (Hillebr.) Byng & Christenh.,
Mappia nimmoniana (J.Grah.) Byng & Stull, comb.
comb. & stat. nov. Basionym: Labordia fagraeoidea
nov. Basionym: Premna nimmoniana J.Grah., Cat. Pl.
Gaudich. var. pumila Hillebr., Fl. Hawaiian Isl.: 290.
Bombay: 155. 1839.
1888.
Mappia obscura (C.Y.Wu) Byng & Stull, comb. nov.
Geniostoma tinifolium (A.Gray) Byng & Christenh.,
Basionym: Nothapodytes obscura C.Y.Wu, Fl. Yunnan.
comb. nov. Basionym: Labordia tinifolia A.Gray, Proc.
1: 170. 1977.
Amer. Acad. Arts. Sci. 4: 322. 1859.
Mappia tomentosa (C.Y.Wu) Byng & Stull, comb. nov.
Geniostoma tinifolium (A.Gray) Byng & Christenh. var.
Basionym: Nothapodytes tomentosa C.Y.Wu, Fl.
lanaiense (Sherff) Byng & Christenh., comb. nov.
Yunnan. 1: 169. 1977.
Basionym: Labordia tinifolia A.Gray var. lanaiensis
Pyrenacantha manchesteriana Byng & Christenh., Sherff, Amer. J. Bot. 25: 583. 1938.
nom. nov. Basionym: Chlamydocarya capitata Baill.,
Geniostoma tinifolium (A.Gray) Byng & Christenh.
Adansonia n.s. 10: 277. 1872, not: Pyrenacantha
var. wahiawaense (H.St.John) Byng & Christenh.,
capitata H.Perrier. Replaced synonym: Pyrenacantha
comb. nov. Basionym: Labordia tinifolia A.Gray var.
occidentalis Byng & Utteridge, Bot. J. Linn. Soc.
wahiawaensis H.St.John, Bull. Torrey Bot. Club 111:
176: 202. 2014, nom. illeg., not: P. occidentalis
481. 1984.
Manchester (fossil). Etymology: Named after Steve
Manchester a paleobotanist who has worked Geniostoma trifolium (Hillebr.) Byng & Christenh.,
extensively on the fossils of Icacinaceae and whose comb. nov. Basionym: Labordia triflora Hillebr., Fl.
fossil species necessitated this new name. Hawaiian Isl.: 293. 1888.
Geniostoma venosum (Sherff) Byng & Christenh.,
comb. nov. Basionym: Labordia venosa Sherff, Amer.
354. Loganiaceae. Reference: Gibbons et al. (2012).
J. Bot. 25: 579. 1938.
Note: Labordia is embedded in Geniostoma. The
Geniostoma waialealae (Wawra) Byng & Christenh.,
species are hereby transferred.
comb. nov. Basionym: Labordia waialealae Wawra,
Geniostoma degeneri (Sherff) Byng & Christenh., Flora 55: 516. 1872.
comb. nov. Basionym: Labordia degeneri Sherff in
Geniostoma waiolani (Wawra) Byng & Christenh.,
Degener, Fl. Hawaii.: Fam. 302. 1938.
comb. nov. Basionym: Labordia waiolani Wawra,
Geniostoma hedyosmifolium (Baill.) Byng & Christenh., Flora 55: 516. 1872.
comb. nov. Basionym: Labordia hedyosmifolia Baill.,
Bull. Mens. Soc. Linn. Paris 1: 240. 1880.
376. Pedaliaceae. Reference: Gormley et al. (2015).
Geniostoma helleri (Sherff) Byng & Christenh., comb.
nov. Basionym: Labordia helleri Sherff, Amer. J. Bot. Note: Ceratotheca Endl. and Josephinia Vent. are
25: 581. 1938. genetically and morphologically indistinct from
Sesamum L. and the species are thus hereby
Geniostoma hirtellum (H.Mann) Byng & Christenh.,
transferred.
comb. nov. Basionym: Labordia hirtella H.Mann,
Proc. Amer. Acad. Arts & Sci. 7: 196. 1867. Sesamum byngianum Christenh., nom. nov. Replaced
synonym: Josephinia africana Vatke, Linnaea 43:
Geniostoma hosakanum (Sherff) Byng & Christenh.,
541. 1882, not: Sesamum africanum Tod. Etymology:

