Plant Ecology and Evolution FastTrack: 1–12, 2010
doi:10.5091/plecevo.2010.437
REGULAR PAPER
Phylogeny and circumscription of Sapindaceae revisited:
molecular sequence data, morphology and biogeography
support recognition of a new family, Xanthoceraceae
Sven Buerki1, Porter P. Lowry II2,3,*, Nadir Alvarez4, Sylvain G. Razaimandimbison5,
Philippe Küpfer6 & Martin W. Callmander2,7
Department of Biodiversity and Conservation, Real Jardin Botanico, CSIC, Plaza de Murillo 2, ES-28014 Madrid, Spain
Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299, U.S.A.
3
Muséum National d’Histoire Naturelle, Case Postale 39, 57 rue Cuvier, FR-75231 05 CEDEX, Paris, France
4
Department of Ecology and Evolution, Biophore, University of Lausanne, CH-1015 Lausanne, Switzerland
5
Department of Botany, Bergius Foundation, SE-10691, Stockholm University, Stockholm, Sweden
6
Institute of Biology, University of Neuchâtel, Rue Emile-Argand 11, CH-2000 Neuchâtel, Switzerland
7
Conservatoire et Jardin botaniques de la ville de Genève, ch. de l’Impératrice 1, CH-1292 Chambésy, Switzerland
*Author for correspondence: pete.lowry@mobot.org
1
2
Background and aims – Recent studies have adopted a broad deinition of Sapindaceae that includes
taxa traditionally placed in Aceraceae and Hippocastanaceae, achieving monophyly but yielding a family
dificult to characterize and for which no obvious morphological synapomorphy exists. This expanded
circumscription was necessitated by the inding that the monotypic, temperate Asian genus Xanthoceras,
historically placed in Sapindaceae tribe Harpullieae, is basal within the group. Here we seek to clarify
the relationships of Xanthoceras based on phylogenetic analyses using a dataset encompassing nearly ¾
of sapindaceous genera, comparing the results with information from morphology and biogeography, in
particular with respect to the other taxa placed in Harpullieae. We then re-examine the appropriateness of
maintaining the current broad, morphologically heterogeneous deinition of Sapindaceae and explore the
advantages of an alternative family circumscription.
Methods – Using 243 samples representing 104 of the 142 currently recognized genera of Sapindaceae s.
lat. (including all in Harpullieae), sequence data were analyzed for nuclear (ITS) and plastid (matK, rpoB,
trnD-trnT, trnK-matK, trnL-trnF and trnS-trnG) markers, adopting the methodology of a recent familywide study, performing single-gene and total evidence analyses based on maximum likelihood (ML) and
maximum parsimony (MP) criteria, and applying heuristic searches developed for large datasets, viz. a new
strategy implemented in RAxML (for ML) and the parsimony ratchet (for MP). Bootstrap analyses were
performed for each method to test for congruence between markers.
Key results – Our indings support earlier suggestions that Harpullieae are polyphyletic: Xanthoceras is
conirmed as sister to all other sampled taxa of Sapindaceae s. lat.; the remaining members belong to three
other clades within Sapindaceae s. lat., two of which correspond respectively to the groups traditionally
treated as Aceraceae and Hippocastanaceae, together forming a clade sister to the largely tropical Sapindaceae s. str., which is monophyletic and morphologically coherent provided Xanthoceras is excluded.
Conclusion – To overcome the dificulties of a broadly circumscribed Sapindaceae, we resurrect the historically recognized temperate families Aceraceae and Hippocastanaceae, and describe a new family, Xanthoceraceae, thus adopting a monophyletic and easily characterized circumscription of Sapindaceae nearly
identical to that used for over a century.
Key words – Aceraceae, Harpullieae, Hippocastanaceae, molecular phylogeny, new family, Sapindaceae,
Xanthoceras, Xanthoceraceae.
All rights reserved. © 2010 National Botanic Garden of Belgium and Royal Botanical Society of Belgium – ISSN 2032-3913
Pl. Ecol. Evol. FastTrack, 2010
INTRODUCTION
The systematics of the family Sapindaceae has challenged
taxonomists for more than a century since its irst comprehensive treatment was published by Radlkofer (1890, 1933).
Until the late 1980s, Sapindaceae were widely treated as
distinct from two closely related families, Hippocastanaceae
and Aceraceae, based primarily on morphology and biogeography (Takhtajan 1987, Cronquist 1988, Dahlgren 1989).
Several recent studies using pollen morphology (Müller &
Leenhouts 1976), phytochemistry (Umadevi & Daniel 1991)
and molecular sequence data (Gadek et al. 1996, Savolainen
et al. 2000, APG II 2003, APG III 2009, Harrington et al.
2005, Buerki et al. 2009) have, however, led to the adoption
of a broader concept in an effort to ensure monophyly, uniting
these entities into a single family, Sapindaceae s. lat.
Sapindaceae s. lat. as currently circumscribed by Harrington et al. (2005), Thorne & Reveal (2007) and Buerki
et al. (2009, 2010) comprise c. 1900 species and 142 genera
distributed among four subfamilies: Dodonaeoideae Burnett,
Hippocastanoideae Burnett, Sapindoideae Burnett and Xanthoceroideae Thorne & Reveal. Recently, Buerki et al. (2009)
demonstrated the para-/polyphyly of all tribes as deined by
Radlkofer (1933), with a single exception, Paullinieae Kunth.
Although they sketched an informal system that recognizes a
dozen monophyletic groups, they did not propose new tribal
limits within the four subfamilies as many potentially important genera of Sapindaceae were not included in their study
due to the lack of sequenceable material.
Historically, Radlkofer (1933) recognized fourteen tribes
within Sapindaceae s. str., ive in Dodonaeoideae and nine in
Sapindoideae (see table 1 in Buerki et al. 2009 for details).
Within Dodonaeoideae, however, he encountered dificulty
assigning nine genera to the four previously described tribes,
ultimately deciding to place them in a new tribe, Harpullieae
Radlk. Within this heterogeneous assemblage, he recognized
two informal groups according to the presence (Delavaya Franchet, Ungnadia Endl. and Xanthoceras Bunge) or absence (Arfeuillea Pierre, Conchopetalum, Eurycorymbus Hand.-Mazz.,
Harpullia Roxb., Magonia A.St.-Hil. and Majidea J.Kirk ex
Oliv.) of a terminal lealet. While revising Radlkofer’s infrafamilial system, largely on the basis of pollen and other morphological features, Müller & Leenhouts (1976) discussed the
possible expansion of Harpullieae to include the three genera
comprising Hippocastanaceae, viz. Aesculus L., Billia L. and
Handeliodendron (these authors did not, however, comment on
the taxonomic status of Aceraceae). In their revised classiication, Müller & Leenhouts (1976) concluded that the connection
between Hippocastanaceae and Harpullieae might involve two
genera in particular, Handeliodendron, originally described in
Sapindaceae (Rehder 1935), and Delavaya, which has always
been placed in Sapindaceae. Müller & Leenhouts (1976) also
regarded Harpullieae as a “heterogeneous assemblage”, with
several genera dificult to connect to the others. For example,
they classiied Harpullia pollen as both type-A and type-H
and Magonia pollen as type-E, whereas other members of the
tribe exclusively exhibit the more common type-A pollen (see
Buerki et al. 2009 for more details on pollen morphology).
Moreover, Harpullieae range from tropical (e.g. Conchopetalum, Delavaya, Magonia) to temperate (Xanthoceras) regions
2
and include both evergreen and deciduous species (Radlkofer
1933, Müller & Leenhouts 1976). Based on wood anatomy,
Klaassen (1999) noted a difference between the temperate
and tropical genera in the tribe, and among the tropical ones
he indicated that Delavaya and Ungnadia stood out because
their wood is similar to that of members of tribe Cupanieae
Reichenb. (Sapindoideae). Buerki et al. (2009) found Harpullieae to be polyphyletic, with Xanthoceras occupying a basal
position within Sapindaceae s. lat., Arfeuillea, Eurycorymbus,
Harpullia and Majidea placed in Dodonaeoideae, Delavaya
occupying a basal position within Sapindoideae, and Conchopetalum resolved in the Macphersonia group (Sapindoideae;
Buerki et al. 2009) closely related to the newly described endemic Malagasy genus Gereaua Buerki & Callm. (Buerki et al.
2010). A close relationship between Delavaya and Ungnadia
was found in an earlier cladistic analysis based on morphology (Judd et al. 1994), which identiied the presence of prolonged basal petal appendages and glabrous stamens as putative synapomorphies, again suggesting that Harpullieae were
far from representing a natural assemblage.
In the present study we seek to (1) clarify the relationships of Xanthoceras within Sapindaceae s. lat. and in particular with respect to the other taxa traditionally and/or currently
placed in Harpullieae, and (2) re-examine the appropriateness
of maintaining the current broadly circumscribed but morphologically heterogeneous deinition of Sapindaceae and
explore the possible advantages of alternative family circumscriptions. Toward this end, we have signiicantly expanded
the dataset of Buerki et al. (2009) to conduct a new set of
phylogenetic analyses, comparing the results with information from morphology and biogeography.
MATERIAL AND METHODS
Sampling, sequence data and phylogenetic analyses
Species names, voucher information, and GenBank accession
numbers for all sequences are provided in the appendix. The
dataset presented in Buerki et al. (2009) was expanded to include a total of 243 samples encompassing more than 70% of
the generic diversity in Sapindaceae s. lat. (104 of the currently
recognized 142 genera; half of the 38 genera not included in
this analysis are monospeciic), representing an increase of
ninety ingroup samples and nineteen genera. To assess the phylogenetic relationships of the taxa placed in tribe Harpullieae
and in the traditionally recognized families Aceraceae and Hippocastanaceae, we sampled at least one species from each genus currently assigned to these groups by adding the following
genera: Magonia and Ungnadia from Harpullieae, plus Billia
and Handeliodendron from Hippocastanaceae (Aesculus, the
third member of this family, was included in the analysis of
Buerki et al. 2009, as were both genera of Aceraceae, Acer and
Dipteronia). The outgroup sampling included one taxon each
from Anacardiaceae (Sorindeia sp., used as the most external
outgroup), Meliaceae (Malleastrum sp.) and Simaroubaceae
(Harrisonia abyssinica Oliv.).
