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Swenson & al. • Chrysophylloideae in Oceania and SE Asia
Towards a natural classification of Sapotaceae subfamily
Chrysophylloideae in Oceania and Southeast Asia based on
nuclear sequence data
Ulf Swenson,1 Stephan Nylinder1 & Jérôme Munzinger2
1 Department of Phanerogamic Botany, Swedish Museum of Natural History, Box 50007, 104 05 Stockholm, Sweden
2 IRD, UMR AMAP, Montpellier, 34000 France
Author of correspondence: Ulf Swenson, ulf.swenson@nrm.se
Abstract Generic limits within subfamily Chrysophylloideae (Sapotaceae) from Oceania and Southeast Asia are reconciled
based on a molecular phylogeny. We analysed sequences of nuclear ribosomal DNA (ETS, ITS) and the nuclear gene RPB2
with BEAST and parsimony jackknifing, using a sample of 168 terminals. Eight morphological characters were traced on a
condensed majorityrule consensus tree to identify diagnostic character combinations for the genera. Accepted genera with
character support are Magodendron, Pichonia, Planchonella, Pycnandra, Sersalisia, and Van-royena, while Beccariella and
Niemeyera require amendment. Beccariella, a widely distributed group, is an illegitimate later homonym and we propose that
the genus Pleioluma is resurrected in its place. The Australian genus Niemeyera is paraphyletic, but it is rendered monophy
letic by reinstating Amorphospermum for N. antiloga. Beauvisagea, Blabeia, Fontbrunea, and Krausella are all segregates of
Planchonella and rejected, while Wokoia is a later synonym of Pichonia. Planchonella baillonii, an endemic species of New
Caledonia, is the sole member of an old lineage and firmly placed as the sister to a clade comprising the other congeners.
Planchonella sandwicensis, a Hawaiian species, previously proposed to be a distinct genus, is a member of Planchonella. In
the Pacific, P. tahitensis (including P. grayana) is a polymorphic species, widely distributed and adapted to a wide range of
habitats. We provide a generic key (excluding Xantolis), diagnostic character combinations for all genera, and the necessary
taxonomic combinations for Pichonia, Planchonella, Pleioluma, and Sersalisia to render each genus monophyletic.
Keywords Amorphospermum; Australasia; Ericales; morphology; Pacific Ocean; phylogeny; Pleioluma; Sapotaceae
Supplementary Material Aligned data matrix is available in the Supplementary Data section of the online version of this
article (http://www.ingentaconnect.com/content/iapt/tax).
Received: 21 Nov. 2012; revision received: 12 Mar. 2103; accepted: 3 July 2013. DOI: http://dx.doi.org/10.12705/624.11
INTRODUCTION
Sapotaceae are an important component of the rainforests
around the world. