Two new genera of Sapindaceae (Cupanieae) from the southern Pacific: Lepidocupania and Neoarytera

Abstract

BUERKI, S., J. MUNZINGER, P.P. LOWRY II & M.W. CALLMANDER (2020). Two new genera of Sapindaceae (Cupanieae) from the southern Pacific: Lepidocupania and Neoarytera. Candollea 75: 269-284. In English, English abstract. DOI: http://dx.doi.org/10.15553/c2020v752a9 Phylogenetic analyses of the family Sapindaceae inferred from nuclear and plastid sequence data have revealed a high level of para-and polyphyly at the subfamilial, tribal, and generic levels. A phylogenetic study focusing on taxa in the southern Pacific belonging to tribe Cupanieae has shown that the two most species-rich genera, Arytera Blume and Cupaniopsis Radlk., are polyphyletic. This study aims to clarify generic limits among the taxa currently placed in these two genera by identifying morphological features that support monophyletic groups suitable for recognition at the generic level. Specimens deposited in major herbaria holding material of these taxa were examined to complement extensive field observations. Careful consideration of morphological features in light of previous taxonomic treatments and the results of phylogenetic analyses enabled us to propose a realigned generic framework for Cupanieae in which two new genera are described to accommodate species previously placed in Arytera and Cupaniopsis: viz., Lepidocupania Buerki, Callm., Munzinger & Lowry (21 species) and Neoarytera Callm., Buerki, Munzinger & Lowry (4 species). A total of 25 new combinations are made, lectotypes are designated for nine names (two first step and seven second-step), and one new synonym is established. A key to the newly circumscribed genera Arytera and Cupaniopsis, along with allied genera, is provided, accompanied by information on the distribution and ecology of each species.

Full text

© CONSERVATOIRE ET JARDIN BOTANIQUES DE GENÈVE 
Addresses of the authors:
SB: Department of Biological Sciences, Boise State University,  University Drive, Boise, ID , USA.
E-mail: svenbuerki@boisestate.edu
JM: AMAP, Université Montpellier, IRD, CIRAD, CNRS, INRA, Montpellier, France.
PPL: Missouri Botanical Garden,  Shaw Blvd., St. Louis, Missouri , USA; Institut de Systématique, Évolution,
et Biodiversité (ISYEB), Muséum National d’Histoire Naturelle, Centre National de la Recherche Scientique, Sorbonne
Université, École Pratique des Hautes Études, Université des Antilles, C.P. ,  rue Cuvier,  Paris, France.
MWC: Conservatoire et Jardin botaniques de la Ville de Genève, ch. de l’Impératrice , C.P. ,  Chambésy, Switzerland.
Submitted on June , . Accepted on August , . First published online on October , .
ISSN : - – Online ISSN : - – Candollea () :  –  ()
Two new genera of Sapindaceae
(Cupanieae) from the southern Pacific:
Lepidocupania and Neoarytera
Sven Buerki, Jérôme Munzinger, Porter P. Lowry II & Martin W. Callmander
Abstract
BUERKI, S., J. MUNZINGER, P.P. LOWRY II & M.W. CALLMANDER (). Two new genera of Sapindaceae (Cupanieae) from the southern
Pacic: Lepidocupania and Neoarytera. Candollea : 
– . In English, English abstract. DOI: http://dx.doi.org/./cva
Phylogenetic analyses of the family Sapindaceae inferred from nuclear and plastid sequence data have revealed a high level
of para- and polyphyly at the subfamilial, tribal, and generic levels. A phylogenetic study focusing on taxa in the southern
Pacic belonging to tribe Cupanieae has shown that the two most species-rich genera, Arytera Blume and Cupaniopsis
Radlk., are polyphyletic. is study aims to clarify generic limits among the taxa currently placed in these two genera
by identifying morphological features that support monophyletic groups suitable for recognition at the generic level.
Specimens deposited in major herbaria holding material of these taxa were examined to complement extensive eld
observations. Careful consideration of morphological features in light of previous taxonomic treatments and the results
of phylogenetic analyses enabled us to propose a re-aligned generic framework for Cupanieae in which two new genera
are described to accommodate species previously placed in Arytera and Cupaniopsis: viz., Lepidocupania Buerki, Callm.,
Munzinger & Lowry ( species) and Neoarytera Callm., Buerki, Munzinger & Lowry ( species). A total of  new
combinations are made, lectotypes are designated for nine names (two rst step and seven second-step), and one new
synonym is established. A key to the newly circumscribed genera Arytera and Cupaniopsis, along with allied genera, is
provided, accompanied by information on the distribution and ecology of each species.
Keywords
SAPINDACEAE – Cupanieae – Arytera – Cupaniopsis – Lepiderema – Lepidocupania – Neoarytera – Synima – Australia
– New Caledonia – New genus – New combination – Typication

Introduction
Phylogenetic analyses of the family Sapindaceae inferred from
nuclear and plastid sequence data have revealed a high level of
para- and polyphyly at the subfamilial, tribal, and generic levels
(H et al., ; B et al., , ). A new
informal infra-familial classication was proposed by B
et al. (), mainly based on molecular data, to accommo-
date these ndings and to provide a framework for developing
improved generic circumscriptions that meet the criterion of
monophyly. Within Sapindaceae, the Cupania group, which
occurs in Australasia, Asia, South America, and Madagas-
car, but is absent in continental Africa, corresponds to the
largest radiation in terms of the number of genera and species
(B et al., ). A forthcoming genus-level phylogeny
of the family based on  nuclear genes will provide the basis
for a new, formal tribal delimitation, in which the Cupania
group corresponds to tribe Cupanieae Radlk. (Buerki et al.,
unpubl. data). To date, the only study that has used a phy-
logenetic framework to test the monophyly of genera in the
Cupania group is that of B et al. (), which focused
on taxa in the islands of the southern Pacic. is study dem-
onstrated that the two most species-rich genera, Arytera Radlk.
and Cupaniopsis Radlk., were both polyphyletic. Within the
Cupania group, they were placed in clade B (see B et al.,
 for more details on the composition of this clade) and
their members were distributed among two subclades (B-III
and B-VI), within which species of both genera were inter-
mixed (see Fig.). Lepiderema Radlk. was shown to be sister
to species of Arytera and Cupaniopsis in clade B-III, whereas
within clade B-IV, Synima Radlk. was inferred to be sister to
other taxa currently placed in Arytera and Cupaniopsis.
Arytera was described by B (), who recog-
nized two species, A. litoralis Blume (designated as the
lectotype of the genus by R, a) (Fig.A) and
A.montana Blume (later transferred to Lepidopetalum Radlk.
by R, a). e rst comprehensive account of the
genus included  species (R, 
– ), and more
recently T () published a monograph in which he
recognized  species (Fig.A
– B). e monophyly of Arytera
was questioned by T (), who established the genus
Mischarytera (Radlk.) H. Turner to accommodate the species
previously placed by R (b) in Arytera sect.
Mischarytera Radlk. T () also divided Arytera sect.
Azarytera Radlk. into two subsections, Arytera subsect. Pacica
H. Turner and Arytera subsect. Distylis H. Turner.
Cupaniopsis was described by R (b) to accom-
modate species originally placed in various other genera, includ-
ing taxa such as C. anacardioides (A. Rich.) Radlk. ( Cupania
anacardioides A. Rich), which was designated as lectotype of
Cupaniopsis by R (). e rst account of the genus
included  species (R, 
– ) (Fig.C – D),
whereas A () recognized  species in his monograph
of Cupaniopsis, ranging from East Malesia to Australia and
several South Pacic islands (Fiji, New Caledonia, and Vanuatu),
with centers of diversity in Australia, New Guinea, and New
Caledonia. R (b) described two sections within
Cupaniopsis, in addition to the nominal section, Cupaniopsis sect.
Mizopetalum Radlk. and Cupaniopsis sect. Macropetalum Radlk.
Neither of these two sections were recognized by A (),
even though the  species of Cupaniopsis sect. Mizopetalum
formed a clade in the cladistic analysis he performed based on
morphological characters (see A, : 
– , g.).
Among the allied genera belonging to the Cupania group,
Lepiderema Radlk. was established by R (b), with
its type being L. papuana Radlk. As currently circumscribed, this
genus includes eight species, six endemic to Australia and two to
New Guinea (R, ; S, ) (Fig F). A second
member of the Cupania group, Synima Radlk., has four species,
three in Australia and one in New Guinea (R, b;
F, ; C et al., ) (Fig.E).
Several authors have questioned the monophyly of Arytera.
R (
– ) commented on the morphological
similarities between species of Arytera sect. Azarytera and
Cupaniopsis sect. Mizopetalum as follows: “Among the Asian
and Oceanic Cupanieae possessing the calyx of Matayba and
only small un-crested petals with  scales or nearly lacking
scales, the genus [Arytera] is remarkable by its capsule [thatis]
270 – Two new genera of Sapindaceae (Cupanieae) from the southern Pacific Candollea 75, 2020
Lepiderema hirsuta
Lepiderema pulchella
Synima serrata
Cupaniopsis agelliformis
0.82
1.00
1.00
1.00
0.71
0.59
1.00
0.94
0.99
0.99
0.98
1.00
1.00
1.00
1.00
1.00
0.90
0.76
0.65
0.95
1.00
1.00
1.00
1.00
0.95
0.99
0.98
0.84
0.99
1.00
0.76
1.00
0.65
1.00
1.00
0.001
Matayba apetala
Cupaniopsis globosa
Cupaniopsis globosa
Cupaniopsis globosa
Cupaniopsis myrmectona
Cupaniopsis myrmectona
Arytera aff. gracilipes
Arytera aff. gracilipes
Arytera lepidota
Cupaniopsis oedipoda
Cupaniopsis fruticosa
Cupaniopsis subfalcata
Arytera collina
Cupaniopsis sylvatica
Cupaniopsis apiocarpa
Cupaniopsis petiolulata
Cupaniopsis petiolulata
Cupaniopsis phalacrocarpa
Cupaniopsis megalocarpa
Cupaniopsis megalocarpa
Cupaniopsis macrocarpa
Cupaniopsis macrocarpa var. polyphylla
Cupaniopsis mackeeana
Cupaniopsis sylvatica
Cupaniopsis petiolulata
BIII
Arytera s.str.
Cupaniopsis s.str.
Cupaniopsis
sect. Mizopetalum
Arytera
sect. Azarytera
Arytera
subsect. Pacica
Cupaniopsis glomeriora
Cupaniopsis glomeriora
Cupaniopsis oedipoda
Cupaniopsis anacardioides
1.00 Arytera chartacea
Arytera divaricata
Arytera littoralis
Arytera microphylla
Arytera littoralis
Arytera neoebudensis
Synima macrophylla
Arytera brackenridgei
Arytera brackenridgei
Matayba domingensis
Cupaniopsis oedipoda
BIV
Fig.1.
–
MrBayes Bayesian halfcompat consensus tree inferred
from eight nuclear and plastid DNA regions adapted from B
UERKI
et
al. (2012) showing relationships within the Cupania group. Bayesian
posterior probability (BPP) support values are indicated. Shades of
grey highlight the polyphyly of Arytera Blume and Cupaniopsis Radlk.