The DNA extraction, ampliication and sequencing protocols used for the nuclear and plastid regions are provided in
Buerki et al. (2009). The nuclear sequences include the whole
ITS region (ITS1, 5.8S and ITS2) and plastid markers include
Buerki et al., Recognition of a new family, Xanthoceraceae
both coding (matK and rpoB) and non-coding regions (the
trnL intron and the intergenic spacers trnD-trnT, trnK-matK,
trnL-trnF and trnS-trnG).
Single-gene, total evidence analyses and their corresponding bootstrap analyses were performed using the maximum parsimony (MP) and maximum likelihood (ML) criteria following
the same procedure as in Buerki et al. (2009). Parsimony ratchet (Nixon 1999) was performed for each partition and for the
combined data set using PAUPRat (Sikes & Lewis 2001). Ten
independent searches were performed with 200 iterations and
15% of the parsimony informative characters perturbed. A strict
consensus tree was constructed based on the shortest equally
parsimonious trees. To assess support at each node, non parametric bootstrap analyses (Felsenstein 1985) were performed
using PAUP* (Swofford 2002) following the same procedure
as in Buerki et al. (2009). Model selection for each partition
was assessed using Modeltest v. 3.7 (Posada & Crandall 1998).
ML analyses were performed using RAxML v. 7.0.0 (Stamatakis 2006, Stamatakis et al. 2008) with 1000 rapid bootstrap
analyses followed by a search for the best-scoring tree in one
single run. These analyses were done using the facilities made
available by the CIPRES portal in San Diego, USA (http://8ball.
sdsc.edu:8888/cipres-web/home).
Topological differences between single-gene phylogenetic
trees were compared by taking into account the level of resolution obtained by each marker and its bootstrap support. Topological differences with bootstrap support (BS) less than 75%
were not considered.
Table 1 – Characteristics of partitions used in the phylogenetic analyses of Sapindaceae s. lat.
IGS = intergenic spacer; MP = maximum parsimony; PI = potentially parsimony informative; CI = constistency index; RI = retention
index; 1for No. of sequences, the total number of samples for the combined analyses is indicated between brackets; 2for mean amount of
phylogenetic information per sample: averaged by alignment size/variable sites number/PI sites number.
Single-gene analysis
ITS
matK
rpoB
trnD-trnT
IGS
trnKmatK
IGS
trnL
intron
trnLtrnF
IGS
trnS-trnG
IGS
8 markers
No. of ingroup
sampled
species/genera
156/89
159/89
174/96
94/69
119/74
169/97
165/93
74/52
214/104
No. sequences1
172
175
199
102
133
192
189
80
1242 (243)
Phylogenetic
information
Sequences: this
study/Genbank
Sequence
length range
Alignment
length
165/5
146/29
199/0
102/0
131/2
188/4
186/3
80/0
-
650–705
1074–1242
357–363
1086–1425
705–753
510–522
380–430
1311–1365
-
1234
1614
363
1925
931
773
661
2156
9657
No. constant
characters (%)
584
(47.3%)
1062
(65.7%)
222
(61.1%)
965
(50.1%)
488
(52.4%)
454
(58.7%)
328
(49.6%)
1577
(73.1%)
5681
(58.8%)
No. variable
characters (%)
653
(52.9%)
552
(34.2%)
141
(38.8%)
960
(49.9%)
443
(47.6%)
319
(41.2%)
333
(50.4%)
579
(26.8%)
3976
(41.2%)
No. potentially
PI characters
(%)
491
(39.8%)
392
(24.3%)
104
(28.7%)
505
(26.2%)
277
(29.8%)
194
(25.1%)
217
(32.8%)
227
(10.5%)
2404
(24.9%)
-
-
-
-
-
-
-
-
16.4/39.7/9.9
214
1967
2010
1975
1703
1500
667
1820
949
4666
1185
281
1858
954
687
705
945
11526
0.272
0.632
0.672
0.694
0.633
0.644
0.646
0.711
0.492
0.673
0.860
0.907
0.790
0.775
0.881
0.858
0.650
0.749
Mean amount
of phylogenetic
information
per sample2
No. trees
retained
Tree length
(steps)
Consistency
Index (CI)
Retention
Index (RI)
3
A
C
Figure 1 – A, best maximum likelihood phylogenetic tree for Sapindaceae s. lat.; B, relationships between and within Aceraceae and Hippocastanaceae; C, relationships within subfamily
Dodonaeoideae. Bootstrap supports are indicated above branches. COS = Cossinieae; CUP = Cupanieae; DOD = Dodonaeeae; DOR = Doratoxyleae; HAR = Harpullieae. The deinition
of the groups follows Buerki et al. (2009).
Pl. Ecol. Evol. FastTrack, 2010
4
B
Buerki et al., Recognition of a new family, Xanthoceraceae
RESULTS
Alignment and phylogenetic analyses
The number of samples and statistics for each partition and
the combined data set are summarized in table 2. The best-it
model for all partitions was the general time reversible (GTR)
with an alpha parameter for the shape of the gamma distribution to account for among-site rate heterogeneity (GTR+G).
The only exception was the ITS region, in which a proportion
of invariable sites was added (GTR+G+I). The MP and ML
single-gene phylogenies provided different levels of resolution, but no differences with a bootstrap support greater than
75% were identiied when compared, so we combined them
in a total evidence approach. Statistics (number of most parsimonious trees; tree length; and consistency and retention
indices) for each analysis are reported in table 1.
For the combined analyses under the MP criterion, nine
of the ten independent PAUPrat searches converged on a best
score of 11526 steps and produced a total of 949 most parsimonious trees, which were used to compile a strict consensus
(not shown); this consensus tree comprised several polytomies, especially near the tips. Under the ML criterion, the
best-it model for the combined matrix was GTR+G+I. This
model was used to perform the single ML run search (log
likelihood = − 79995.7), followed by rapid bootstrap analyses.
When compared, analyses compiled under the MP and
ML criteria yielded very similar topologies. Moreover, as no
moderately to strongly supported differences were observed
between the two phylogenetic trees, only the ML topology
will be presented and discussed hereafter (igs 1 & 2).
Phylogenetic relationships
With the addition of the ninety ingroup samples used in the
present analysis, including representatives of several genera
of Sapindaceae s. lat. that had not previously been sequenced,
the phylogenetic relationships revealed here are highly congruent with the informal system proposed by Buerki et al.
(2009). Based on sampling that includes at least one representative of all genera traditionally placed in Sapindaceae
tribe Harpullieae, Aceraceae and Hippocastanaceae, our results further conirm that Xanthoceras sorbifolium Bunge
(previously assigned to Harpullieae by Radlkofer, 1933) is
resolved as sister to the remaining sampled members of Sapindaceae s. lat. (however with a low BS; ig. 1A). Our results also indicate that the other genera of Harpullieae belong
to three additional clades, one in subfam. Dodonaeoideae
and two in subfam. Sapindoideae (igs 1 & 2), conirming
the polyphyly of the tribe. Within Dodonaeoideae, ive of
the genera currently assigned to Harpullieae belong to the
Dodonaea group, viz. Arfeuillea, Eurycorymbus, Harpullia,
Magonia and Majidea (ig. 1C), and Harpullia itself appears
to be polyphyletic, with the three species sampled occupying
very different positions within the phylogeny (expanded sampling to include additional members of the genus are, however, needed to conirm this inding). Within Sapindoideae, two
of the three remaining genera assigned to Harpullieae (Delavaya and Ungnadia) are placed in the Delavaya group, the
basal most lineage within the subfamily, and the third genus
(Conchopetalum) belongs to the Macphersonia group (ig. 2).
The inclusion of Billia and Handeliodendron in our
analysis, along with additional species of Acer and Aesculus,
strengthens support for the monophyly of both Aceraceae and
Hippocastanaceae and conirms their sister relationship (ig.
1A & B). Our results suggest the possible paraphyly of Acer
(with respect to Dipteronia) and of Aesculus (with respect to
Billia and Handeliodendron), although this inding should be
tested further with additional sampling. Support for the clade
comprising Sapindaceae s. str. (i.e. Dodonaeoideae plus Sapindoideae) is likewise stronger in the present analysis (BS
= 88) than in that of Buerki et al. (2009; BS = 69; ig. 1).
Moreover, Diplokeleba N.E.Br., long regarded as a member
of Sapindoideae (tribe Cupanieae), is instead placed within
Dodonaeoideae (ig. 1C).
DISCUSSION
Polyphyly of Harpullieae
The results presented above clearly show that the tribe Harpullieae (as well as all other sapindaceous tribes with the
exception of Paullinieae), as deined initially by Radlkofer
(1890, 1933) and modiied by Müller & Leenhouts (1976),
is highly polyphyletic, with members placed in no fewer than
four clades scattered among various parts of Sapindaceae
s. lat. Harrington et al. (2005) and Buerki et al. (2009) argued that additional sampling (especially of Harpullieae)
was required before taking a deinitive stand regarding the
phylogenetic and taxonomic status of Xanthoceras. Although
we have now analyzed more than 70% of the genera and included all those that are putatively related to Xanthoceras,
its precise phylogenetic position within Sapindaceae is not
strongly supported (BS < 50; ig. 1A). However, both the MP
and ML analyses presented here clearly point toward Xanthoceras comprising a basal lineage with Sapindaceae s. lat.
(ig. 1A). Moreover, a study comparing the performance of
supertree methods based on an identical dataset (Buerki et
al. in press) produced the same result, with both the Matrix
Representation with Parsimony and MinFlip supertree methods placing Xanthoceras as the most basally branching lineage. This phylogenetic pattern might be explained either by
a higher rate of extinction in the lineage that now comprises
only Xanthoceras than in the other lineages, or alternatively
by a rapid diversiication or radiation of these other lineages
resulting in a loss of phylogenetic signal (Judd & Olmstead
2004). In the case of Sapindaceae s. lat., the former hypothesis seems more likely based on preliminary divergence time
estimations that place the origin of the clade in the Late Cretaceous (c. 110 My), with divergence among the four lineages
occurring between 90 and 80 My (Buerki et al. in prep.).