145
Named for James W. Byng, for his contributions to several clades of this genus embedded in Verbena
African botany. L. Morphologically the two genera are very similiar,
Sesamum celebicum (Blume) Byng & Christenh., comb. differing mainly in style length, and there is evidence
nov. Basionym: Josephinia celebica Blume, Bijdr. Fl. of horizontal gene transfer between the ‘genera’.
Ned. Ind. 14: 779. 1826. We propose an expanded Verbena and transfer any
remaining names. Additionally, Xolocotzia Miranda
Sesamum christenhuszianum Byng, nom. nov.
is placed in Petrea L. and Baillonia Bocq. is merged
Replaced synonym: Josephinia grandiflora R.Br.,
with Citharexylum B.Juss.
Prodr. Fl. Nov. Holland.: 520. 1810, not: Sesamum
grandiflorum Schinz. Etymology: The species is Citharexylum amabile (Bocq. ex Baill.) Christenh.
named for Maarten J. M. Christenhusz, for his & Byng, comb. nov. Basionym: Baillonia amabilis
contributions to Australian botany. Bocq. ex Baill., Adansonia 2: 252. 1862.
Sesamum eriocarpum (Decne.) Byng & Christenh., Petrea xolocotzia Christenh. & Byng, nom. nov.
comb. nov. Basionym: Pretrea eriocarpa Decne., Replaced synonym: Xolocotzia asperifolia Miranda,
Duch. Rev. Bot. 1: 517. 1846. Bol. Soc. Bot. México 29: 40. 1965, not: Petrea
asperifolia Brongn. ex Moldenke. Etymology: Named
Sesamum eugeniae (F.Muell.) Byng & Christenh.,
for the former genus Xolocotzia, now a synonym of
comb. nov. Basionym: Josephinia eugeniae F.Muell.,
Petraea.
Hooker’s J. Bot. Kew Gard. Misc. 9: 370, t. 11. 1857.
Verbena alejandrana (B.L.Turner) Christenh. & Byng,
Sesamum forbesii (Decne.) Byng & Christenh., comb.
comb. nov. Basionym: Glandularia alejandrana
nov. Basionym: Pretrea forbesii Decne., Ann. Sci.
B.L.Turner, Lundellia 2: 57. 1999.
Nat., Bot. sér. 5, 3: 334. 1865.
Verbena angustilobata (Moldenke) Christenh. & Byng,
Sesamum imperatrice (Vent.) Byng & Christenh.,
comb. nov. Basionym: Verbena cabrerae Moldenke
comb. nov. Basionym: Josephinia imperatricis Vent.,
var. angustilobata Moldenke, Phytologia 28: 195.
Jard. Malmaison 2: t. 67. 1804.
1974.
Sesamum integribracteatum (Engl.) Byng & Christenh.,
Verbena brachyrhynchos (G.L.Nesom & Vorobik)
comb. nov. Basionym: Ceratotheca integribracteata
Engl., Bot. Jahrb. Syst. 19: 156. 1894.
Glandularia brachyrhynchos G.L.Nesom & Vorobik,
Sesamum papillosum (W.Fitzg.) Byng & Christenh., Sida 13: 31. 1988.
comb. nov. Basionym: Josephinia papillosa W.Fitzg.,
Verbena guaibensis (P.Peralta & V.Thode) Christenh. &
J. Proc. Roy. Soc. Western Australia 3: 207. 1918.
Byng, comb. nov. Basionym: Glandularia guaibensis
Sesamum reniforme (Abels) Byng & Christenh., P.Peralta & V.Thode, Rodriguésia 61: S30. 2010.
comb. nov. Basionym: Ceratotheca reniformis
Verbena malpaisana (T.Van Devender & G.L.Nesom)
Abels, Morphol., Geogr. Taxon., Dicerocaryum &
Ceratotheca (Hamburg): 177. 1973.
Glandularia malpaisana T.Van Devender &
Sesamum saxicola (E.A.Bruce) Byng & Christenh., G.L.Nesom, Phytoneuron 2012-65: 1. 2012.
comb. nov. Basionym: Ceratotheca saxicola
Verbena turneri (G.L.Nesom) Christenh. & Byng, comb.
E.A.Bruce, Bothalia 6: 233. 1951.
nov. Basionym: Glandularia turneri G.L.Nesom,
Sesamum senecioides (Klotzsch) Byng & Christenh., Phytologia 72: 383. 1992.
comb. nov. Basionym: Pretrea senecioides Klotzsch,
Naturw. Reise Mossambique [Peters] 6(Bot., 1): 189.
1861. 408. Adoxaceae. Reference: Liu et al. (2000).
Sesamum sesamoides (Endl.) Byng & Christenh., comb. Note: Based on molecular data Sinadoxa C.Y.Wu, Z.L.Wu
nov. Basionym: Ceratotheca sesamoides Endl., & R.F.Huang and Tetradoxa CY.Wu form a clade with
Linnaea 7: 5. 1832. Adoxa L. and the three genera are morphologically
very similar,. The variability in the other genera in
Sesamum trilobum (Bernh.) Byng & Christenh., comb.
Adoxaceae (Sambucus L. and Viburnum L.) are far
nov. Basionym: Sporledera triloba Bernh., Linnaea
greated and merging these genera are in line with
16: 42. 1842. (Figure 16F)
traditional generic circumscription in this family.
Sesamum zanguebarium (Lour.) Byng & Christenh., Tetradoxa was already described as Adoxa, so only
comb. nov. Basionym: Martynia zanguebaria Lour., one new combination is needed.
Fl. Cochinch. 2: 386. 1790.
Adoxa corydalifolia (C.Y.Wu, Z.L.Wu & R.F.Huang)
Christenh. & Byng, comb. nov. Basionym: Sinadoxa
382. Verbenaceae. References: Yuan & Olmstead corydalifolia C.Y.Wu, Z.L.Wu & R.F.Huang, Acta
(2008), Marx et al. (2010).
Note: Glandularia J.F.Gmel. is paraphyletic with