Candollea 75, 2020 Two new genera of Sapindaceae (Cupanieae) from the southern Pacific – 271
Fig.2.
–
A. Arytera litoralis Blume (Halmahera, North Maluku, Indonesia); B. Arytera divaricata F. Muell. (Queensland, Australia);
C. Cupaniopsis macrocarpa var. polyphylla Adema (New Caledonia); D. Cupaniopsis flagelliformis (F.M. Bailey) Radlk. (Queensland, Australia);
E.Synima serrata (S.T. Reynolds) Callm. & Buerki (Queensland, Australia); F. Lepiderema punctulata (F. Muell.) Radlk. (Queensland, Australia).
[A: Bangun 440; B: Gray et al. 9741; C: Lowry et al. 7319; D: Gray et al. 9693; E: Gray et al. 9856] [Photos: A: I.A. Haris; B, D: B. Gray;
C: P. Lowry; E: S. & A. Pearson; F: G. Sankowsky]
C
A B
D
FE

more or less divaricately lobed with cocci not at all winged, or
somewhat compressed-obovate [and] usually sessile […], its
unappendaged aril usually completely enclosing the seed, the
compact structure of its leaves and in the species of section IV
[Arytera sect. Azarytera] by the lepidote leaets, by which char-
acter this section agrees with the genus Cupaniopsis section III
Mizopetalum […].” (R, 
– : ; translated
from Latin by R. Gereau). T (: ) mentioned that
Arytera, as he circumscribed it, is only recognizable based “on
a polythetic set of [six] character states”, and excluded three
species (A. bullata H. Turner, A. lautereriana (F.M. Bailey)
Radlk., and A. macrobotrys (Merr. & L.M. Perry) R.W. Ham),
which he transferred to his new genus Mischarytera (see above).
In light of the long-recognized issues regarding the delimi-
tation of Arytera and Cupaniopsis, coupled with the recent
molecular phylogenetic analyses that have clearly shown both
of them to be polyphyletic, as currently circumscribed, a thor-
ough review of both genera seems appropriate. In the present
study, we aim to revise generic limits so that they correspond
to monophyletic groups, and to place all of the taxa currently
assigned to Arytera and Cupaniopsis within this new generic
framework, supported by morphological features that charac-
terize the clades dened by B et al. (). Taxa belong-
ing to the phylogenetically closely related genera Lepiderema
and Synima have also been included in an eort to clarify their
relationships and taxonomic identities. Our goal is to resolve
generic delimitations within this challenging group and thereby
further Radlkofer’s quest, which started nearly  years ago.
Material & Methods
In order to identify potentially informative morphological char-
acters that support monophyletic groups and to assess ani-
ties among the taxa currently placed in Arytera, Cupaniopsis,
Lepiderema, and Synima, we examined material from the fol-
lowing herbaria: BM, BRI, CNS, G, K, L, MO, MPU, NOU, P,
SING, and SUVA. is was complemented by accessing digital
images of type specimens at other herbaria through the Global
Plants website [https://plants.jstor.org]. To encompass the full
range of morphological variation within species across their
distributional ranges, we also consulted the descriptions pro-
vided in regional oras (i.e. Flora Malesiana, A et al., ;
Flora of Australia, R, b) and generic monographs
(i.e. Cupaniopsis, A, ; Arytera, T, ), as well
as the seminal works of R (a, b, 
– ).
Data were compiled on key morphological characters for each
taxon. Finally, between  and , we also conducted eld-
work throughout much of the range of the study group, includ-
ing in Australia, Borneo, the Fijian archipelago, the Moluccas,
New Caledonia, Peninsular Malaysia, Singapore, and Vanuatu,
in order to observe and collect material of indigenous species,
and to gain insights into their morphology and ecology, as well
as their relationships with other members of the family.
Phylogenetic results and morphological
characters
A key step toward resolving generic delimitations within the
closely related Sapindaceae that were the focus of this study
involved aligning our morphological ndings with previous
taxonomies and discussing the results in light of the phylo-
genetic framework presented in B et al. (). Bioge-
ography was also taken into consideration as a criterion for
delimiting genera. The presentation of our findings given
below follows the sequence of clades presented in B et
al. (). We also have sought to corroborate our taxonomic
ndings by expanding phylogenetic analyses to include DNA
sequences from several key species, although the results of this
work are not formally included in the present paper and will
instead be the subject of a forthcoming publication focusing
on the evolution and biogeography of the Cupania clade in
the Pacic islands and neighboring regions. Our goal here is
to provide the new generic classication required as a basis for
conducting evolutionary and biogeographical analyses.
Clade B-III identified by B et al. () (Fig.)
contains all sampled taxa of Lepiderema, which form a
subclade that is sister to another subclade comprising all
sequenced species of Cupaniopsis sect. Mizopetalum (including
species, ve of which were originally placed in this section
by Radlkofer and the rest were subsequently added by A
() in his treatment of the genus). In this treatment, we are
only recognizing  of these species (see taxonomic treatment
below), together with all four sampled species of Arytera sect.
Azarytera. Although the sampling from these groups used by
B et al. () was limited, their phylogenetic results
strongly suggested that species of Arytera sect. Azarytera were
nested within Cupaniopsis sect. Mizopetalum, which precluded
the possibility of recognizing either of these infrageneric taxa
as a separate genus. The presence of lepidote scales on the
vegetative and reproductive organs of these species represents
a clear morphological synapomorphy for clade B-III. Two
taxonomic interpretations are possible given this phyloge-
netic context: ) recognize one large genus encompassing
all the members of Lepiderema, Arytera sect. Azarytera, and
Cupaniopsis sect. Mizopetalum; or ) treat Lepiderema as one
genus and place the remaining species in another genus. We
prefer the second option because species of Lepiderema can
easily be distinguished from the other members of clade B-III
by their lack of petal and ovary scales (vs. petal and ovary scales
present in the other taxa). Moreover, Lepiderema is restricted
to Australia and New Guinea, whereas the other taxa occur
in New Caledonia, Fiji and Samoa (with the exception of
A. brackenridgei Radlk., which has a wide distribution in the
272 – Two new genera of Sapindaceae (Cupanieae) from the southern Pacific Candollea 75, 2020