The pattern observed here, in which resolution between
lineages remains problematic even after sequencing a large
number of markers from a broad sampling of taxa, has been
observed in many other angiosperm groups, especially among
the rosids (Bello et al. 2009 and references within), such as
Fabales, where the relationships among the currently accepted families remain unsolved. In order to clarify the situation
within Sapindaceae s. lat. and provide a practical classiica5
Pl. Ecol. Evol. FastTrack, 2010
Figure 2 – Relationships within subfamily Sapindoideae. Bootstrap supports are indicated above branches. CUP = Cupanieae; KOE =
Koelreuterieae; HAR = Harpullieae; MEL = Melicocceae; NEP = Nephelieae; SCH = Schleichereae. The deinition of the groups follows
Buerki et al. (2009).
6
Buerki et al., Recognition of a new family, Xanthoceraceae
tion that circumscribes easily recognizable groups, we suggest that other criteria should be considered in addition to
monophyly. In addition to representing the most basal lineage
within the family, Xanthoceras presents a unique and highly
distinctive combination of morphological characters, including imparipinnate leaves (vs. paripinnate, evergreen leaves in
most genera), large lowers with petals c. 1.5–2 cm long (vs.
small lowers with petals < 1.5 cm long), 5-horn-like appendages protruding from the nectary disk (vs. no appendages
protruding from the disk), 7–8 ovules per locule (vs. generally 1–2 ovules per locule) and > 15 seeds (vs. 1–3 seeds).
Moreover, if Xanthoceras is included within Sapindaceae s.
str., it stands out as the sole member with a north-temperate
distribution, whereas all other genera occur in the tropics and/
or subtropics.
The tropical Chinese genus Delavaya, traditionally assigned to Harpullieae, has been viewed by several authors
(e.g. Müller & Leenhouts 1976, Cronquist 1988) as a “link”
between Sapindaceae and Hippocastanaceae through the
temperate genus Handeliodendron. The molecular analyses
presented here failed to conirm this hypothesis (igs 1 & 2;
see below). Instead, they indicate that Handeliodendron belongs to the Hippocastanaceae clade, a placement previously
suggested by Forest et al. (2001) based on the presence of
simple, opposite leaves, whereas Delavaya occupies a basal
position within subfam. Sapindoideae along with Ungnadia
from Texas and Florida, another genus originally assigned
to Harpullieae (these two genera thus forming the Delavaya
group; ig. 2). As indicated above, a close relationship between Delavaya and Ungnadia was previously suggested
by Klaassen (1999) and Judd et al. (1994) based on wood
anatomy and morphological cladistic analyses, respectively.
A majority (ive out of nine) of the genera traditionally assigned to Harpullieae (viz., Arfeuillea, Eurycorymbus, Harpullia, Magonia and Majidea) belong to subfam. Dodonaeoideae, and in particular to the Dodonaea group, a inding that
is consistent with Radlkofer’s (1890, 1933) original placement of Harpullieae. The basal most branch of the Dodonaea
group includes the South American genus Magonia (ig. 1C),
associated by Müller & Leenhouts (1976) with the temperate
Asian Xanthoceras on the basis of their sharing seven or eight
ovules per locule. The close relationship between Averrhoidium, Diplokeleba (previously assigned to Sapindoideae) and
Magonia might be relected in part by seed morphology; the
irst two genera are the only members of Sapindaceae to have
winged seeds (Radlkofer 1933).
Finally, the results of the phylogenetic analyses presented
in this study are in agreement with the indings of Buerki et
al. (2009) with regard to the position of the last genus traditionally assigned to Harpullieae, the Malagasy endemic Conchopetalum, which is conirmed to belong to the Macphersonia group (Sapindoideae; ig. 2; see Buerki et al. in press
for more details). Relationships between this taxon and other
members of Sapindaceae are discussed in Buerki et al. (2009).
Harpullieae have traditionally been considered to represent a “link” between Aceraceae, Hippocastanaceae and
Sapindaceae, but as mentioned above, the molecular analyses
presented here fail to conirm this hypothesis (igs 1 & 2).
Our analyses do not support the long-held view that Acera-
ceae and Sapindaceae are closely related (Radlkofer 1890,
1933, Müller & Leenhouts 1976, Umadevi & Daniel 1991).
Instead, they show that the two genera currently placed in
Aceraceae form a strongly supported group, and that they are
more closely related to Hippocastanaceae than to the clade
comprising Sapindaceae s. str. (ig. 1B), as earlier suggested
by Harrington et al. (2005), Thorne & Reveal (2007) and
Buerki et al. (2009).
The present analyses further conirm that (i) Sapindaceae
s. lat. constitute a monophyletic entity that is supported by
molecular (but not morphological) synapomorphies; (ii) the
three traditionally recognized families Aceraceae, Hippocastanaceae and Sapindaceae, as circumscribed by Radlkofer
(1933), are each monophyletic and moderately to strongly
supported, provided that Xanthoceras is excluded from Sapindaceae; and (iii) Xanthoceras sorbifolium is sister to the
clade comprising these three families (ig. 1A). The concept
of a broadly deined Sapindaceae that includes Aceraceae,
Hippocastanaceae and Xanthoceras, recently adopted by the
Angiosperm Phylogeny Group (APG II 2003) and followed
by Harrington et al. (2005), Buerki et al. (2009) and APG
III (2009), is consistent with the phylogenetic relationships
revealed in earlier studies and conirmed here. However, this
broad circumscription of Sapindaceae presents several conceptual problems. First, no clear morphological synapomorphies have been identiied for Sapindaceae s. lat. (Harrington
et al. 2005, Thorne & Reveal 2007, Buerki et al. 2009) and
the high level of heterogeneity that results from the inclusion
of Xanthoceras and the taxa traditionally placed in Aceraceae
and Hippocastanaceae makes it dificult to characterize the
family. Second, treating Sapindaceae broadly reduces these
easily identiied and widely recognized families to synonymy, changing the long-established family assignment of several well known, emblematic and widely cultivated genera,
most notably Acer and Aesculus.
Classiication
Two alternative approaches are available to address the family level circumscription of the taxa currently placed in Sapindaceae s. lat.: (i) retain the broad deinition recently proposed
by the Angiosperm Phylogeny Group (APG II 2003, APG III
2009) or (ii) resurrect the temperate families Aceraceae and
Hippocastanaceae, restrict Sapindaceae s. str. slightly by excluding Xanthoceras, and describe a new family to accommodate this genus. Both interpretations are consistent with
the primary principle of classiication as deined by Backlund
& Bremer (1998), which requires the monophyly of taxonomic entities. However, the second approach is clearly preferable when two other principles proposed by these authors
are taken into consideration, maximizing ease of identiication and maintaining nomenclatural stability. While an argument could be made that it is preferable to avoid adding a new
name at the family rank (Stevens 1997), in the present case
we believe that this is signiicantly outweighed by the clear
advantages of maintaining Aceraceae, Hippocastanaceae and
Sapindaceae (excluding Xanthoceras) as the morphologically
and biogeographically coherent entities that have been recognized for well over a century. In order to render Sapindaceae
7
Pl. Ecol. Evol. FastTrack, 2010
Key to distinguish Xanthoceraceae from closely related families
Leaves alternate, compound (rarely unifoliate in some Allophylus and Dodonaea species, both
Sapindaceae)..........................................................................................................................................2
1’. Leaves opposite, simple or compound......................................................................................................3
2. Flower large, petals c. 2 cm long; disc with 5-horn-like appendages; ovules 7 or 8 per locule, all fertile;
seeds > 15 per fruit; leaves imparipinnately compound, plants deciduous; temperate regions of eastern
Asia, from inner Mongolia across China to Korea........................................................Xanthoceraceae
2’. Flower small, petals < 1.5 cm long; disc lacking horn-like appendages; ovules 1 or 2 per locule (7 or
8 in the South American genus Magonia), fertile ovule generally 1 per locule; seeds 1 to 3 per fruit;
leaves paripinnately compound (rarely imparipinnate, unifoliolate or simple), plants evergreen; tropical
to subtropical regions........................................................................................................Sapindaceae
3. Flower actinomorphic, leaves palmately lobed or pinnately compound; fruit a schizocarpic fruits with
1-seeded samaroid mericarps..................................................................................................Aceraceae
3’. Flower zygomorphic, leaves palmately compound; fruit a 3-carpellate, usually 1-seeded
capsule.......................................................................................................................Hippocastanaceae
1.
s. str. monophyletic, we therefore propose to place Xanthoceras in a new monotypic family, Xanthoceraceae.
Taxonomy
Xanthoceraceae Buerki, Callm. & Lowry fam. nov. ≡ Sapindaceae subfam. Xanthoceroideae Thorne & Reveal
(Thorne & Reveal 2007: 119).
Haec familia ab Aceraceis Hippocastanaceis et Sapindaceis
lore grandi (petalis c. 2 cm longis) appendicibus 5 corniformibus e disco nectarifero exsertis, ovulis (omnibus fertilibus) in quoque loculo 6 ad 8 atque seminibus exarillatis
plus quam 15 distinguitur. – Type: Xanthoceras sorbifolium
Bunge (von Bunge 1833: 85).
Shrubs or small trees, andromonoecious. Leaves imparipinnately compound; lealets serrate; bracts ovate, large.
Flowers in dense racemes 15–20 cm long, staminate and
perfect lowers borne in separate inlorescences, all actinomorphic. Sepals 5, oblong, imbricate. Petals 5, large, c. 2
cm long, broadly obovate, shortly clawed at base, scale absent. Disk 5-lobed, lobes alternate to petals, apex with ive
suberect, obtuse, abaxial horn-like appendage. Stamens 8,
not exserted; anthers ellipsoid, gland-tipped. Ovary ellipsoid,
3-locular; ovules 7 or 8 per locule, in 2 lines, all fertile; style
terminal, erect; stigma papillate. Fruit a globose to pyriform
capsule, 3-ridged, pericarp very thick, dehiscing loculicidally
into 3 schizocarps. Seeds numerous (> 15), compressed-globose, arillode absent. 2n = 30.