Phytotax. Sin. 19: 208. 1981. Soc. 71: 227. 1975.

Valeriana ×aurigerana (Giraudias) Christenh. & Byng,
409. Caprifoliaceae. References: Bell & Donoghue comb. nov. Basionym: Centranthus ×aurigeranus
(2005), Hidalgo et al. (2010), Giraudias, Bull. Soc. Études Sci. Angers 19: 107. 1889.
Note: Valeriana L. is widely polyphyletic with Valeriana battandieri (Maire) Christenh. & Byng,
Belonanthus Graebn., Centranthus Lam. & DC., comb. nov. Basionym: Centranthus battandieri
Fedia Gaertn., Phuodendron (Graebn.) Dalla Torre Maire, Bull. Soc. Hist. Nat. Afrique N. 17: 115. 1926.
& Harms, Plectritis (Lindl.) DC. and Valerianella Mill. Valeriana belonantha Christenh. & Byng, nom. nov.
embedded in it. Theoretically the Valeriana celtica Replaced synonym: Belonanthus angustifolius
L. clade can be segratated to maintain several of the Schmale, Notizbl. Bot. Gart. Berlin-Dahlem 13: 25.
genera, but as Valeriana longiflora Willk. is sister 1936, not: Valeriana angustifolia Mill. Etymology:
to the entire clade and not all species of Valeriana Named for the genus Belonanthus, now a synonym
have yet been sequenced, it is likely that recognising of Valeriana, from βελόνη, needle, and άνθος,
these as separate genera defies the purpose and flower.
it will be difficult to find morphological characters Valeriana brachystemon (Fisch. & C.A.Mey.) Byng
that distinguish these new genera. Therefore we & Christenh., comb. nov. Basionym: Plectritis
propose to treat Valeriana in the broad sense, a brachystemon Fisch. & C.A.Mey., Index Semin. (St.
morphologically coherent group, and we hereby Petersburg) 2: 47. 1836.
make the necessary combinations. Valeriana cadevallii (Sennen) Christenh. & Byng, nom.
Valeriana aegaea Christenh. & Byng, nom. nov. nov. Basionym: Centranthus cadevallii Sennen,
Replaced synonym: Valerianella hirsutissima Link, Treb. Inst. Catalana Hist. Nat.: 83. 1917. Replaced
Linnaea 9: 580. 1835, not: Valeriana hirsutissima synonym: Centranthus macrosiphon Boiss., Diagn.
Killip. Etymology: Named for the Aegean region, Pl. Orient. ser. 1, 3: 57. 1843, not: Valeriana
where this species occurs. macrosiphon E.Vilm.
Valeriana aligera Christenh. & Byng, nom. nov. Valeriana calycina (Maire) Christenh. & Byng, comb.
Replaced synonym: Aligera eichleriana Suksd., nov. Basionym: Fedia calycina Maire, Bull. Soc. Hist.
Deutsche Bot. Monatsschr. 15: 147. 1897, not: Nat. Afrique N. 22: 291. 1931.
Valeriana eichleriana (Muell.) Graebn. Etymology: Valeriana carinata (Loisel.) Christenh. & Byng, comb.
Named for the genus Aligera Suksd., now a synonym nov. Basionym: Valerianella carinata Loisel., Not. Fl.
of Valeriana. France: 149. 1810.
Valeriana amarella (Lindh. ex Engelm.) Christenh. & Valeriana caucasiana (Boiss.) Christenh. & Byng,
Byng, comb. nov. Basionym: Fedia amarella Lindh. comb. nov. Basionym: Betckea caucasiana Boiss., Fl.
ex Engelm., Boston J. Nat. Hist. 6: 217. 1857. Orient. 3: 94. 1875.
Valeriana amazonum (Fridl. & A.Raynal) Christenh. & Valeriana chenopodiifolia (Pursh) Christenh. & Byng,
Byng, comb. nov. Basionym: Centranthus amazonum comb. nov. Basionym: Fedia chenopodiifolia Pursh,
Fridl. & A.Raynal, Adansonia sér. 3, 20: 328. 1998. Fl. Amer. Sept. 2: 727. 1814.
Valeriana aphanoptera (A.Gray) Byng & Christenh., Valeriana ciliosa (Greene) Byng & Christenh., comb.
comb. nov. Basionym: Valerianella aphanoptera nov. Basionym: Valerianella ciliosa Greene, Proc.
A.Gray, Proc. Amer. Acad. Arts & Sci. 19: 83. 1883. Acad. Nat. Sci. Philadelphia 1895: 548. 1896.
Valeriana aretiastra Christenh. & Byng, nom. nov. Valeriana congesta (Lindl.) Byng & Christenh., comb.
Replaced synonym: Aretiastrum maximum Graebn., nov. Basionym: Valerianella congesta Lindl., Bot.
Repert. Spec. Nov. Regni Veg. 15: 323. 1918, not: Reg. 13: t. 1094. 1827.
Valeriana maxima Killip. Etymology: Named for
Valeriana coronata (L.) Christenh. & Byng, comb. nov.
Aretiastrum, now a synonym of Valeriana, from
Basionym: Valeriana locusta L. var. coronata L., Sp.
Greek αρετή, virtue.
Pl. 1: 34. 1753.
Valeriana arkansana Christenh. & Byng, comb. nov.
Valeriana davyana (Jeps.) Christenh. & Byng, comb.
Basionym: Valerianella palmeri Dyal, Rhodora
nov. Basionym: Plectritis davyana Jeps., Fl. W. Calif.:
40: 201. 1938, not: Valeriana palmeri A.Gray ex
475. 1901.
S.Watson. Etymology: Named for the US state
Arkansas, where the type of this species was Valeriana dorotheae (Weberl.) Christenh. & Byng,
collected. comb. nov. Basionym: Phyllactis dorotheae Weberl.,
Ber. Deutsch. Bot. Ges. 94: 306. 1981.
Valeriana atlantica (I.Richardson) Christenh. & Byng,
comb. nov. Basionym: Centranthus longiflorus Valeriana edwardsensis Christenh. & Byng, nom.
Steven subsp. atlanticus I.Richardson, Bot. J. Linn. nov. Replaced synonym: Valerianella texana Dyal,
Rhodora 40: 198. 1938, not: Valeriana texana