Solomon Islands, Vanuatu, Fiji, Wallis and Futuna (Horn
Islands), Tonga and Samoa). In order to accommodate the
group that is sister to Lepiderema, which comprises the species
previously placed in Arytera sect. Azarytera and Cupaniopsis
sect. Mizopetalum, a new genus is required, which we described
below as Lepidocupania Buerki, Callm., Munzinger & Lowry.
e taxa belonging to clade B-IV of B et al. ()
(Fig.) can easily be distinguished from those in clade B-III by
the absence of lepidote scales on their vegetative and reproduc-
tive organs. Synima is inferred to occupy a basal position in clade
B-IV. is genus is restricted to Australia, with the exception of
a single species in New Guinea, viz. S. cordieri (F. Muell.) Radlk.
(R, b). Synima is characterized by having crested
scales on its petals and seeds, the latter fully covered with a sar-
cotesta (R, a; F, ; C et al.,
). e remainder of clade B-IV comprises three subclades,
which are fully aligned with previously recognized taxonomic
entities. e species of Arytera subsect. Pacica correspond to
the rst subclade, which is sister to the two other subclades, one
containing the type of the genus (A. litoralis) and all species cur-
rently assigned to Arytera subsect. Arytera and Arytera subsect.
Distylis, and a third subclade comprising the type of Cupaniopsis
(C. anacardioides) and all species currently placed in this genus,
with the exception of those belonging to Cupaniopsis sect.
Mizopetalum (see above). Since each of these three subclades
is consistent with a currently recognized taxonomic entity and
is also morphologically coherent, we have opted to recognize
them as a separate, well-dened genus. Two of the subclades
correspond to Arytera and Cupaniopsis because they contain
their respective types, whereas the third subclade represents
a new genus, which we formally describe here as Neoarytera
Callm., Buerki, Munzinger & Lowry.
Below we provide a key to the genera, which includes the
two new genera, and we also include an appendix that presents
a synopsis of currently accepted species of Arytera, Cupaniopsis,
Lepidocupania, and Neoarytera, with their respective distribu-
tions. e taxonomy of the two new genera follows the com-
prehensive monographs by A () and T (),
with the exception of Cupaniopsis rotundifolia Adema, which
is not accepted here. e synonymies proposed by Adema and
Turner are not repeated here except for two names that require
nomenclatural clarification: Arytera pachyphylla Radlk. and
Cupaniopsis ganophloea Radlk.
A key to the newly circumscribed Arytera,
Cupaniopsis and allied genera
e following key is adapted from R (b).
. Calyx lobes free (sometimes sepals basally united), orbicu-
lar, elliptic or obovate, concave .................................... 
a. Calyx shortly cupular, the lobes toothed or partite, usually
ovate ............................................................................ 
. Scales absent from petals ............................ Lepiderema
a. Scales present on petals ............................................... 
. Scales present on vegetative and fertile organs ...............
............................................................... Lepidocupania
a. Scales absent from vegetative and fertile organs .............
................................................................... Cupaniopsis
. Petal scales crested ............................................. Synima
a. Petal scales not crested ................................................ 
. Petal scales adnate to the petal margin or free from the
petals; central axis of fruit not thickened .......... Arytera
a. Petal scales comprising minute enations; central axis of
fruit distinctly thickened ............................. Neoarytera
Taxonomy
Description and synopsis of Lepidocupania
Lepidocupania Buerki, Callm., Munzinger & Lowry, gen. nov.
(Fig.A
– C).
Type: Lepidocupania lepidota (Radlk.) Buerki, Callm.,
Munzinger & Lowry ( Arytera lepidota Radlk.).
= Cupaniopsis sect. Mizopetalum Radlk. in Sitzungs-
ber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss.
München : . . Type: Cupaniopsis fruticosa
Radlk. (Lepidocupania fruticosa (Radlk.) Buerki,
Callm., Munzinger & Lowry) (lectotype designated
by A, : ).
= Arytera sect. Azarytera Radlk. in Sitzungsber. Math.-
Phys. Cl. Königl. Bayer. Akad. Wiss. München : .
. Type: Arytera arcuata Radlk. ( Lepidocupania
arcuata (Radlk.) Buerki, Callm., Munzinger & Lowry)
(lectotype designated by T, : ).
Trees or shrubs. Indument of short, straight, patent or
appressed trichomes; glandular scales present on vegetative
parts, inorescence axes, pedicels, abaxial surface of calyx, pistil,
and fruits; buds “varnished”. Leaves alternate, 
– -jugate;
leaflets opposite to alternate, subsessile to petiolulate,
margin entire to coarsely dentate. Inorescences axillary or
pseudo-terminal. Flowers zygomorphic or actinomorphic
(in L.arcuata, L. brackenridgei, L. gracilipes, and L. lepidota),
functionally unisexual; sepals (
–)(– ), free and imbri-
cate or united (in L. arcuata, L. brackenridgei, L. gracilipes,
and L. lepidota) to form a dentate calyx cup; petals  ( in
L.glomeriora), with  distinct scales; disc lobed or not (in
L.arcuata, L. brackenridgei, L. gracilipes, and L. lepidota), rim
glabrous to pilose; stamens (
–)
– , anthers basixed; ovary

– -locular. Fruit a capsule, with 
–  well developed lobes,
rarely  (in L. concolor, L.guillauminii, and L. samoensis),
Candollea 75, 2020 Two new genera of Sapindaceae (Cupanieae) from the southern Pacific – 273

dehiscence loculicidal, glabrous to puberulous and rugose to
verrucose outside, glabrous to pilose inside; seed ellipsoid or
ovoid to globose, sarcotesta covering half to all of the seed,
esh-membranaceous.
Distribution. – Lepidocupania comprises  species occur-
ring in the Caroline Islands, Fiji, New Caledonia, Samoa, the
Solomon Islands, and Vanuatu (Fig.).
Notes. – Lepidocupania shares the presence of lepidote scales
on its vegetative and reproductive organs with Lepiderema.
However, Lepidocupania can easily be distinguished from
Lepiderema by the presence (vs. absence) of petal and ovary
scales, and it differs from Cupaniopsis by the presence (vs.
absence) of glandular scales on its vegetative and fertile organs.
Lepidocupania arcuata (Radlk.) Buerki, Callm., Munzinger
& Lowr y, comb. nov.
 Arytera arcuata Radlk. in Sitzungsber. Math.-Phys. CI.
Konigl. Bayer. Akad. Wiss. Munich : . .
Lectotypus (designated by T, : ): New
Caledonia. Prov. Sud: Nouméa, X., Balansa
150 (M[M]!; isolecto-: FI [FI]!, NY
[NY, NY]!, P [P, P]!).
Distribution and ecology. – According to T’s ()
concept, Lepidocupania arcuata is endemic to the New Caledo-
nian archipelago, where it occurs from sea level to m, pre-
dominantly on calcareous soils (Loyalty Islands, Ile des Pins),
but also on sand, clay and schist. It is found in mesophyll and
sclerophyll forest and scrub.
Notes. – Additional taxonomic analyses are required, espe-
cially with regard to material from the Loyalty Islands, which
exhibits very peculiar indument compared to that from the
main island of New Caledonia [Grande Terre]. Moreover,
T () tentatively identied a specimen from Tonga
(Parks 16317: L) as L. arcuata, which, if conrmed,
would significantly expand the geographic range of this
species.
Lepidocupania brackenridgei (A. Gray) Buerki, Callm., Mun-
zinger & Lowry, comb. nov. (Fig.B).
 Cupania brackenridgei A. Gray in Wilkes, U.S. Expl.
Exped., Phan. : . .  Arytera brackenridgei
(A.Gray) Radlk. in Sitzungsber. Math.-Phys. CI.
Konigl. Bayer. Akad. Wiss. Munich : . .
Holotypus: Fiji: Ovalau, 
– , Wilkes s.n.
(US[US] image seen; iso-: P [P]!).
Distribution and ecology. – Lepidocupania brackenridgei is
widespread in the Solomon Islands, Vanuatu, Fiji, Wallis and
Futuna (Horn Islands), Tonga and Samoa, where it occurs
from sea level to m (A, ). is common species
occurs in primary and secondary rainforest, but is also found
in savannah; it grows on limestone and lava elds.
Lepidocupania concolor (Gillespie) Buerki, Callm., Munzinger
& Lowr y, comb. nov.
 Guioa concolor Gillespie in Bull. Bernice P. Bishop Mus.
: . .  Arytera concolor (Gillespie) A.C. Sm. in
J. Arnold Arbor. : . .  Cupaniopsis concolor
(Gillespie) R.W. Ham in Blumea : . .
Holotypus: Fiji: Taveuni, vicinity of Waiyevo, .III.
Gillespie 4794 (BISH [BISH] image seen; iso-:
BISH [BISH, BISH] images seen,
K[K]!, NY [NY]!, GH [GH]
image seen, US [US] image seen).
Distribution and ecology. – Lepidocupania concolor is endemic
to Fiji, where it is known from the three main islands of Viti
Levu, Vanua Levu, and Taveuni (S, ).
Lepidocupania fruticosa (Radlk.) Buerki, Callm., Munzinger
& Lowr y, comb. nov.
 Cupaniopsis fruticosa Radlk. in Sitzungsber. Math.-Phys.
CI. Konigl. Bayer. Akad. Wiss. Munich : . .
Lectotypus (designated by A, : ): Ne w
Caledonia: sine loco, s.d., Pancher 142 (M [M]
image seen; iso-: MEL [MEL] image seen,
P[P, P, P, P]!).
Distribution and ecology. – Lepidocupania fruticosa is
endemic to the southwestern part of Grande Terre, where
it grows in dense humid forest, usually on serpentinite, but
sometimes also on schist or laterite, from  to (– ) m
(A, ).
Notes. – Based on our current knowledge on this species,
additional taxonomic studies appear to be needed to clarify its
circumscription. is should also include material currently
assigned to L. subfalcata and L. tontoutensis (see below for
additional details).
Lepidocupania glabra (Adema) Buerki, Callm., Munzinger &
Lowr y, comb. nov.
 Cupaniopsis glabra Adema in Leiden Bot. Ser. : .
.
Holotypus: New Caledonia. Prov. Sud: Basse Ton-
touta, rive gauche, terrain serpentineux, m, .II.,
274 – Two new genera of Sapindaceae (Cupanieae) from the southern Pacific Candollea 75, 2020