Included genus: Xanthoceras Bunge.
Distribution and habitat – Xanthoceras sorbifolium is
widespread in north temperate China, where it grows on hills
and slopes in Gansu, Hebei, Henan, Nei Mongol, Ningxia,
Shaanxi, Shandong and Shanxi provinces; it is also found in
Korea (Xia & Gadek 2007).
Etymology – Von Bunge (1833) published the name Xanthoceras in reference to the unique yellow horn-like appendages found on the loral disc in the sole member of this genus,
X. sorbifolium, and coined a common name in English, yel8
lowhorn. In Chinese Xanthoceras is referred to as wen guan
guo (Xia & Gadek 2007).
Ethnobotany – Xanthoceras is used as a folk medicine in
China for treating various diseases (Chan & Mak 2006) as
well as enuresis in children (Chan et al. 2008). Its seeds are
sweet and are prized as a food. Acylated triterpene saponins
were recently isolated from the husk of X. sorbifolium (Chan
et al. 2008) and exhibited cytotoxicity towards several human
cancer cell lines (Chan & Mak 2006).
Notes – The new family Xanthoceraceae can be distinguished
from Aceraceae, Hippocastanaceae and Sapindaceae (as recircumscribed here) by a number of morphological features,
as summarized in the key presented above.
SUPPLEMENTARY DATA
Supplementary data are available at Plant Ecology and evolution, Supplementary Data Site (http://www.ingentaconnect.
com/content/botbel/plecevo/supp-data), and consist of a tabular version of the appendix (pdf format).
ACKNOWLEDGMENTS
We are grateful to the following persons and institutions for
supplying plant material: the Conservatoire et Jardin botaniques de la Ville de Genève (Switzerland); the Malagasy
staff of the Missouri Botanical Garden in Antananarivo
(Madagascar); Caroline Weckerle (University of Zürich,
Switzerland); Laura Vary (University of California-Irvine,
USA) and Jean-Noël Labat (Muséum National d’Histoire
Naturelle, Paris, France). The irst author is also grateful to
Yong-Ming Yuan (University of Neuchâtel, Switzerland) for
providing laboratory support and plant material. We thank
Jérôme Munzinger (Laboratoire de Botanique, Centre IRD
de Nouméa, New Caledonia), Roy E. Gereau, Nicholas Turland, Peter B. Phillipson (Missouri Botanical Garden) and
Félix Forest (Royal Botanic Gardens, Kew) for valuable discussions and/or specimen identiications. The irst author received inancial support from the Laboratory of Evolutionary
Buerki et al., Recognition of a new family, Xanthoceraceae
Botany (University of Neuchâtel, Switzerland) and a postdoctoral fellowship awarded by the Swiss National Science
Foundation (n° PBNEP3-129903).
REFERENCES
Angiosperm Phylogeny Group (APG II) (2003) An update of the
Angiosperm Phylogeny Group classiication for the orders and
families of lowering plants: APG II. Botanical Journal of the
Linnean Society 141: 399–436.
Angiosperm Phylogeny Group (APG III) (2009) An update of the
Angiosperm Phylogeny Group classiication for the orders and
families of lowering plants: APG III. Botanical Journal of the
Linnean Society 161: 105–121.
Backlund A., Bremer, K. (1998) To be or not to be – principles of
classiication of monotypic families. Taxon 47: 391–400.
Bello M.A., Bruneau A., Forest F., Hawkins J.A. (2009) Elusive
relationships within order Fabales: phylogenetic analyses using
matK and rbcL sequence data. Systematic Botany 34: 102–114.
Buerki S., Forest F., Acevedo-Rodríguez P., Callmander M.W., Nylander J.A.A.,Harrington M., Sanmartín I., Küpfer P., Alvarez
N. (2009) Plastid and nuclear DNA markers reveal intricate relationships at subfamilial and tribal levels in the soapberry family (Sapindaceae). Molecular Phylogenetics and Evolution 51:
238–258.
Buerki S., Phillipson P.B., Lowry II P.P., Callmander M.W. (2010)
Molecular phylogenetics and morphological evidence support
recognition of Gereaua, a new endemic genus of Sapindaceae
from Madagascar. Systematic Botany 35: 172–180.
Buerki S., Forest F., Salamin, N., Alvarez N. (in press) Comparative performance of supertree algorithms in large datasets using
the soapberry family (Sapindaceae) as a case study. Systematic
Biology.
Chan P.-K., Mak E. (2006) A novel compound isolated from Xanthoceras sorbifolia inhibits ovarian cancer: Identiication of
diangeloyl groups in saponin essential for anti-tumor activity.
Proceedings of the American Association for Cancer Research
47: 452–453.
Chan P.-K., Zhao M., Che C.-T., Mak E. (2008) Cytotoxic acylated
triterpene saponins from the husks of Xanthoceras sorbifolia.
Journal of Natural Products 71: 1247–1250.
Cronquist A. (1988) The evolution and classiication of lowering
plants, Ed. 2. New York, New York Botanical Garden Press.
Dahlgren G. (1989) An updated system of classiication. Botanical
Journal of the Linnean Society 100: 197–203.
Felsenstein J. (1985) Conidence-limits on phylogenies – an approach using the bootstrap. Evolution 39: 783–791.
Forest F., Drouin J. N., Charest R., Brouillet L., Bruneau A. (2001)
A morphological phylogenetic analysis of Aesculus L. and Billia Peyr. (Sapindaceae). Canadian Journal of Botany 79: 154–
169.
Gadek P.A., Fernando E.S., Quinn C.J., Hoot S.B., Terrazas T.,
Sheahan M.C., Chase M.W. (1996) Sapindales: molecular delimitation and infraordinal groups. American Journal of Botany
83: 802–811.
Harrington M.G., Edwards K.J., Johnson S.A., Chase M.W., Gadek
P.A. (2005) Phylogenetic inference in Sapindaceae s. lat. using
plastid matK and rbcL DNA sequences. Systematic Botany 30:
366–382.
Judd W.S., Olmstead R.G. (2004) A survey of tricolpate (eudicot)
phylogenetic relationships. American Journal of Botany 91:
1627–1644.
Judd W.S., Sanders R.W., Donoghue M.J. (1994) Angiosperm family pairs: preliminary phylogenetic analyses. Harvard Papers in
Botany 5: 1–51.
Klaassen R. (1999) Wood anatomy of the Sapindaceae. International Association of Wood Anatomists Journal 2 (suppl.): 1–214.
Müller J., Leenhouts P.W. (1976) A general survey of pollen types
in Sapindaceae in relation to taxonomy. In: Ferguson I.K.,
Müller J. (eds) The evolutionary signiicance of the exine: 407–
445. London, Academic Press.
Nixon K.C. (1999) The Parsimony Ratchet, a new method for rapid
parsimony analysis. Cladistics 15: 407–414.
Posada D., Crandall K.A. (1998) Modeltest: testing the model of
DNA substitution. Bioinformatics 14: 817–818.
Radlkofer L. (1890) Ueber die Gliederung der Familie der Sapindaceen. Sitzungsberichte der Königl. Bayerischen Akademie der
Wissenschaften zu München 20: 105–379.
Radlkofer L. (1933) Sapindaceae. In: Engler A. (ed.) Das Planzenreich: Regni Vegetabilis Conspectus (IV) 165 (Heft 98ah).
Leipzig, Verlag von Wilhelm Engelmann.
Rehder A. (1935) Handeliodendron, a new genus of the Sapindaceae. Journal of the Arnold Arboretum 16: 65–67.
Savolainen V., Fay M.F., Albach D.C., Backlund A., van der Bank
M., Cameron K.M., Johnson S.A., Lledó M.D., Pintaud J.-C.,
Powell M., Sheahan M.C., Soltis D.E., Soltis P.S., Weston P.,
Whitten W.M., Wurdack K.J., Chase M.W. (2000) Phylogeny of
the eudicots: a newly complete familial analysis based on rbcL
gene sequences. Kew Bulletin 55: 257–309.
Sikes D.S., Lewis P.O. (2001) PAUPRat: PAUP* implementation
of the parsimony ratchet. beta software, v. 1. Distributed by the
authors. Storrs, Department of Ecology and Evolutionary Biology, University of Connecticut.
Stamatakis A. (2006) RAxML-VI-HPC: Maximum Likelihoodbased phylogenetic analyses with thousands of taxa and mixed
models. Bioinformatics 22: 2688–2690.
Stamatakis A., Hoover P., Rougemont J. (2008) A rapid bootstrap
algorithm for the RAxML web-servers. Systematic Biology 57:
758–771.
Stevens P.F. (1997) What kind of classiication should the practising
taxonomist use to be saved? In: Dransield J., Coode M.J.E.,
Simpson D.A. (eds) Plant diversity in Malesia III, Proceedings
of the third international Flora Malesiana symposium: 295–320.
Kew, Royal Botanical Garden.
Swofford D.L. (2002) PAUP*: Phylogenetic analysis using parsimony (*and other methods), v. 4. Sunderland, Massachusetts,
Sinauer Associates.
Takhtajan A. (1987) Systema Magnoliophytorum. Leningrad, Soviet Sciences Press.
Thorne R.F., Reveal J.L. (2007) An update classiication of the
class Magnoliopsida (“Angiospermae”). Botanical Review 73:
67–182.
Umadevi I., Daniel M. (1991) Chemosystematics of the Sapindaceae. Feddes Repertorium 102: 607–612.
von Bunge A.A. (1833) Enumeratio plantarum, quas in China Boreali collegit. Mémoires de l’Académie Impériale des Sciences
de Saint Pétersbourg 2: 75–147.
Xia N., Gadek PA. (2007) Sapindaceae. In: Zhengyi W., Raven
P.H., Deyuan, H. (eds) Flora of China: 5–24. Beijing, Science
Press; St. Louis, Missouri Botanical Garden Press.
Manuscript received 28 Mar. 2010; accepted in revised version 14
Jun. 2010.
Communicating Editor: Steven Dessein.
9
Pl. Ecol. Evol. FastTrack, 2010
Appendix – Voucher information and GenBank accession numbers for taxa used in the phylogenetic analysis of Sapindaceae s. lat. (including outgroups).