147
Steyerm. Etymology: Named for the Edwards nov. Basionym: Aligera jepsonii Suksd., Erythea 6:
Plateau in Texas, where this species occurs. 24. 1898.
Valeriana erikae (Graebn.) Christenh. & Byng, comb. Valeriana juncea (Boiss. & Heldr.) Christenh. & Byng,
nov. Basionym: Stangea erikae Graebn., Bot. Jahrb. comb. nov. Basionym: Centranthus junceus Boiss. &
Syst. 37: 450. 1906. Heldr., Diagn. Pl. Orient. 2: 73. 1849.
Valeriana eriocarpa (Desv.) Christenh. & Byng, comb. Valeriana kellereri (Stoj., Stef. & T.Georgiev) Christenh.
nov. Basionym: Valerianella eriocarpa Desv., J. Bot. & Byng, comb. nov. Basionym: Centranthus
(Paris) 2: 314. 1809. longiflorus var. kellereri Stoj., Stef. & T.Georgiev, Ber.
Valeriana erotica Christenh. & Byng, nom. nov. Bulg. Bot. Ges. 11: 37. 1928.
Replaced synonym: Centranthus longiflorus Steven, Valeriana kotschyi (Boiss.) Christenh. & Byng, comb.
Observ. Pl. Ross. 2: 76. 1824 [1828], not: Valeriana nov. Basionym: Valerianella kotschyi Boiss., Diagn.
longiflora Willk. Etymology: Named for Eros, Greek Pl. Orient. ser. 1, 3: 60. 1843
god of desire and erotic love. Valeriana lanata Christenh. & Byng, nom. nov.
Valeriana extincta Christenh. & Byng, nom. nov. Replaced synonym: Fedia lasiocarpa Steven, Mém.
Replaced synonym: Valerianella affinis Balf.f., Proc. Soc. Imp. Naturalistes Moscou 5: 50. 1817, not
Roy. Soc. Edinb. 11: 837. 1882, not: Valeriana affinis Valeriana lasiocarpa Griseb. Etymology: From Latin
Opiz. Etymology: This species is now extinct, but lanatus, woolly, for the woolly fruit.
used to occur only on Socotra Island. Valeriana langei Christenh. & Byng, nom. nov.
Valeriana florifera (Shinners) Christenh. & Byng, comb. Replaced synonym: Valerianella divaricata Lange,
nov. Basionym: Valerianella florifera Shinners, Field Vidensk. Meddel. Naturhist. Foren. Kjøbenhavn ser.
& Lab. 21(2): 93. 1953. 2, 1861 (vol. 23): 61. 1861, not: Valeriana divaricata
Valeriana gibbosa (Suksd.) Christenh. & Byng, comb. Hinter. Etymology: Named for Danish botanist Johan
nov. Basionym: Plectritis gibbosa Suksd., Werdenda M.C. Lange, who discovered and described this
1: 43. 1927. species from Grenada.
Valeriana graciliflora (Fisch. & C.A.Mey.) Byng & Valeriana lecoquii (Jord.) Christenh. & Byng, comb.
Christenh., comb. nov. Basionym: Fedia graciliflora nov. Basionym: Centranthus lecoqii Jord., Pugill. Pl.
Fisch. & C.A.Mey., Index Semin. (St. Petersburg) 6: Nov.: 76. 1852.