Candollea 75, 2020 Two new genera of Sapindaceae (Cupanieae) from the southern Pacific – 275
Fig.3.
–
A. Lepidocupania glomeriflora (Radlk.) Buerki, Callm., Munzinger & Lowry (New Caledonia); B. Lepidocupania brackenridgei (A. Gray)
Buerki, Callm., Munzinger & Lowry (Vanua Levu, Fiji); C. Lepidocupania lepidota (Radlk.) Buerki, Callm., Munzinger & Lowry (New Caledonia);
D.Neoarytera collina (Pancher & Sebert) Callm., Buerki, Munzinger & Lowry (New Caledonia); E. Neoarytera neoebudensis (Guillaumin) Callm.,
Buerki, Munzinger & Lowry (New Caledonia); F. Neoarytera chartacea (Radlk.) Callm., Buerki, Munzinger & Lowry (New Caledonia).
[A: Callmander et 788; B: Munzinger 379; C: Munzinger 7700; D: Munzinger 7395; E: Munzinger 7404; F: Hequet 3525]
[Photos: A
–
B, D
–
E: P. Lowry; C: J. Munzinger; F: V. Hequet]
C
A B
D
FE

MacKee 40234 (L [L]!; iso-: NOU [NOU]!,
P [P]!).
Distribution and ecology. – Lepidocupania glabra is endemic
to the Tontouta River valley (Grande Terre). It is found in
maquis and continuous tall maquis (“maquis paraforestier”,
see MC et al.,  for details) dominated by Gymnostoma
chamaecyparis ( J. Poiss.) L.A.S. Johnson (Casuarinaceae) on
ultramafic substrate, primarily alluvium, between  and
m (A, ; D'A, ).
Notes. – is species has been assigned an IUCN risk of
extinction status of “Critically Endangered” [CR] based on its
narrow distribution and reduction in population size (IUCN,
).
Lepidocupania globosa (Adema) Buerki, Callm., Munzinger
& Lowr y, comb. nov.
 Cupaniopsis globosa Adema in Leiden Bot. Ser. : .
.
Holotypus: New Caledonia. Prov. Sud: Bourail, les
Montagnes Blanches, .X., Suprin 2080 (P not found;
iso-: L [L]!, NOU [NOU]!).
Distribution and ecology. – Lepidocupania globosa is endemic
to the west coast of Grande Terre, where it is restricted to
sclerophyll forest (B et al., ).
Notes. – is species has been assigned an IUCN risk of
extinction status of “Vulnerable” (IUCN, ). See note
under L. pennelii.
Lepidocupania glomeriora (Radlk.) Buerki, Callm., Munz-
inger & Lowry, comb. nov. (Fig.A).
 Cupaniopsis glomeriora Radlk. in Sitzungsber. Math.-
Phys. CI. Konigl. Bayer. Akad. Wiss. Munich : .
.
Lectotypus (first step designated by A, :
; second step designated here): New Caledonia.
Prov. Sud: Bourail, dans les bois, III., Balansa 1447
(P[P]!; isolecto-: P [P, P]!).
Distribution and ecology. – Lepidocupania glomeriflora
is endemic to New Caledonia, where it occurs on Grande
Terre, the Ile des Pins, and the three larger Loyalty Islands
(A, ; B, ). It is found in maquis vegeta-
tion, dry to mesic forests, or forest remnants, along the coast,
on hills or mountainsides, often along rivers, and occurs on
calcareous substrates, serpentinite and schist, but appears not
to favor peridotitic substrates and is completely absent from
Grande Terre’s large southern ultramac massif. Lepidocupania
glomeriora has been recorded from  to m.
Notes. – Nine syntype collections were cited in the protologue
of Cupaniopsis glomeriflora: Balansa 153 p.p., 1447, Baudouin
354 p.p., “Culta in hort. Paris”, Deplanche 83, Labillardière 169,
Pancher 782, Vieillard 228, and 233. A (: ) desig-
nated Balansa 1447 as the lectotype. Original material at P is,
however, mounted on three sheets, necessitating the second step
lectotypication designated here, for which we have selected the
most complete and best-preserved sheet [P].
Lepidocupania gracilipes (Radlk.) Buerki, Callm., Munzinger
& Lowr y, comb. nov.
 Arytera gracilipes Radlk. in Repert. Spec. Nov. Regni
Veg. : . .
Lectotypus (designated by T, : ): New
Caledonia. Prov. Nord: “montagnes de Panloïtch, près
Gatop”, s.d., Vieillard 2403 (K [K]!; isolecto-:
FI [FI]!, NY [NY, NY]!,
M[M] image seen, P [P, P,
P, P, P, P,
P]!).
Distribution and ecology. – Lepidocupania gracilipes is
endemic to Grande Terre, where it grows in gallery forest,
continuous tall maquis (“maquis paraforestier”, see MC et
al.,  for details), and thickets on (rocky) serpentine terrain,
and sometimes along streams on alluvium (T, ). It
seems to be restricted to serpentine, hyper-magnesium brown
soils. e species has been recorded from sea level to m.
Lepidocupania grandiora (Adema) Buerki, Callm., Munz-
inger & Lowry, comb. nov.
 Cupaniopsis grandiora Adema in Bull. Mus. Natl. Hist.
Nat., B, Adansonia, : . .
Holotypus: New Caledonia. Prov. Sud: Mont Do,
m, s.d., McPherson 3805 (MO [MO]!; iso-:
NOU [NOU]!).
Distribution and ecology. – Lepidocupania grandiflora is
endemic to Grande Terre, where is has a peculiar distribu-
tion and edaphic range; most collections are from the central
region (Mont Do and around the Col d’Amieu), but others
have been made in the Pouébo area, some km to the north-
west. Lepidocupania grandiora grows in wet forest and maquis
vegetation, on ultramac and non-ultramac substrates, from
 to m.
Notes. – A (: ) suggested a close affinity
between this species and L. oedipoda based on vegetative
276 – Two new genera of Sapindaceae (Cupanieae) from the southern Pacific Candollea 75, 2020