For each taxon, voucher information (between brackets) is listed in the following order: voucher collector and collection number, institution where the voucher
was deposited, country. Accession numbers are listed in the following order: ITS, matK, rpoB, trnD-trnT, trnK-matK, trnL, trnL-F, trnS-trnG. Hyphens indicate
that no data are available. The entire table is available as an electronic appendix (see Supplementary Data).
Ingroup: Acer campestre L. (s.n., -, Spain) -, AJ438796, -, -, AJ438796, -, -, -; Acer cissifolium (Siebold & Zucc.) K.Koch (-, -, -) AF241483, -, -, -, -, -, -, -; Acer
erianthum Schwer. (Chase 19983, K, China) EU720501, -, EU720843, EU720980, -, EU721271, EU721459, -; Acer glabrum Torr. (Morris Arb. 93-277B, MO,
-) -, -, -, -, -, DQ978532, DQ978532, -; Acer henryi Pax (-, -, -) AF401141, -, -, -, -, -, -, -; Acer opalus Mill. (Grimm GG01305, -, -) AM238317, -, -, -, -, -, -, -;
Acer platanoides L. (-, -, -) -, AJ438788, -, -, AJ438788, -, -, -; Acer saccharum Marshall (Chase 106, K, Cult. source, Orange Co.) EU720502, -, EU720844, -,
-, EU721272, EU721460, -; Aesculus chinensis Bunge (Xiang 305, -, -) -, AY724267, -, -, -, -, -, -; Aesculus lava Sol. (98-48, -, -) -, AY968670, -, -, -, -, -, -;
Aesculus glabra Willd. (J.C. Raulston Accn. No. 960612, -, -) -, AY968671, -, -, -, -, -, -; Aesculus hippocastanum L. (living collection 6911289263, -, -) -,
AY724266, -, -, -, -, -, -; Aesculus indica (Wall. ex Cambess.) Hook. (Chase 19987, K, India) EU927392, -, EU720845, EU720981, -, EU721273, EU721461, -;
Aesculus pavia Castigl. (Chase 503, K, -) -, -, EU720846, EU720982, -, EU721274, EU721462, -; Aesculus wangii Hu (-, -, -) -, -, -, -, -, AF411085, AF411085,
-; Alectryon connatus Radlk. (Chase 2047, K, Australia) EU720415, EU720577, EU720732, EU720928, EU721025, EU721169, EU721357, EU721534;
Alectryon coriaceus Radlk. (Edwards KE59, JCT, Australia) -, EU720599, EU720756, -, EU721039, EU721192, EU721380, EU721548; Alectryon excelsus
Gaertn. (-, -, -) EF635451, -, -, -, -, -, -, -; Allophyllus arboreus Choux (Rakotovao 2812, MO, Madagascar) EU720508, -, EU720852, -, -, -, -, -; Allophyllus
arboreus Choux (Ravelonarivo 1618, MO, Madagascar) EU720515, EU720665, EU720859, -, EU721102, -, -, -; Allophyllus arboreus Choux (Rakotovao 3131,
MO, Madagascar) EU720531, EU720681, EU720879, -, EU721118, -, -, -; Allophyllus cobbe (L.) Rauesch (Callmander 462, MO, Madagascar) EU720517, -,
EU720861, -, -, -, -, -; Allophyllus longipes Radlk. (-, -, -) -, -, -, -, -, AY207572, AY207572, -; Allophyllus natalensis De Winter (Edwards KE227, JCT, South
Africa) -, AY724268, -, -, -, -, -, -; Allophyllus sp. (Edwards KE273, JCT, Tanzania) -, AY724269, -, -, -, -, -, -; Allophyllus trichodesmus Radlk. (Rakotovao 2897,
MO, Madagascar) EU720511, -, EU720855, -, -, -, -, -; Allophyllus trichodesmus Radlk. (Ravelonarivo 1619, MO, Madagascar) EU720516, -, EU720860, -, -,
-, -, -; Amesiodendron chinensis (Merr.) Hu (Yuan s.n., NEU, China) EU720403, -, EU720718, EU720917, -, EU721155, EU721344, EU721525; Arfeuillea
arborescens Pierre (Chase 2122, K, Bogor, BG) EU720461, EU720629, EU720793, EU720962, EU721067, EU721229, EU721417, -; Arytera divaricata
F.Muell. (Edwards KE010, JCT, Australia) -, AY724271, -, -, -, -, -, -; Arytera littoralis Blume (Yuan s.n., NEU, China) EU720405, EU720566, EU720720,
EU720919, EU721018, EU721157, EU721346, EU721527; Arytera littoralis Blume (Chase 2123, K, Bogor, BG) EU720462, EU720630, EU720794,
EU720963, EU721068, EU721230, EU721418, -; Arytera microphylla (Benth.) Radlk. (Edwards KE60, JCT, Australia) -, AY724270, -, -, -, -, -, -; Atalaya alata
(Sim) H.Forbes (Edwards KE228, JCT, South Africa) EU720425, EU720593, EU720748, EU720939, EU721036, EU721184, EU721372, EU721543; Atalaya
angustifolia S.T.Reynolds (West 5349, ANH, Australia) -, EU720569, EU720723, -, -, EU721160, EU721348, -; Atalaya capense R.A.Dyer (Edwards KE509,
JCT, South Africa) EU720429, -, EU720752, -, -, EU721188, EU721376, -; Atalaya salicifolia (DC.) Blume (Edwards KE58, JCT, Australia) -, AY724272, -, -,
-, -, -, -; Athyana weinmannifolia (Griseb.) Radlk. (Pennington 17581, MO, Peru) EU720487, EU720649, EU720824, EU720975, EU721086, EU721257,
EU721445, EU721576; Averrhoidium dalyi Acev.-Rodr. & Ferrucci (Weckerle 00/03/18-1/1, Z, Peru) EU720495, -, EU720836, -, -, EU721268, EU721456, -;
Beguea apetala Capuron (Buerki 149, NEU, Madagascar) EU720491, EU720652, EU720828, EU720978, EU721089, EU721261, EU721449, -; Beguea apetala
Capuron (Vary 40, MO, Madagascar) EU720512, EU720663, EU720856, -, EU721100, EU721281, EU721469, -; Billia sp. (Hammel 20075, -, -) -, AY724275,
-, -, -, -, -, -; Blighia sapida K.D.Koenig (Edwards KE86, JCT, West Africa) EU720416, EU720578, EU720733, EU720929, EU721026, EU721170, EU721358,
EU721535; Blighia unijugata Baker (Edwards KE274, JCT, Tanzania) -, AY724276, -, -, -, -, -, -; Blomia prisca (Standl.) Lundell (Acevedo 12242, US, Mexico,
Yucatán) EU720444, EU720611, EU720772, -, EU721050, EU721208, EU721396, -; Bridgesia incisifolia Bertero ex Cambess. (Killip & Pisano 39778, K,
Chile) EU720476, EU720645, EU720811, EU720973, EU721082, EU721247, EU721435, -; Cardiospermum grandilorum Sw. (Edwards KE207, JCT, -) -,
EU720588, EU720743, EU720935, -, EU721179, EU721367, -; Cardiospermum microcarpum Kunth (Yuan s.n., NEU, China) -, -, EU720712, EU720911, -,
EU721149, EU721338, -; Cardiospermum sp. (Yuan s.n., NEU, China) EU720399, -, EU720713, EU720912, -, EU721150, EU721339, -; Castanospora
alphandii (F.Muell.) F.Muell. (Edwards KE88, JCT, Australia) -, AY724279, -, -, -, -, -, -; Chytranthus carneus Radlk. (Chase 2868, RBG, -) EU720477,
EU720646, EU720812, EU720974, EU721083, EU721248, EU721436, EU721575; Chytranthus prieurianus Baill. (Edwards KE272, JCT, Tanzania) -,
AY724280, -, -, -, -, -, -; Conchopetalum brachysepalum Capuron (Rabarimanarivo 8, MO, Madagascar) EU720530, EU720680, EU720877, -, EU721117,
EU721299, EU721487, EU721586; Cossinia pinnata Comm. ex Lam. (Lorence 4510, MO, Mauritius) -, -, EU720820, -, -, EU721253, EU721441, -; Cubilia
cubili (Blanco) Adelb. (Chase 2125, K, Bogor, BG) EU720463, EU720631, EU720795, EU720964, EU721069, EU721231, EU721419, EU721567; Cupania
dentata DC. (Acevedo 12241, US, Mexico, Yucatán) EU720523, EU720670, EU720867, EU720988, EU721107, EU721289, EU721477, EU721581; Cupania
guatemalensis (Turcz.) Radlk. (Davidse 35743, MO, Belize) -, EU720678, EU720875, EU720993, EU721115, EU721297, EU721485, -; Cupania hirsuta Radlk.
(Acevedo 1101, US, French Guiana) EU720521, EU720668, EU720865, -, EU721105, EU721287, EU721475, -; Cupania rubiginosa (Poir.) Radlk. (Mori 8868,
MO, French Guiana) EU720481, -, EU720817, -, -, EU721251, EU721439, -; Cupania scrobiculata Rich. (Acevedo 11100, US, French Guiana) EU720524,
EU720671, EU720868, EU720989, EU721108, EU721290, EU721478, -; Cupaniopsis anacardioides Radlk. (Chase 217, K, Australia) EU720438, EU720605,
EU720763, EU720946, EU721045, EU721199, EU721387, EU721552; Cupaniopsis lagelliformis (Bailey) Radlk. (Edwards KE42, JCT, Australia) EU720432,
EU720598, EU720755, EU720942, -, EU721191, EU721379, EU721547; Cupaniopsis fruticosa Radlk. (Munzinger 564, MO, New Caledonia) EU720533, -,
EU720881, -, EU721119, EU721302, EU721490, -; Cupaniopsis sp. (Munzinger 710, MO, New Caledonia) EU720532, -, EU720880, EU720996, -, EU721301,
EU721489, EU721587; Cupaniopsis sp. (Munzinger 1103, MO, New Caledonia) EU720507, EU720660, EU720851, -, EU721097, EU721278, EU721466, -;
Deinbollia borbonica Scheff. (Edwards KE197, JCT, Tanzania) EU720412, EU720574, EU720729, -, -, EU721166, EU721354, EU721532; Deinbollia
macrocarpa Capuron (Antilahimena 4293, MO, Madagascar) -, EU720626, EU720790, -, EU721064, EU721226, EU721414, EU721565; Deinbollia macrocarpa
Capuron (H. Razaindraibe 118, MO, Madagascar) EU720535, EU720683, EU720883, -, EU721121, EU721304, EU721492, EU721589; Deinbollia macrocarpa
Capuron (Buerki 144, NEU, Madagascar) EU720503, EU720656, EU720847, -, EU721093, EU721275, EU721463, -; Deinbollia oblongifolia (E.Mey. ex Arn.)
Radlk. (Edwards KE233, JCT, South Africa) EU720427, EU720595, EU720750, -, -, EU721186, EU721374, EU721545; Deinbollia pervillei (Blume) Radlk.
(Phillipson 5919, MO, Madagascar) EU720395, EU720560, EU720708, -, EU721012, EU721145, EU721334, -; Deinbollia pervillei (Blume) Radlk.
(Callmander 688, MO, Madagascar) EU720514, -, EU720858, -, -, EU721283, EU721471, -; Delavaya yunnanensis Franch. (Forrest 20682, MO, China,
Yunnan) EU720484, -, EU720821, -, -, EU721254, EU721442, -; Diatenopteryx sorbifolia Radlk. (Zardini 43371, MO, Paraguay) EU720534, EU720682,
EU720882, -, EU721120, EU721303, EU721491, EU721588; Diatenopteryx sorbifolia Radlk. (Tressens 3504, K, Argentina) -, -, EU720810, -, -, EU721246,
EU721434, -; Dictyoneura obtusa Blume (Edwards KE142, JCT, Australia) EU720428, -, EU720751, -, -, EU721187, EU721375, -; Dilodendron bipinnatum
Radlk. (Acevedo 11129, US, Bolivia) -, EU720677, EU720874, -, EU721114, EU721296, EU721484, EU721584; Dimocarpus australianus Leenh. (Edwards
KE34, JCT, Australia) EU720433, -, EU720757, -, -, , EU721381, -; Dimocarpus longan Lour. (Yuan s.n., NEU, China) -, EU720563, EU720714, EU720913,
EU721015, EU721151, EU721340, EU721521; Dimocarpus longan Lour. (Edwards KE502, JCT, Asia) -, EU720590, EU720745, -, -, EU721181, EU721369,
-; Dimocarpus longan Lour. (-, -, Thailand) EF532337, -, -, -, -, -, -, -; Dimocarpus longan Lour. (Chase 1351, K, Bogor, BG) -, EU720615, EU720777,
EU720953, EU721053, EU721213, EU721401, EU721559; Diploglottis campbelli Cheel (Chase 2048, K, Australian, BG) EU720457, EU720624, EU720788,
EU720960, EU721062, EU721224, EU721412, -; Diploglottis diphyllostegia (F.Muell.) Radlk. (Edwards KE001, JCT, Australia) -, AY724287, -, -, -, -, -, -;
Mischarytera sp. (Edwards KE159, JCT, Australia) EU720417, EU720579, EU720734, EU720930, EU721027, EU721171, EU721359, -; Diploglottis smithii
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Buerki et al., Recognition of a new family, Xanthoceraceae
S.T.Reynolds (BG838, BBG, Australia) -, AY724288, -, -, -, -, -, -; Diplokeleba loribunda N.E.Br. (Acevedo 11130, US, Bolivia, St Cruz) -, -, EU720773,
EU720950, -, EU721209, EU721397, -; Diplopeltis huegelii Endl. (Chase 2192, K, Australia) EU720473, EU720642, EU720807, EU720971, EU721079,
EU721243, EU721431, -; Dipteronia sinensis Oliv. (Chase 502, RBG, -) EU720445, EU720612, EU720774, -, -, EU721210, EU721398, -; Dodonaea lanceolata
F.Muell. (Edwards KE120, JCT, Australia) -, AY724290, -, -, -, -, -, -; Dodonaea madagascariensis Radlk. (Bocksberger GB028, NEU, Madagascar) EU720518,
-, EU720862, EU720984, -, EU721284, EU721472, -; Dodonaea viscosa Jacq. (Razaitsalama 956, MO, Madagascar) EU720519, EU720666, EU720863,
EU720985, EU721103, EU721285, EU721473, -; Dodonaea viscosa Jacq. (Merello 1077, MO, Peru) EU720536, EU720684, EU720884, EU720997, EU721122,
EU721305, EU721493, -; Dodonaea viscosa Jacq. (Yuan s.n., NEU, China) EU720406, EU720567, EU720721, EU720920, EU721019, EU721158, EU721347,
-; Doratoxylon chouxii Capuron (Labat JNL3543, P, Madagascar) EU720394, EU720559, EU720707, EU720908, EU721011, EU721144, EU721333, -;
Doratoxylon chouxii Capuron (Callmander 679, MO, Madagascar) EU720513, EU720664, EU720857, -, EU721101, EU721282, EU721470, -; Elattostachys
apetala Radlk. (Munzinger 692, MO, New Caledonia) EU720537, EU720685, EU720885, EU720998, EU721123, EU721306, EU721494, EU721590;
Elattostachys apetala Radlk. (McPherson 18184, MO, New Caledonia) EU720538, EU720686, EU720886, EU720999, EU721124, EU721307, EU721495,
EU721591; Elattostachys megalantha S.T.Reynolds (Irvine IRV507, CSIRO, Atherton) -, EU720609, EU720768, -, EU721048, EU721204, EU721392, -;
Elattostachys microcarpa S.T.Reynolds (Edwards KE98, JCT, Australia) EU720409, EU720571, EU720726, -, -, EU721163, EU721351, -; Elattostachys
nervosa (F.Muell.) Radlk. (Chase 2022, K, Australian, BG) EU720455, EU720622, EU720786, EU720959, EU721060, EU721222, EU721410, EU721563;
Elattostachys sp. (Lowry 5650A, MO, New Caledonia) EU720529, EU720679, EU720876, EU720994, EU721116, EU721298, EU721486, EU721585;
Eriocoelum kerstingii Gilg ex Engl. (Merello 1586, MO, Ghana) EU720539, EU720687, EU720887, EU721000, EU721125, EU721308, EU721496, EU721592;
Eriocoelum microspermum Radlk. (Bradley 1025, MO, Gabon) EU720540, EU720688, EU720888, EU721001, EU721126, EU721309, EU721497, EU721593;
Erythrophysa aesculina Baill. (Randrianasolo 625, MO, Madagascar) -, -, -, -, -, EU721329, -, -; Euphorianthus longifolius Radlk. (Chase 2126, K, Bogor, BG)
EU720464, -, EU720796, -, -, EU721232, EU721420, -; Eurycorymbus cavalerieri (H.Lév.) Rehder & Hand.-Mazz. (Yuan s.n., NEU, China) EU720404,
EU720565, EU720719, EU720918, EU721017, EU721156, EU721345, EU721526; Filicium decipiens (Wight & Arn.) Thwaites (Chase 2128, K, Bogor, BG)
EU720466, EU720633, EU720798, -, -, EU721234, EU721422, -; Filicium longifolium (H.Perrier) Capuron (Rabenantonadro 1113, MO, Madagascar)
EU720541, -, EU720889, -, -, EU721310, EU721498, -; Filicium thouarsianum (A.DC.) Capuron (Antilahimena 5021, MO, Madagascar) EU720493, -,
EU720832, -, -, EU721265, EU721453, -; Ganophyllum falcatum Blume (Chase 2129, K, Bogor, BG) EU720467, EU720634, EU720799, -, EU721071,
EU721235, EU721423, -; Ganophyllum falcatum Blume (BH9269, CSIRO, Australia) -, -, -, -, -, EU721330, -, -; Glenniea pervillei (Baill.) Leenh.
(Andriamihajarivo 1053, MO, Madagascar) EU720490, EU720651, EU720827, EU720977, EU721088, EU721260, EU721448, -; Glenniea pervillei (Baill.)
Leenh. (Andriamihajarivo 1025, MO, Madagascar) -, EU720653, EU720829, -, EU721090, EU721262, EU721450, -; Gongrodiscus bilocularis H.Turner
(Munzinger 749, MO, New Caledonia) EU720542, EU720689, EU720890, -, EU721127, EU721311, EU721499, -; Guindilia trinervis Gilles ex Hook. (Chase
802, K, Chile) -, EU720613, EU720775, EU720951, EU721051, EU721211, EU721399, EU721557; Guioa acutifolia Radlk. (Edwards KE14, JCT, Australia)
-, AY724297, -, -, -, -, -, -; Guioa glauca Radlk. (McPherson 18230, MO, New Caledonia) EU720545, EU720692, EU720893, -, EU721130, EU721315,
EU721503, -; Guioa lasioneura Radlk. (BG1888, BBG, Australia) -, AY724298, -, -, -, -, -, -; Guioa microsepala Radlk. (Munzinger 744, MO, New Caledonia)
EU720546, EU720693, EU720894, -, EU721131, EU721316, EU721504, EU721596; Guioa semiglauca (F.Muell.) Radlk. (Chase 2058, K, Australia, BG)
EU720458, EU720625, EU720789, -, EU721063, EU721225, EU721413, -; Guioa villosa Radlk. (McPherson 18040, MO, New Caledonia) EU720544,
EU720691, EU720892, EU721003, EU721129, EU721314, EU721502, EU721595; Guioa sp. (Munzinger 945, MO, New Caledonia) EU720505, EU720658,
EU720849, -, EU721095, EU721277, EU721465, -; Handeliodendron bodinieri (H.Lév.) Rehder (QYXiang 302 / C. Ming 050923, -, -) -, AY724299, -, -, -,
EF186776, -, -; Haplocoelopsis africana F.G.Davies (Edwards KE276, JCT, Tanzania) EU720441, EU720608, EU720767, EU720949, -, EU721203, EU721391,
EU721555; Haplocoelum foliosum (Hiern) Bullock (Friis 1894, MO, Ethiopia) EU720479, -, EU720815, -, -, EU721250, EU721438, -; Haplocoelum foliosum
subsp. foliosum (Hiern) Bullock (Edwards KE195, JCT, Tanzania) EU720410, EU720572, EU720727, EU720924, -, EU721164, EU721352, EU721530;
Haplocoelum gallaense (Engl.) Radlk. (Edwards KE501, JCT, South Africa) -, EU720583, -, -, -, -, -, -; Haplocoelum perrieri Capuron (Rakotomalaza 1165,
MO, Madagascar) EU720396, -, EU720709, EU720909, -, EU721146, EU721335, EU721519; Harpullia arborea (Blanco) Radlk. (Chase 1353, K, Bogor, BG)
EU720448, -, EU720779, -, -, EU721215, EU721403, -; Harpullia cupanioides Roxb. (-, -, -) AY2075669, -, -, -, -, -, -, -; Harpullia rhyticarpa C.White &
Francis (Edwards KE003, JCT, -) -, AY724303, -, -, -, -, -, -; Hippobromus paucilorus Radlk. (Edwards KE229, JCT, -) -, AY724305, -, -, -, EU721331,
EU721517, -; Hypelate trifoliata Sw. (R. Rankin HABJ72057, K, -) -, -, EU720813, -, -, -, -, -; Jagera javanica (Blume) Blume ex Kalkman (Chase 2130, K,
Bogor, BG) EU720468, EU720635, EU720800, -, EU721072, EU721236, EU721424, EU721569; Jagera javanica subsp. australiana Leenh. (Edwards KE178,
JCT, Australia) EU720442, -, EU720769, -, -, EU721205, EU721393, EU721556; Jagera pseudorhus var. pseudorhus f. pilosiuscula Radlk. (Edwards KE41,
JCT, Australia) -, EU720606, EU720764, EU720947, EU721046, EU721200, EU721388, EU721553; Koelreuteria paniculata Laxm. (Harder 5668, MO,
Vietnam) EU720548, EU720695, EU720896, -, EU721133, EU721318, EU721506, -; Koelreuteria paniculata Laxm. (Yuan CN2006-3, NEU, China) EU720397,
EU720561, EU720710, -, EU721013, EU721147, EU721336, EU721520; Koelreuteria paniculata Laxm. (Wilson 1476, RBG, -) -, AY724308, -, -, -, -, -, -;
Koelreuteria sp. (Harder 5724, MO, Vietnam) EU720547, EU720694, EU720895, EU721004, EU721132, EU721317, EU721505, -; Laccodiscus klaineanus
Pierre ex Engl. (Walters 1269, MO, Gabon) EU720549, EU720696, EU720897, -, EU721134, EU721319, EU721507, -; Lecaniodiscus fraxinifolius Baker
(Edwards KE194, JCT, Tanzania) EU720418, EU720580, EU720735, EU720931, EU721028, EU721172, EU721360, EU721536; Lepiderema hirsuta
S.T.Reynolds (Edwards KE36, JCT, Australia) EU720435, EU720601, EU720759, -, EU721041, EU721195, EU721383, EU721549; Lepiderema pulchella
Radlk. (Chase 2020, K, Australian, BG) EU720454, -, EU720785, EU720958, -, EU721221, EU721409, -; Lepidopetalum fructoglabrum Welzen (Edwards
KE139, JCT, Australia) EU720408, -, EU720724, EU720922, -, EU721161, EU721349, EU721528; Lepisanthes alata (Blume) Leenh. (Chase 1355, K, Bogor,
BG) EU720450, EU720618, EU720781, -, EU721056, EU721217, EU721405, -; Lepisanthes feruginea (Radlk.) Leenh. (Chase 1354, K, Bogor, BG) EU720449,
EU720617, EU720780, -, EU721055, EU721216, EU721404, -; Lepisanthes rubiginosa (Roxb.) Leenh. (Chase 1350, K, Bogor, BG) EU720446, EU720614,
EU720776, EU720952, EU721052, EU721212, EU721400, EU721558; Lepisanthes senegalensis (Poir.) Leenh. (Callmander 627, MO, Madagascar) EU720492,
EU720654, EU720830, EU720979, EU721091, EU721263, EU721451, EU721577; Litchi chinensis Sonn. (Yuan s.n., NEU, China) EU720400, EU720564,
EU720715, EU720914, EU721016, EU721152, EU721341, EU721522; Llagunoa mollis Kunth (Jaramillollejia 3199, MO, Colombia) EU720482, -, EU720818,
-, -, EU721252, EU721440, -; Llagunoa nitida Ruiz & Pav. (Pennington 17552, MO, Peru) EU720486, -, EU720823, -, -, EU721256, EU721444, -; Loxodiscus
coriaceus Hook.f. (Bradford 1136, MO, New Caledonia) EU720488, -, EU720825, -, -, EU721258, EU721446, -; Macphersonia chapelieri (Baill.) Capuron
(Buerki 138, NEU, Madagascar) EU720459, EU720627, EU720791, EU720961, EU721065, EU721227, EU721415, EU721566; Macphersonia gracilis
O.Hoffm. (Rabenantoandro 1081, MO, Madagascar) EU720550, EU720697, EU720898, EU721005, EU721135, EU721320, EU721508, EU721597; Magonia
pubescens A.St.-Hil. (Mori 16966, MO, Brazil) EU720483, -, EU720819, -, -, -, -, -; Majidea zanguebarika Kirk ex Oliv. (TH275, MO, Madagascar) EU720552,
-, EU720900, EU721006, -, EU721322, EU721510, -; Matayba apetala Radlk. (Acevedo 11929, US, Jamaica) EU720526, EU720674, EU720871, -, EU721111,
EU721293, EU721481, EU721583; Matayba cf. opaca Radlk. (Acevedo 11118, US, French Guiana) EU720522, EU720669, EU720866, EU720987, EU721106,
EU721288, EU721476, EU721580; Matayba domingensis (DC.) Radlk. (Taylor 11819, MO, Caribbean) EU720551, EU720698, EU720899, -, EU721136,
EU721321, EU721509, EU721598; Matayba elaeagnoides Radlk. (Zardini 43278, MO, Paraguay) EU720553, EU720699, EU720901, -, EU721137, EU721323,
EU721511, -; Matayba guianensis Aubl. (Acevedo 12342, US, French Guiana) EU720527, EU720675, EU720872, -, EU721112, EU721294, EU721482, -;
Matayba laevigata Radlk. (Acevedo 12357, US, French Guiana) EU720528, EU720676, EU720873, EU720992, EU721113, EU721295, EU721483, -;
Melicoccus bijugatus Jacq. (Acevedo s.n., US, Puerto Rico ) EU927391, EU720610, EU720771, -, EU721049, EU721207, EU721395, -; Melicoccus lepidopetalus
11
Pl. Ecol. Evol. FastTrack, 2010
Radlk. (Acevedo 11128, US, Bolivia) EU720443, -, EU720770, -, -, EU721206, EU721394, -; Mischarytera lautereriana (F.M.Bailey) H.Turner (Edwards
KE1302, JCT, Australia) -, -, EU720742, -, -, -, -, -; Mischarytera macrobotrys (Merr. & L.M.Perry) H.Turner (BH6631, CSIRO, Australia) -, AY724313, -, -, -,
-, -, -; Mischarytera sp. (Edwards KE159, JCT, Australia) EU720417, EU720579, EU720734, EU720930, EU721027, EU721171, EU721359, -; Mischocarpus
exangulatus (F.Muell.) Radlk. (Edwards KE30, JCT, Australia) EU720434, EU720600, EU720758, EU720943, EU721040, EU721194, EU721382, -;
Mischocarpus grandissumus Radlk. (Edwards KE37, JCT, Australia) EU720437, EU720604, EU720762, EU720945, EU721044, EU721198, EU721386,
EU721551; Mischocarpus pentapetalus (Roxb.) Radlk. (Chase 2133, K, Bogor, BG) EU720470, EU720637, EU720802, EU720966, EU721074, EU721238,
EU721426, EU721571; Mischocarpus pyriformis (F.Muell.) Radlk. (Chase 2059, K, Australian, BG) EU720460, EU720628, EU720792, -, EU721066,
EU721228, EU721416, -; Molinaea petiolaris Radlk. (Rabenantoandro 1448, MO, Madagascar) EU720554, EU720700, EU720902, EU721007, EU721138,
EU721324, EU721512, -; Molinaea sp. nov. (Antilahimena 4301, MO, Madagascar) EU720510, EU720662, EU720854, EU720983, EU721099, EU721280,
EU721468, EU721578; Neotina coursii Capuron (H. Razaindraibe 119, MO, Madagascar) EU720543, EU720690, EU720891, EU721002, EU721128,
EU721313, EU721501, EU721594; Nephelium lappaceum L. (Edwards KE222, JCT, Asia ) -, EU720584, EU720738, EU720932, EU721030, EU721175,
EU721363, EU721537; Nephelium lappaceum L. (Yuan s.n., NEU, China) EU720401, -, EU720716, EU720915, -, EU721153, EU721342, EU721523;
Nephelium mutabile Blume (Chase 2134, K, Bogor, BG) -, AY724316, -, -, -, -, -, -; Pancovia golungensis (Hiern) Exell & Mendonça (Edwards KE231, JCT,
Tanzania) EU720411, EU720573, EU720728, EU720925, EU721022, EU721165, EU721353, EU721531; Pappea capensis Eckl. & Zeyh. (Edwards KE232,
JCT, South Africa) EU720424, EU720592, EU720747, EU720938, EU721035, EU721183, EU721371, EU721542; Paranephelium macrophyllum King (Chase
1356, K, Bogor, BG) EU720451, EU720619, EU720782, EU720955, EU721057, EU721218, EU721406, -; Paranephelium xestophyllum Miq. (Edwards
KE503, JCT, Asia) EU720420, EU720582, EU720737, -, EU721029, EU721174, EU721362, -; Paullinia alata subsp. alata G.Don (Weckerle 00/03/09-2/1, Z,
Peru) -, -, EU720834, -, -, -, -, -; Paullinia elegans Cambess. (Weckerle 00/05/27-1/1, Z, Peru) -, -, EU720835, -, -, EU721267, EU721455, -; Paullinia eriocarpa
Triana & Planch. (Weckerle 00/06/13-1/5, Z, Peru) EU720497, -, EU720839, -, -, -, -, -; Paullinia faginea (Triana & Planch.) Radlk. (Weckerle 00/05/27-1/5, Z,
Peru) -, -, EU720837, -, -, -, -, -; Paullinia faginea (Triana & Planch.) Radlk. (Weckerle 00/06/13-1/3, Z, Peru) EU720496, -, EU720838, -, -, -, -, -; Paullinia
pachycarpa Benth. (Weckerle 01/01/26-1/1, Z, Peru) EU720500, -, EU720842, -, -, -, -, -; Paullinia pinnata L. (Edwards KE199, JCT, Tanzania) EU720413,
EU720575, EU720730, EU720926, EU721023, EU721167, EU721355, -; Paullinia subauriculata Radlk. (Weckerle 00/03/19-1/1, Z, Peru) EU720494, -,
EU720833, -, -, EU721266, EU721454, -; Plagioscyphus aff. louvelii Danguy & Choux (Lowry 6034, MO, Madagascar) EU720555, EU720701, EU720903,
EU721008, EU721139, EU721325, EU721513, EU721599; Plagioscyphus unijugatus Capuron (Buerki 145, NEU, Madagascar) EU720475, EU720644,
EU720809, EU720972, EU721081, EU721245, EU721433, EU721574; Podonephelium homei Radlk. (McPherson 18156, MO, New Caledonia) -, -, -, -, -,
EU721312, EU721500, -; Podonephelium homei Radlk. (Pillon 156, MO, New Caledonia) EU720489, EU720650, EU720826, EU720976, EU721087,
EU721259, EU721447, -; Pometia pinnata J.R.Forst. & G.Forst. (Chase 2135, K, Bogor, BG) EU720471, EU720638, EU720803, EU720967, EU721075,
EU721239, EU721427, EU721572; Pometia pinnata J.R.Forst. & G.Forst. (Yuan s.n., NEU, China) EU720402, -, EU720717, EU720916, -, EU721154,
EU721343, EU721524; Pseudima sp. (McPherson 15867, MO, Panama) EU720556, EU720702, EU720904, EU721009, EU721140, EU721326, EU721514,
EU721600; Pseudopteris decipiens Baill. (Kakazomannjary 12529-SF, MO, Madagascar) EU720480, -, EU720816, -, -, -, -, -; Rhysotoechia mortoniana
(F.Muell.) Radlk. (Edwards KE117, JCT, Australia) EU720414, EU720576, EU720731, EU720927, EU721024, EU721168, EU721356, EU721533; Rhysotoechia
robertsonii (F.Muell.) Radlk. (Edwards KE277, JCT, Australia) -, EU720570, EU720725, EU720923, EU721021, EU721162, EU721350, EU721529; Sapindus
oligophyllus Merr. & Chun (Yuan s.n., NEU, China) EU720407, EU720568, EU720722, EU720921, EU721020, EU721159, -, -; Sapindus saponaria L. (Chase
2136, K, Bogor, BG) -, EU720639, EU720804, EU720968, EU721076, EU721240, EU721428, -; Sapindus trifoliatus L. (Edwards KE504, JCT, Asia) -,
EU720586, EU720740, EU720934, EU721032, EU721177, EU721365, EU721538; Sarcopteryx martyana (F.Muell.) Radlk. (Irvine IRV1810, CSIRO,
Australia) EU720426, EU720594, EU720749, EU720940, EU721037, EU721185, EU721373, EU721544; Sarcopteryx reticulata S.T.Reynolds (Gray BG1137,
CSIRO, Australia) EU720421, EU720587, EU720741, -, EU721033, EU721178, EU721366, EU721539; Sarcopteryx sp. (Edwards KE49, JCT, Australia)
EU720439, EU720607, EU720765, EU720948, EU721047, EU721201, EU721389, EU721554; Sarcotoechia serrata S.T.Reynolds (Edwards KE31, JCT,
Australia) EU720436, EU720603, EU720761, EU720944, EU721043, EU721197, EU721385, EU721550; Sarcotoechia villosa S.T.Reynolds (Edwards KE102,
JCT, Australia) EU720419, EU720581, EU720736, -, -, EU721173, EU721361, -; Schleichera oleosa (Lour.) Oken (Chase 2137, K, Bogor, BG) EU720423,
EU720591, EU720746, EU720937, -, EU721182, EU721370, EU721541; Scyphonichium cf. multilorum (Mart.) Radlk. (Acevedo 11102, US, French Guiana)
-, EU720672, EU720869, EU720990, EU721109, EU721291, EU721479, -; Serjania altissima (Poepp.) Radlk. (Weckerle 00/07/02-1/4, Z, Peru) EU720498, -,
EU720840, -, -, EU721269, EU721457, -; Serjania communis Cambess. (Chase 2138, K, Bogor, BG) EU720472, EU720640, EU720805, EU720969, EU721077,
EU721241, EU721429, -; Serjania glabrata Kunth (Merello 1058, MO, Peru) EU720557, EU720703, EU720905, EU721010, EU721141, EU721327,
EU721515, -; Serjania mexicana (L.) Willd. (Davidse 35748, MO, Belize) -, EU720704, EU720906, -, EU721142, -, -, -; Serjania triquetra Radlk. (-, -, -)
AY207571, -, -, -, -, -, -, -; Smelophyllum capense Radlk. (Edwards KE506, JCT, South Africa) -, AY724330, -, -, -, -, -, -; Stadmannia oppositifolia (Lam.) Poir.
(Edwards KE505, JCT, Madagascar) -, AY724331, -, -, -, -, -, -; Storthocalyx leioneurus Radlk. (Munzinger 1100, MO, New Caledonia) EU720506, EU720659,
EU720850, -, EU721096, -, -, -; Storthocalyx sp. (Munzinger 960, MO, New Caledonia) EU720504, EU720657, EU720848, -, EU721094, EU721276,
EU721464, -; Synima cordieri (F.Muell.) Radlk. (Edwards KE29, JCT, Australia) -, EU720602, EU720760, -, EU721042, EU721196, EU721384, -; Synima
macrophylla S.T.Reynolds (Edwards KE19, JCT, Australia) EU720430, EU720596, EU720753, EU720941, -, EU721189, EU721377, EU721546; Talisia
angustifolia Radlk. (Zardini 43668, MO, Paraguay) EU720558, EU720705, EU720907, -, EU721143, EU721328, EU721516, -; Talisia nervosa Radlk.
(Pennington 628, MO, -) EU720474, EU720643, EU720808, -, EU721080, EU721244, EU721432, -; Talisia obovata A.C.Sm. (R. Lombello 13, MO, Brazil)
EU720485, EU720648, EU720822, -, EU721085, EU721255, EU721443, -; Thouinia acuminata S.Watson (Liston 633-2, MO, Mexico, Jalisco) EU720478,
EU720647, EU720814, -, EU721084, EU721249, EU721437, -; Tina isaloensis Drake (Ranirison PR827, G, Madagascar) EU720520, EU720667, EU720864,
EU720986, EU721104, EU721286, EU721474, EU721579; Tina striata Radlk. (Vary 45, MO, Madagascar) EU720509, EU720661, EU720853, -, EU721098,
EU721279, EU721467, -; Tinopsis apiculata Radlk. (Buerki 131, NEU, Madagascar) EU720422, EU720589, EU720744, EU720936, EU721034, EU721180,
EU721368, EU721540; Toechima daemelianum Radlk. (JC66, CSIRO, Australia) -, AY724334, -, -, -, -, -, -; Toechima erythrocarpum (F.Muell.) Radlk. (Edwards
KE20 , JCT, Australia) EU720431, EU720597, EU720754, -, EU721038, EU721190, EU721378, -; Toechima plurinerve Radlk. (Chase 1357, K, Bogor, BG)
EU720452, EU720620, EU720783, EU720956, EU721058, EU721219, EU721407, EU721561; Toechima tenax (Cunn. ex Benth.) Radlk. (Chase 2046, K,
Australian, BG) EU720456, EU720623, EU720787, -, EU721061, EU721223, EU721411, EU721564; Toechima tenax (Cunn. ex Benth.) Radlk. (Chase 2132,
K, Bogor, BG) EU720469, EU720636, EU720801, EU720965, EU721073, EU721237, EU721425, EU721570; Tristira triptera (Blanco) Radlk. (Chase 2139,
K, Asia) -, EU720585, EU720739, EU720933, EU721031, EU721176, EU721364, -; Tristiropsis acutangula Radlk. (Chase 1358, K, Bogor, BG) EU720453,
EU720621, EU720784, EU720957, EU721059, EU721220, EU721408, EU721562; Ungnadia speciosa Endl. (Chase 2854, RBG, -) -, EU720706, -, -, -,
EU721332, EU721518, -; Urvillea ulmaceae Kunth (Weckerle 00/07/05-1/1, Z, Peru) EU720499, EU720655, EU720841, -, EU721092, EU721270, EU721458,
-; Vouarana guianensis Aubl. (Lucas 109, MO, French Guiana) EU720525, EU720673, EU720870, EU720991, EU721110, EU721292, EU721480, EU721582;
Xanthoceras sorbifolium Bunge (Yuan CN2006, NEU, China) EU720398, EU720562, EU720711, EU720910, EU721014, EU721148, EU721337, -;
Xerospermum noronhianum (Blume) Blume (Chase 2130, K, Bogor, BG) -, EU720641, EU720806, EU720970, EU721078, EU721242, EU721430, EU721573;
Outgroups: Sorindeia sp. (Buerki 137, NEU, Madagascar) -, -, EU720831, -, -, EU721264, EU721452, -; Harrisonia abyssinica Oliv. (Edwards KE510, JCT,
Tanzania) EU720440, -, EU720766, -, -, EU721202, EU721390, -; Malleastrum sp. (Rakotovao 2609, MO, Madagascar) -, -, EU720878, EU720995, -,
EU721300, EU721488, -.
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