59. 1840. Valeriana macrocera (Torr. & A.Gray) Byng &
Valeriana heterocarpa (Pomel) Christenh. & Byng, Christenh., comb. nov. Basionym: Plectritis
comb. nov. Basionym: Fedia heterocarpa Pomel, macrocera Torr. & A.Gray, Fl. N. Amer. 2: 50. 1841.
Nouv. Mat. Fl. Atl. 1: 71. 1874. Valeriana macroptera (Suksd.) Byng & Christenh.,
Valeriana hidalgoana Christenh. & Byng, nom. nov. comb. nov. Basionym: Aligera macroptera Suksd.,
Replaced synonym: Aligera californica Suksd., Deutsche Bot. Monatsschr. 15: 146. 1897.
Werdenda 1: 44. 1927, not: Valeriana californica Valeriana maroccana (Rouy) Christenh. & Byng, comb.
A.Heller. Etymology: Named for Oriane Hidalgo, nov. Basionym: Centranthus maroccanus Rouy, Fl.
for her contribution to the understanding of France [Rouy & Foucaud] 8: 80. 1903.
relationships in Valeriana. Valeriana martini (Losco) Christenh. & Byng, comb.
Valeriana hirsuta (Emb. & Maire) Christenh. & Byng, nov. Basionym: Valerianella martini Losco, Trat. Pl.
comb. nov. Basionym: Fedia cornucopiae (L.) Gaertn. Arag. 1: 23. 1876.
var. hirsuta Emb. & Maire, Mém. Soc. Sci. Nat. Maroc Valeriana matheziana Christenh. & Byng, nom. nov.
21–22: 35. 1930. Replaced synonym: Fedia longiflora Torr. & A.Gray.,
Valeriana ibika Christenh. & Byng, nom. nov. Replaced Fl. N. Amer. 2: 51. 1841, not: Valeriana longiflora
synonym: Fedia puberula Bertol. ex Guss. Pl. Rar. Sic.: Willk. Etymology: Named for Joël Mathez, for his
15. 1826, not: Valeriana puberula Piper. Etymology: contribution to the knowledge of Fedia, now a
From Greek ηβικός, pubic. synonym of Valeriana.
Valeriana insignis (Suksd.) Christenh. & Byng, comb. Valeriana minor (Hook.) Byng & Christenh., comb. nov.
nov. Basionym: Aligera insignis Suksd., Deutsche Basionym: Plectritis congesta (Lindl.) DC. var. minor
Bot. Monatsschr. 15: 147. 1897. Hook., Fl. Bor.-Amer. 1: 291. 1833.
Valeriana ×interjecta Christenh. & Byng, nom. nov. Valeriana narda Christenh. & Byng, nom. nov.
Replaced synonym: Centranthus × intermedius Replaced synonym: Valerianella intermedia Dyal,
(Schltdl.) Rapin, Guide Bot. Vand ed. 2: 277. 1862, Rhodora 40: 202. 1938, not: Valeriana intermedia
not: Valeriana intermedia Vahl. Etymology: Latin for Vahl. Etymology: From Greek, νάρδος, a spikenard.
‘in between’. Valeriana nevadensis (Boiss.) Christenh. & Byng,
Valeriana jepsonii (Suksd.) Christenh. & Byng, comb. comb. nov. Basionym: Centranthus nevadensis