characters. eir morphological similarity points toward the
need for further analyses, especially focusing on collections
from the Pouébo area.
Lepidocupania guillauminii (Kaneh.) Buerki, Callm., Munz-
inger & Lowry, comb. nov.
 Mischocarpus guillauminii Kaneh. in Bot. Mag. (Tokyo)
: . .  Cupaniopsis guillauminii (Kaneh.)
Adema in Leiden Bot. Ser. : . .
Holotypus: Caroline islands: Truk [Chuuk] atoll,
VI., Kanehira 1268 (FU; iso-: A [A] image
seen, BISH [BISH] image seen, P [P]!).
Distribution and ecology. – Lepidocupania guillauminii
is endemic to Chuuk atoll in the Caroline Islands, where it
grows in lowland evergreen forests on volcanic soil and humus
(A, ).
Note. – While A (: ) noted that the avail-
able material of this species is rather incomplete and that it
resembles both L. concolor and L. samoensis, he recognized it
as distinct based on several dierences in the amount of indu-
ment and in features of the fruits.
Lepidocupania inoplea (Radlk.) Buerki, Callm., Munzinger &
Lowr y, comb. nov.
 Cupaniopsis inoplea Radlk. in Sitzungsber. Math.-Phys.
CI. Konigl. Bayer. Akad. Wiss. Munich : . .
Lectotypus (designated here): New Caledonia. Prov.
Nord: Mt. Poum, V., Balansa 3307 (P [P]!;
isolecto-: P [P, P]!).
Distribution and ecology. – Lepidocupania inoplaea, as cir-
cumscribed by A (), is endemic to the archipelago
of New Caledonia, where it occurs in the northwestern part
of Grande Terre and two of the Loyalty Islands (Lifou and
Maré). It is found in maquis vegetation and gallery forest on
serpentinite and schist, from sea level to m.
Notes. – Cupaniopsis inoplea was described based on Balansa
3307. Original material at P is mounted on three sheets, two
of which [P, P] bear the following note in
Radlkofer’s hand: “Cupaniopsis inoplaea m. Radlk.” Here we
designate the most complete and best-preserved of these two
sheets [P] as the lectotype.
It has come to our attention that the original spelling of
the epithet, ‘inoplea’, was changed to ‘inoplaea’ by G
() and A (), but the rules of nomenclature do
not justify this change, and we therefore retain the original
spelling, as proposed by R (b).
A () did not indicate that this species grows on
calcareous substrate, although two collections cited in his mon-
ograph from the Loyalty Islands (Schmid 677 and MacKee (Leg.
Suprin) 43447) were gathered without doubt from calcareous
sites. Further study will be needed to determine whether these
collections belong to Lepidocupania inoplea or another species.
Lepidocupania lepidota (Radlk.) Buerki, Callm., Munzinger
& Lowr y, comb. nov. (Fig.C).
 Arytera lepidota Radlk. in Sitzungsber. Math.-Phys. CI.
Konigl. Bayer. Akad. Wiss. Munich : . .
Lectotypus (first step designated by T, :
; second step designated here): New Caledonia.
Prov. Sud: Mont Dore, s.d., Pancher [Mus. Néocal.] 222
(P[P]!; isolecto-: C [C] image seen,
K [K]!, MEL [MEL] image seen;
NY[NY] image seen, P [P]!).
Distribution and ecology. – Lepidocupania lepidota is endemic
to Grande Terre, where it is restricted to dense humid forest on
ultramac substrates, from  to m. Most collections are
from the main ultramac massif of the south, although it is also
recorded from a few isolated localities in the north, including
Mont Do and Cap Bocage (T, ), and more recently
from Kantalupaik (M et al., ).
Notes. – Seven syntype collections were cited in the proto-
logue of Arytera lepidota: Balansa 1445, 2841, Baudouin 134A,
Pancher [Mus. Néocal.] 222, Vieillard 205, and 206. T
(: ) designated Pancher [Mus. Neocal.] 222 as the lecto-
type. e original material deposited at P is, however, mounted
on two sheets, necessitating the second step lectotypication
designated here, for which we have selected the most complete
and best-preserved sheet [P].
Lepidocupania mouana (Guillaumin) Buerki, Callm., Munz-
inger & Lowry, comb. nov.
 Cupaniopsis mouana Guillaumin in Mém. Mus. Natl.
Hist. Nat., Sér. B, Bot. : . .
Holotypus: New Caledonia. Prov. Nord: Mt. Mou,
.II., Baumann-Bodenheim [leg. Baas Becking] 6036
(P [P]!; iso-: L [L]!, Z [Z-]
image seen).
Distribution and ecology. – Lepidocupania mouana is
endemic to Grande Terre, where it is restricted to Mont Mou
in the southwest. A () cited a single specimen, the
holotype, collected in , while an additional collection was
made in  (Grignon & Munzinger 256). is species grows
in continuous tall maquis (“maquis paraforestier”, see MC
Candollea 75, 2020 Two new genera of Sapindaceae (Cupanieae) from the southern Pacific – 277

et al.,  for details) dominated by species of Gymnostoma
L.A.S. Johnson, on peridotitic substrate.
Note. – Lepidocupania mouana was assigned an IUCN risk of
extinction status of “Endangered” [EN] by J et al. ().
Lepidocupania myrmoctona (Radlk.) Buerki, Callm., Munz-
inger & Lowry, comb. nov.
 Cupaniopsis myrmoctona Radlk. in Sitzungsber. Math.-
Phys. CI. Konigl. Bayer. Akad. Wiss. Munich : .
.
Lectotypus (designated by A, : ; second
step designated here): New Caledonia: sine loco, s.d.,
Labillardière s.n. (G [G]!; isolecto-: FI [FI
FI]!, G [G, G]!, K).
Distribution and ecology. – Lepidocupania myrmoctona is
endemic to Grande Terre, where it is abundant along the
east coast and grows in dense humid forest, mostly on non-
ultramac substrate (schist), but it has also been recorded in
montane ecosystems in the main southern ultramac massif
(A, ).
Notes. – Cupaniopsis myrmoctona was described based on
“Labillardière (Hb. Webb, Hook., Deless.)” (R, b:
). A (: ) designated material from G as the
lectotype. A second step lectotypication is, however, required
because the original material at G comprises three sheets origi-
nating from three dierent herbaria: Delessert [G],
Moricand [G], and Ventenat [G]. Vente-
nat’s herbarium was originally part of the Delessert herbarium
(C et al., ) and both [G] and
[G] are annotated in Radlkofer’s hand: “Cupaniopsis
myrmoctona m. Radlk.”. Here we designate the most complete
and best-preserved sheet [G], originating from the
Delessert herbarium, as the lectotype.
Lepidocupania oedipoda (Radlk.) Buerki, Callm., Munzinger
& Lowr y, comb. nov.
 Cupaniopsis oedipoda Radlk. in Sitzungsber. Math.-Phys.
CI. Konigl. Bayer. Akad. Wiss. Munich : . .
Lectotypus (rst step designated by A, : ;
second step designated here): New Caledonia. Prov.
Sud: escarpements du Cougui [Koghi], m, s.d.,
Pancher s.n. (P [P]!; isolecto-: M [M]
image seen; P [P, P, P]!).
= Cupaniopsis ganophloea Radlk. in Sitzungsber. Math.-
Phys. CI. Konigl. Bayer. Akad. Wiss. Munich : .
. Lectotypus (designated here): New Caledonia.
Prov. Nord: “Balade”, 
– , Veillard 230
(P[P]!; probable isolecto-: P [P]!).
Distribution and ecology. – Lepidocupania oedipoda is
restricted to Grande Terre, extending to the northwest in the
Belep archipelago (Art Island); it grows in maquis vegeta-
tion, humid forests, gallery forests, and forest remnants, often
along streams, usually on hills or mountainsides, on serpentine,
greywacke and schist, from sea level to m (A, ).
Notes. – Five syntypes were cited in the protologue of
Cupaniopsis oedipoda: Balansa 153 p.p., 1441 and 2257, Baudouin
354 p.p. and Pancher s.n. A (: ) designated Pancher
s.n. as the lectotype. Original material of this collection at P is,
however, mounted on four sheets, necessitating a second step
lectotypication. Here we designate the fruiting material with
a note in Radlkofer’s hand: “Cupaniopsis oedipoda m. Radlk.”
as the lectotype.
Cupaniopsis ganophloea was described based on four syn-
types: Labillardière s.n., Pancher 777, Vieillard 227, 230 and
231. e most complete and best-preserved material bearing,
in Radlkofer’s hand, “Cupaniopsis ganophloea m. Radlk.”
is [P]. is collection has no label data and could
represent material of either Veillard 230 or 231. Veillard 230 in
P[P] only has a few fruits with the note “Cupani-
opsis ganophloea m. Radlk.” in Radlkofer’s hand. We prefer to
designate the later sheet as lectotype and consider [P]
as a probable isolectotype.
Lepidocupania pennelii (Guillaumin) Buerki, Callm., Munz-
inger & Lowry, comb. nov.
 Cupaniopsis pennelii Guillaumin in Bull. Soc. Bot.
Fr.: . .
Holotypus: New Caledonia. Prov. Sud: env. de
Bourail, .II., Pennel 403 (P [P]!; isolecto-:
P [P, P]!).
Distribution and ecology. – Lepidocupania pennelii is
endemic to Grande Terre, where it is restricted to the west-
central coastal area between Bourail and Moindou, and one
site along the east coast at the same latitude, around Saint Pol.
It grows in forests on limestone, from  to m in elevation
(A, ).
Notes. – Cupaniopsis pennelii was described based on Pennel
403. Material of the collection at P is mounted on three sheets,
but only one of them [P] bears the name of Guil-
laumin’s new species in his own hand, and we therefore regard
it as the holotype.
Lepidocupania pennelii is morphologically similar to
L.globosa and L. rosea (the latter known only from the type
specimen), whose ecological preferences and distributions are
nearly the same. ey likely form a species complex and will
require further taxonomic work.
278 – Two new genera of Sapindaceae (Cupanieae) from the southern Pacific Candollea 75, 2020

Lepidocupania rosea (Adema) Buerki, Callm., Munzinger &
Lowr y, comb. nov.
 Cupaniopsis rosea Adema in Leiden Bot. Ser. : . .
Holotypus: New Caledonia. Prov. Nord: km E of
Col de Crève-Coeur on road between Canala and io,
c.m, .IX., McPherson 1905 (L [L]!;
iso-: MO [MO]!, NOU [NOU]!,
P[P]!).
Distribution and ecology. – Lepidocupania rosea is only
known from the type specimen, collected on Grande Terre,
near Nakety, in a forest around m.
Notes. – See note under Lepidocupania pennelii.
Lepidocupania samoensis (Christoph.) Buerki, Callm., Mun-
zinger & Lowry, comb. nov.
 Cupaniopsis samoensis Christoph. in Bernice P. Bishop
Mus. Bull. : . .
Holotypus: Samoa: Savai`i, above Matavanu,
.VIII., Christophersen & Hume 2045 (BISH
[BISH] image seen; iso-: A [A] image
seen, BISH [BISH, BISH] images seen,
K[K]!, P [P]!, UC [UC]
image seen, US [US] image seen).
Distribution and ecology. – Lepidocupania samoensis is
endemic to the two main islands of Samoa, Savai`i and Upolu,
where it grows in primary evergreen forests between  to
m (A, ).
Lepidocupania squamosa (Adema) Buerki, Callm., Munzinger
& Lowr y, comb. nov.
 Cupaniopsis squamosa Adema in Bull. Mus. Natl. Hist.
Nat., B, Adansonia, : . .
Holotypus: New Caledonia. Prov. Nord: massif
de la Tiébaghi, c. m, .XII., McPherson
6176 (MO[MO]!; iso-: L [L]!,
NOU[NOU]!, P[P]!).
Distribution and ecology. – Lepidocupania squamosa is
endemic to Grande Terre, where it grows in dense maquis-like
scrub vegetation on serpentine, from  to m (A,
, and recent collections) and on isolated ultramac moun-
tains of the northeast (Boulinda, Kopéto, Tiébaghi, and Poum).
Notes. – Lepidocupania squamosa was assigned an IUCN
risk of extinction status of “Endangered” by J et al.
(). Since this assessment was conducted, a large portion
of the vegetation on the Tiébaghi massif has been cleared for
mining, which has surely led to further population decline
(especially concerning since this area contained the largest
subpopulation of L. squamosa).
Lepidocupania subfalcata (Adema) Buerki, Callm., Munzinger
& Lowr y, comb. nov.
 Cupaniopsis subfalcata Adema in Leiden Bot. Ser. :
. .
Holotypus: New Caledonia. Prov. Nord: summit
plateau Mt. Koniambo, 
– m, .III., MacKee
4297 (L [L]!; iso-: A [A] image seen,
K[K]!, P [P]!).
Distribution and ecology. – When A () published
his monograph of Cupaniopsis, this species was thought to
be endemic to Grande Terre and was known only from the
type collection from Mt. Koniambo, between 
– m.
However, one year later, Adema identied a second specimen
to this species (Jaré 2944) in L [L.] from the Mé
Adéo road, about km southeast of Koniambo.
Notes. – e holotype of this species was initially identied
by Guillaumin as C. sebertii Guillaumin, a name considered a
synonym of C. fruticosa ( Lepidocupania fruticosa) by A
(). Our examination of the available herbarium material
suggests that further taxonomic analysis is needed to clarify
Candollea 75, 2020 Two new genera of Sapindaceae (Cupanieae) from the southern Pacific – 279
New Caledonia
Lepidocupania 17 spp.
Neoarytera 4 spp.
Vanuatu
Lepidocupania 1 sp.
Neoarytera 1 sp.
Fiji
Lepidocupania 2 spp.
Samoa
Lepidocupania 1 sp.
Caroline Islands
Lepidocupania 1 sp.
Solomon Islands
Lepidocupania 1 sp.
Wallis et Futuna
Lepidocupania 1 sp.
Fig.4.
–
Distribution map of Lepidocupania Buerki, Callm.,
Munzinger & Lowry (black line) and Neoarytera Callm., Buerki,
Munzinger & Lowry (white line) in the southern Pacific.
[Oceania Region Map by Vemaps.com]

species delimitations within this group, which probably rep-
resents a species complex.
Lepidocupania tontoutensis (Guillaumin) Buerki, Callm.,
Munzinger & Lowry, comb. nov.
 Cupaniopsis tontoutensis Guillaumin in Mém. Mus.
Natl. Hist. Nat., Sér. B, Bot. : . .
Holotypus: New Caledonia. Prov. Sud: cours moyen
de la Tontouta, rive droite, c.m, .I., Virot 1448
(P[P]!; iso-: P [P]!).
= Cupaniopsis rotundifolia Adema in Leiden Bot. Ser. :
. . Holotypus: New Caledonia. Prov. Sud:
colline surplombant la Tontouta, .VIII., Jaré
2531 (NOU [NOU]!; iso-: P [P,
P]!), syn. nov.
Distribution and ecology. – Lepidocupania tontoutensis is
endemic to Grande Terre, where it is restricted to the Tontouta
River valley. It is found in maquis vegetation on serpentine sub-
strate, mostly alluvium, between  and m (A, ).
Notes. – Cupaniopsis tontoutensis was described based on
Virot 1448. Original material at P is, however, mounted on
two sheets, only one of which [P] bears the name of
Guillaumin’s new species in his own hand, along with the word
“type”. We regard this specimen as the holotype.
Cupaniopsis rotundifolia Adema was only known from the
type specimen from the Tontouta River valley, with ecological
and edaphic preferences similar to those of C. tontoutensis (
Lepidocupania tontoutensis), as mentioned by A (:
), who wrote: “Probably closely related to C. fruticosa and
C. tontoutensis”. Cupaniopsis tontoutensis was said to dier from
C. rotundifolia by its fewer, wider leaets with the secondary
venation oriented at a wider angle to the midrib, and by the
sti trichomes on the endocarp of its fruits. However, several
recent collections made in the lower Tontouta River valley as
part of a study of rare species from this area (D’A, )
show a perfect continuum in leaet shape (as exemplied by
Lannuzel & D'Angelo 49 [MPU], Lannuzel & D'Angelo
34 [MPU], and Lannuzel & D'Angelo 39 [MPU])
and fruits (cf. Lannuzel & D'Angelo 32 [MPU]), with
leaves clearly corresponding to L. tontoutensis and the presence
of sti trichomes on the endocarp. We have therefore placed
C. rotundifolia in synonymy under L. tontoutensis.
Description and synopsis of Neoarytera
Neoarytera Callm., Buerki, Munzinger & Lowry, gen. nov.
(Fig.D
– F).
Type: Neoarytera chartacea (Radlk.) Callm., Buerki, Mun-
zinger & Lowry ( Arytera chartacea Radlk.).
= Arytera subsect. Pacica H. Turner in Blumea, Suppl.
: . . Type: Arytera collina Pancher & Sebert (
Neoarytera collina (Pancher & Sebert) Callm., Buerki,
Munzinger & Lowry).
Trees. Indument comprising short, straight, appressed
trichomes; glandular scales absent; buds not “varnished”.
Leaves alternate, 
– -jugate; leaets opposite to subopposite,
subsessile to petiolulate, margin entire. Inorescences axil-
lary or pseudo-terminal. Flowers actinomorphic, functionally
unisexual; calyx -dentate, teeth equal; petals , with a distinct
claw, scales minute, petal margin with minute enations; disc
more or less distinctly five-lobed, rim subpilose; stamens
(
–)(– ), anthers basifixed; ovary (
–)-locular. Fruit a
capsule, with  or  well developed lobes, dehiscence loculi-
cidal, central axis distinctly thickened, glabrous to puberulous
and rugose to verrucose outside, pilose inside, calyx persistent;
seed ellipsoid, sarcotesta covering all or sometimes only half of
the seed (N. neoebudensis), eshy-membranaceous, comprising
a single layer. Radicle bearing trichomes, at least basally.
Distribution. – Neoarytera comprises four species, three
endemic to New Caledonia and one occurring on both
NewCaledonia and Vanuatu (Fig.).
Notes. – Neoarytera is characterized by the distinctly thick-
ened central axis of its fruit (vs. not thickened in Arytera and
Cupaniopsis), petal scales comprising minute enations (vs. scales
adnate to the margins of the petals or free from the petals in
the other two genera), and its -, rarely -locular ovary and fruit
(vs. ovary and fruit - or -locular in the two other genera).
Arytera, as re-circumscribed here, can easily be distinguished
by its -layered sarcotesta (vs. single-layer in Cupaniopsis and
Neoarytera) (T, : ). Finally, Cupaniopsis diers from
Arytera and Neoarytera by having free or nearly free calyx lobes,
which are usually -seriate, orbicular, elliptic or obovate, and
concave (vs. calyx shortly cupular, the lobes toothed or divided,
and usually ovate in the other two genera) (R, a).
Neoarytera chartacea (Radlk.) Callm., Buerki, Munzinger &
Lowr y, comb. nov. (Fig.F).
 Arytera chartacea Radlk. in Sitzungsber. Math.-Phys.
Cl. Königl. Bayer. Akad. Wiss. München : . .
Lectotypus (rst step designated by T, : ;
second step designated here): Nouvelle-Calédonie.
Prov. Sud: Port des Français près de Nouméa, IX.,
Balansa 147 (P [P]!; isolecto-: K [K
K]!, M [M, M] images
seen, NY [NY] image seen, P [P,
P]!).
280 – Two new genera of Sapindaceae (Cupanieae) from the southern Pacific Candollea 75, 2020

Distribution and ecology. – Neoarytera chartacea is endemic
to the west coast of Grande Terre, where it grows in sclerophyll
forest or various types of substrate, in particular limestone and
serpentine (T, ).
Notes. – Three syntypes were cited in the protologue of
Arytera chartacea: Balansa 147, 1442 and Pancher 610. T
(: ) designated Balansa 147 as the lectotype. Original
material at P is, however, mounted on three sheets, necessitat-
ing a second step lectotypication. Here we designate the most
complete and best-preserved sheet [P] as the lectotype.
Neoarytera collina (Pancher & Sebert) Callm., Buerki, Mun-
zinger & Lowry, comb. nov. (Fig.D).
 Cupania collina Pancher & Sebert in Rev. Marit. Colon.
: . .  Arytera collina (Pancher & Sebert)
Radlk. in Not. Syst. (Paris) : . .
Lectotypus (rst step designated by T, : ;
second step designated here): New Caledonia: sine
loco, s.d., Pancher [Bois] 79 (P [P]!; isolecto-:
P[P, P, P]!).
= Arytera pachyphylla Radlk. in Sitzungsber. Math.-
Phys. Cl. Königl. Bayer. Akad. Wiss. München
: . . Lectotypus (designated here): New
Caledonia. Prov. Sud: “environs de Nouméa, Pont
des Français”, 
– , Deplanche 280 [Vieillard
2391] (P [P]!; isolecto-: K [K]!,
G[G, G]!, P [P,
P, P, P, P,
P, P]!).
Distribution and ecology. – Neoarytera collina is endemic to
New Caledonia, where it occurs on the southwestern coast of
Grande Terre, Ile des Pins, and Maré Island in the Loyalties.
It grows in sclerophyll forest on various types of substrate, in
particular limestone and serpentine (T, ).
Notes. – T (: ) designated Pancher [Bois] 79
as the lectotype, since it was cited by P & S
(: ) as material associated to Cupania collina when they
described it in their Notice sur les bois de la Nouvelle Calédonie.
Original material at P is mounted on four sheets, necessitating
a second step lectotypication, for which we have here chosen
the most complete and best-preserved sheet [P].
Six syntypes were cited in the protologue of Ar ytera
pachyphylla: Balansa 148, Baudouin 690, Deplanche 280, 447,
Pancher [Mus. Néocal.] 215, 778 and Vieillard 247. T
(: ) cited only two of these syntypes, Baudouin 690
and Deplanche 280. Original material of these two collections
at P is, however, mounted on three sheets, from among which
we designate the most complete and best-preserved collection
of Deplanche 280 as the lectotype [P].
Neoarytera nekorensis (H. Turner) Callm., Buerki, Munzinger
& Lowr y, comb. nov.
 Arytera nekorensis H. Turner in Blumea, Suppl. : .
.
Holotypus: New Caledonia. Prov. Nord: Poya, forêt de
Nekoro, .VIII., MacKee 42137 (L [L]!; iso-:
P [P, P]!).
Distribution and ecology. – Neoarytera nekorensis is endemic
to the Nekoro forest in the centre-west region of Grande
Terre, where it grows in sclerophyll forest on black clay soil
(T, ).
Notes. – is species has been assigned an IUCN risk of
extinction status of “Vulnerable” (IUCN, ).
Neoarytera neoebudensis (Guillaumin) Callm., Buerki, Mun-
zinger & Lowry, comb. nov. (Fig.E).
 Cupaniopsis neoebudensis Guillaumin in J. Arnold Arbor.
: . .  Arytera neoebudensis (Guillaumin)
H.Turner in Blumea : . .
Holotypus: Va n u a t u . Prov. Taféa: Erromango Isl., Dillon
Bay, .VI., Kajewski 381 (A; iso-: BISH, BRI [BRI-
AQ]!, K [K]!, NY, P [P]!).
Distribution and ecology. – Neoarytera neoebudensis, as
currently circumscribed, occurs in New Caledonia (Grande
Terre and the Loyalty islands), Walpole Island, and Vanuatu.
It grows on rocky slopes near lagoons and in lowland evergreen
forests on volcanic soils (T, ).
Acknowledgments
e authors wish to thank the following herbarium curators and
collaborators for their help: MarkCarine and RaneePrakash
(BM), PaulForster (BRI), FrankZich (CNS), NicolasFumeaux
and LaurentGautier (G), AlanPaton (K), FritsAdema and
NicolienSol (L), JimSalomon and PetePhillipson (MO),
CarolineLoup (MPU), VanessaHequet and JacquelineTinel
(NOU), MarcJeanson (P), SerenaLee and YeeWenLow
(SING), and MarikaTuiwawa and AliviretiNaikatini (SUVA).
We are very grateful to the Idaho Botanical Research Founda-
tion for its generous support of our eldwork; Guillaume Lan-
nuzel for crucial collections in the Toutouta valley; ChrisDavid-
son and SharonChristoph for fruitful discussions and their
enthusiastic interest in our research; RoyGereau for his help
with translating Radlkofer’s Latin diagnoses into English; and
Bruce and JoyGray, YeeWenLow, and AndrewFord for their
Candollea 75, 2020 Two new genera of Sapindaceae (Cupanieae) from the southern Pacific – 281

kind assistance in Queensland. Finally, we thank BruceGray,
IdrisA.Haris, VanessaHequet, Steve and AlisonPearson, and
GarrySankowsky for granting permission to use their photos,
and FelixForest and FredStauffer for improving an earlier
version of this manuscript.
Bibliography
A, F. (). Cupaniopsis Radlk. (Sapindaceae): a monograph.
Leiden Bot. Ser. .
A, F., P.W. L & P.C.  W (). Sapin-
daceae. Fl. Malesiana ser. , ().
B, C.L. (). Arytera. Rumphia : 
– .
B, P., T. J & J.M. V (). Plant extinction in
New Caledonia: protection of sclerophyll forests urgently needed.
Biodivers. Conserv. : 
– .
B, S., F. F, P. A-R, M.W. C,
J.A.A. N, M. H, I. S, P. K
& N. A (). Plastid and nuclear DNA markers reveal
intricate relationships at subfamilial and tribal levels in the soap-
berry family (Sapindaceae). Molec. Phylogenet. Evol. : 
– .
B, S., P.P. L II, N. A, S.G. R,
P. K & M.W. C (). Phylogeny and cir-
cumscription of Sapindaceae revisited: molecular sequence data,
morphology and biogeography support recognition of a new fam-
ily, Xanthoceraceae. Pl. Ecol. Evol. : 
– .
B, S., F. F, M.W. C, P.P. L II, D.S.D
& J. M (). Phylogenetic inference of New Caledo-
nian lineages of Sapindaceae: molecular evidence requires a reas-
sessment of generic circumscriptions. Taxon : 
– .
B, J.-F. (). Flore des îles Loyauté (Nouvelle-Calédonie):
plantes patrimoniales, plantes envahissantes et espaces naturels
remarquables. Rapport Conservation International, Nouméa &
Province des Iles Loyauté, Lifou.
C, M.W., O.D. D, H.W. L, P. B,
P. M & L. G (). Etienne-Pierre Ventenat
(
– ) and the gardens of Cels and Empress Joséphine.
Candollea : 
– .
C, M.W., A.J. F & S. B (). New com-
binations for two species in the genus Synima (Sapindaceae,
Cupanieae) from Queensland (Australia). Candollea : 
– .
D'A, S. (). Ecologie et conservation de trois espèces rares et
menacées de Nouvelle-Calédonie. Mémoire de Master. Université
de Lorraine-UMR ECOFOG, Kourou & IAC, Païta.
F, P.I. (). Synima reynoldsiae P.I. Forst. (Sapindaceae),
a new species from the ‘Wet Tropics’ of north-east Queensland.
Austrobaileya : 
– .
G, A. (). Sapindacées. Flore analytique et synoptique
de la Nouvelle-Calédonie, phanérogames: 
– . Office de la
Recherche Scientique Coloniale, Paris.
H, M.G., K.J. E, S.A. J, M.W. C
& P.A. G (). Phylogenetic inference in Sapindaceae
sensu lato using plastid matK and rbcL DNA sequences. Syst.
Bot. : 
– .
IUCN (). e IUCN Red List of threatened species. [https://www.
iucnredlist.org]
J, T., P. B & J.M. V (). reatened plants
of New Caledonia: is the system of protected areas adequate?
Biodivers. Conserv. : 
– .
MC, S.G., T. J, F. R & J.E. A (). Fire and
succession in the ultramac maquis of New Caledonia. J. Biogeog.
: 
– .
M, J., M. P & D. B (). Flore & végétation du
Katalupaik. In: P, O. (ed.), La Planète Revisitée, Nouvelle-
Calédonie 2016
– 18, Volet «Forêt» 2017 en province Nord. Rapport
d ’ét ap e : 
– . ProNatura, MNHN, Paris.
P, J.A.I. & H. S (). Notice sur les bois de la Nouvelle
Calédonie. Paris.
R, L. (a). Ueber die Sapindaceen Holländisch-Indi-
ens. Actes du congrès international de botanistes, d’horticulteurs,
de négociants et de fabricants de produits du règne végétal tenu à
Amsterdam 1877: 
– ,  – . A. W. Sijtho, Leide.
R, L. (b). Ueber Cupania und damit verwandte
Panzen. Sitzungsber. Math.-Phys. CI. Konigl. Bayer. Akad. Wiss.
München : 
– .
R, L. (
– ). Sapindaceae. In: E, A. (ed.), Das
Panzenreich a-h. W. Engelmann, Leipzig.
R, S.T. (). Notes on Sapindaceae II. Austrobaileya :