Boiss., Diagn. Pl. Orient. ser. 2, 2: 120. 1856. Centranthus sibthorpii Heldr. & Sartori ex Boiss.,
Valeriana nuttallii (Walp.) Christenh. & Byng, comb. Diagn. Pl. Orient. ser. 2, 2: 119. 1856.
nov. Basionym: Valerianella nuttallii Walp., Repert. Valeriana sieberi (Heldr.) Christenh. & Byng, comb.
Bot. Syst. 2: 527. 1843. nov. Basionym: Centranthus sieberi Heldr., Verh. K.K.
Valeriana obtusiloba (Boiss.) Christenh. & Byng, comb. Zool.-Bot. Ges. Wien 40: 295. 1890.
nov. Basionym: Valerianella obtusiloba Boiss., Valeriana stenocarpa (Krok) Christenh. & Byng, comb.
Diagn. Pl. Orient. ser. 1, 3: 59. 1843. nov. Basionym: Valerianella stenocarpa Krok, Kongl.
Valeriana oxyrhyncha (Fisch. & C.A.Mey.) Christenh. & Vetensk. Acad. Nya Handl. 5(1): 64. 1864.
Byng, comb. nov. Basionym: Valerianella oxyrhyncha Valeriana sulcata (Pomel) Christenh. & Byng, comb.
Fisch. & C.A.Mey., Index Semin. (St.Petersburg) 4: 51. nov. Basionym: Fedia sulcata Pomel, Nouv. Mat. Fl.
1837. Atl. 1: 72. 1874.
Valeriana ozarkana (Dyal) Christenh. & Byng, comb. Valeriana szovitsiana (Fisch. & C.A.Mey.) Christenh. &
nov. Basionym: Valerianella ozarkana Dyal, Rhodora Byng, comb. nov. Basionym: Valerianella szovitsiana
40: 208. 1938. Fisch. & C.A.Mey., Index Semin. (St. Petersburg) 3:
Valeriana pallescens (L.) Byng & Christenh., comb. 48. 1836.
nov. Basionym: Fedia cornucopiae (L.) Gaertn. var. Valeriana triplaris (Boiss. & Buhse) Christenh. & Byng,
pallescens Maire, Bull. Soc. Hist. Nat. Afrique N. 16: comb. nov. Basionym: Valerianella triplaris Boiss. &
34. 1925. Buhse, Nouv. Mém. Soc. Imp. Naturalistes Moscou
Valeriana paulae (Graebn.) Christenh. & Byng, comb. 12: 111. 1860.
nov. Basionym: Stangea paulae Graebn., Bot. Jahrb. Valeriana tuberculata (Boiss.) Christenh. & Byng,
Syst. 37: 450. 1906. comb. nov. Basionym: Valerianella tuberculata
Valeriana plagiostephana (Fisch. & C.A.Mey.) Boiss., Diagn. Pl. Orient. ser. 1, 3: 59. 1843.
Christenh. & Byng, comb. nov. Basionym: Valeriana turgida (Steven) Christenh. & Byng, comb.
Valerianella plagiostephana Fisch. & C.A. Mey., nov. Basionym: Fedia turgida Steven, Mém. Soc.
Index Semin. (St. Petersburg) 2: 52. 1835. Imp. Naturalistes Moscou 5: 345. 1817.
Valeriana pleurota Christenh. & Byng, nom. nov. Valeriana umbilicata (Sull.) Christenh. & Byng, comb.
Replaced synonym: Fedia costata Steven, Mém. nov. Basionym: Fedia umbilicata Sull., Amer. J. Sci.
Soc. Imp. Naturalistes Moscou 5: 344. 1817, not: Arts 42: 50. 1842.
Valeriana costata Schmale. Etymology: From Greek Valeriana valeria Byng & Christenh., nom. nov.
πλευρωτός, ribbed. Basionym: Valerianella anomala A.Gray, Proc.
Valeriana pontica (Velen.) Christenh. & Byng, comb. Amer. Acad. Arts & Sci. 19: 83. 1883, not: Valeriana
nov. Basionym: Valerianella pontica Velen., anomala Eastw. Etymology: Derived from Latin
Sitzungsber. Königl. Böhm. Ges. Wiss. 37: 34. 1893 valere, to force or to be healthy or strong.
(1894). Valeriana valerianella Christenh. & Byng, nom. nov.
Valeriana pulvinata (Rauh & Willer) Christenh. & Replaced synonym: Valeriana locusta L. var. radiata
Byng, comb. nov. Basionym: Phyllactis pulvinata L., Sp. Pl. 1: 34. 1753, not: Valeriana radiata Willd.
Rauh & Willer, Bot. Jahrb. Syst. 82: 268. 1963. Etymolgy: Named for the former genus Valerianella,
Valeriana rimosa (Bastard) Christenh. & Byng, comb. meaning a small Valeriana.
nov. Basionym: Valerianella rimosa Bastard, J. Bot. Valeriana wandae (Graebn. & Tessend.) Christenh.
(Paris) 3: 20. 1814. & Byng, comb. nov. Basionym: Stangea wandae
Valeriana rubens (Suksd.) Christenh. & Byng, comb. Graebn. & Tessend., Bot. Jahrb. Syst. 37: 450. 1906.
nov. Basionym: Aligera rubens Suksd., Deutsche Bot. Valeriana woodsiana (Torr. & A.Gray) Christenh. &
Monatsschr. 15: 146. 1897. Byng, comb. nov. Basionym: Fedia woodsiana Torr.
Valeriana salata Christenh. & Byng, nom. nov. & A.Gray, Fl. N. Amer. 2: 52. 1841.
Replaced synonym: Valerianella microcarpa Loisel.,
Not. Fl. France: 151. 1810, not: Valeriana microcarpa Acknowledgements
Sumnev. Etymology: From Greek σαλάτα, salad,
because the leaves can be eaten as salad. We thank the curators of A, BM, GH, K, L, P and U
Valeriana sclerocarpa (Fisch. & C.A.Mey.) Christenh. & for access to herbarium collections and literature
Byng, comb. nov. Basionym: Valerianella sclerocarpa whilst preparing this manuscript. JWB would like to
Fisch. & C.A.Mey., Index Semin. (St.Petersburg) 2: thank Harvard University Herbarium and the Alberta
53. 1835. Mennega Stichting foundation for travel grants to visit
A, GH and P herbaria. EB is grateful to Didier Geffard-
Valeriana sibthorpii (Heldr. & Sartori ex Boiss.)
Kuriyama from UMS 2700 for help taking images of
Oxygyne confusa under the microscope.