– .
R, S.T. (). Notes on Sapindaceae III. Austrobaileya :

– .
R, S.T. (a). Notes on Sapindaceae IV. Austrobaileya :

– .
R, S.T. (b). Sapindaceae. In: G, A.S. (ed.), Fl.
Australia : 
– .
S, A. (). e two New Guinea species of Lepiderema Radlk.
(Sapindaceae). Blumea : 
– .
S, A.C. (). Flora vitiensis nova . Pacic Tropical Garden,
Kauai.
T, H. (). Cladistic and biogeographic analyses of Arytera
Blume and Mischarytera gen. nov. (Sapindaceae) with notes on
methodology and a full taxonomic revision. Blumea, Suppl. .
282 – Two new genera of Sapindaceae (Cupanieae) from the southern Pacific Candollea 75, 2020

Candollea 75, 2020 Two new genera of Sapindaceae (Cupanieae) from the southern Pacific – 283
Appendix
–
Synopsis of accepted species of Arytera Blume, Cupaniopsis Radlk., Lepidocupania Buerki, Callm., Munzinger & Lowry,
and Neoarytera Callm., Buerki, Munzinger & Lowry, with their respective distributions.
Genus Species Distribution
Arytera Blume
A. bifoliolata S.T. Reynolds Indonesia (Western New Guinea), Australia
A. brachyphylla Radlk. Papua New Guinea
A. densiflora Radlk. Papua New Guinea
A. dictyoneura S.T. Reynolds Australia
A. distylis (Benth.) Radlk. Australia
A. divaricata F. Muell. Australia
A. foveoleata F. Muell. Australia
A. lineosquamulata H.Turner Papua New Guinea, Australia
A. litoralis Blume From India across SE Asia throughout Malesia up to the Solomon Islands
A. microphylla (Benth.) Radlk. Australia
A. miniata H.Turner Papua New Guinea
A. morobeana H.Turner Papua New Guinea
A. multijuga H.Turner Papua New Guinea
A. musca H.Turner Papua New Guinea
A. novaebrittanniae H.Turner Papua New Guinea, Solomon Islands
A. pauciflora S.T. Reynolds Australia
A. pseudofoveolata H.Turner Papua New Guinea, Australia
Cupaniopsis Radlk.
C. acuticarpa Adema Papua New Guinea
C. amoena A.C. Sm. Fiji
C. anacardioides (A.Rich.) Radlk. Indonesia (Western New Guinea), Papua New Guinea, Australia
C. apiocarpa Radlk. New Caledonia
C. azantha Radlk. New Caledonia
C. baileyana Radlk. Australia
C. bilocularis Adema Papua New Guinea
C. bullata Adema Papua New Guinea
C. celebica Adema Indonesia (Sulawesi)
C. chytradenya Radlk. New Caledonia
C. cooperorum P.I. Forst. Australia
C. crassivalvis Radlk. New Caledonia
C. curvidens Radlk. Indonesia (Western New Guinea), Papua New Guinea
C. dallachyi S.T. Reynolds Australia
C. diploglottoides Adema Australia
C. euneura Adema Papua New Guinea
C. flagelliformis Radlk. Australia
C. fleckeri S.T. Reynolds Australia
C. foveolata Radlk. Australia
C. grisea Adema New Caledonia
C. hypodermatica Radlk. New Caledonia
C. kajewskii Merr. & L.M. Perry Papua New Guinea, Solomon Islands
C. leptobotrys Radlk. Vanuatu, Fiji
C. mackeeana Adema New Caledonia
C. macrocarpa Radlk. New Caledonia

284 – Two new genera of Sapindaceae (Cupanieae) from the southern Pacific Candollea 75, 2020
Genus Species Distribution
C. macropetala Radlk. Indonesia (Western New Guinea), Papua New Guinea
C. megalocarpa Adema New Caledonia
C. napaensis Adema Papua New Guinea
C. newmanii S.T. Reynolds Australia
C. petiolulata Radlk. New Caledonia
C. phalacrocarpa Adema New Caledonia
C. phanerophlebia Merr. & L.M. Perry Papua New Guinea
C. platycarpa Radlk. Indonesia (Western New Guinea), Papua New Guinea
C. rhytodocarpa Adema Papua New Guinea
C. serrata Radlk. Australia
C. shirleyana Radlk. Australia
C. stenopetala Radlk. Indonesia (Moluccas), Papua New Guinea
C. strigosa Adema Indonesia (Sulawesi)
C. sylvatica Guillaumin New Caledonia
C. tomentella (F.Muell. ex Benth.) S.T. Reynolds Australia
C. trigonocarpa Radlk. New Caledonia
C. vitiensis Radlk. Fiji
C. wadsworthii Radlk. Australia
Lepidocupania Buerki et al.
L. arcuata (Radlk.) Buerki et al. New Caledonia
L. brackenridgei (A. Gray) Buerki et al. Vanuatu, Fiji, Wallis et Futuna, Samoa, Solomon Islands
L. concolor (Gillespie) Buerki et al. Fiji
L. fruticosa (Radlk.) Buerki et al. New Caledonia
L. glabra (Adema) Buerki et al. New Caledonia
L. globosa (Adema) Buerki et al. New Caledonia
L. glomeriflora (Radlk.) Buerki et al. New Caledonia
L. gracilipes (Radlk.) Buerki et al. New Caledonia
L. grandiflora (Adema) Buerki et al. New Caledonia
L. guillauminii (Kaneh.) Buerki et al. Caroline Islands
L. inoplea (Radlk.) Buerki et al. New Caledonia
L. lepidota (Radlk.) Buerki et al. New Caledonia
L. mouana (Guillaumin) Buerki et al. New Caledonia
L. myrmoctona (Radlk.) Buerki et al. New Caledonia
L. oedipoda (Radlk.) Buerki et al. New Caledonia
L. pennelii (Guillaumin) Buerki et al. New Caledonia
L. rosea (Adema) Buerki et al. New Caledonia
L. samoensis (Christoph.) Buerki et al. Samoa
L. squamosa (Adema) Buerki et al. New Caledonia
L. subfalcata (Adema) Buerki et al. New Caledonia
L. tontoutensis (Guillaumin) Buerki et al. New Caledonia
Neoarytera Callm. et al.
N. chartacea (Radlk.) Callm. et al. New Caledonia
N. collina (Pancher & Sebert) Callm. et al. New Caledonia
N. nekorensis (H. Turner) Callm. et al. New Caledonia
N. neoebudensis (Guillaumin) Callm. et al New Caledonia, Vanuatu