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© 2018 Plant Gateway Ltd.

The Global Flora ISSN 2398-6336 eISSN 2398-6344

i Published online 9 February 2018

THE GLOBAL FLORA

GLOVAP Nomenclature Part 1

© Plant Gateway 2018

British Library Cataloguing in Publication data

For information or to purchase other Plant Gateway titles please visit

Maarten J.M. Christenhusz

Ehoarn Bidault (Burmanniaceae)

John E. Burrows (Myrtaceae)

Florian Jabbour (Ranunculaceae)

Vincent S.F.T. Merckx (Burmanniaceae, Triuridaceae)

Greg Stull (Icacinaceae)

Background any of the standard databases cited above and hence

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2 Published online 9 February 2018

taxonomic changes to reflect a monophyletic group, Taxonomic list

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4 Published online 9 February 2018

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6 Published online 9 February 2018

Alsophila valdesquamata (Janssen & Rakotondr.) F21. Dennstaedtiaceae

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8 Published online 9 February 2018

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10 Published online 9 February 2018

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12 Published online 9 February 2018

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22 Published online 9 February 2018

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24 Published online 9 February 2018

Athyrium philippinense (M.Kato) Christenh., comb. 234. 1862.

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26 Published online 9 February 2018

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28 Published online 9 February 2018

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30 Published online 9 February 2018

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32 Published online 9 February 2018

Thelypteris lawakii (Holttum) Christenh., comb. Thelypteris marojejyensis (Rakotondr.) Christenh.,

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34 Published online 9 February 2018

Thelypteris novae-brittaniae (Holttum) Christenh., Thelypteris paraochthodes (Ching ex K.H.Shing &

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36 Published online 9 February 2018

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38 Published online 9 February 2018

Thelypteris sumbawensis (C.Chr.) Christenh., comb. Blumea 22: 240. 1975.

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Basionym: Lellingeria affinis Labiak, Fl. Neotrop. Hook.

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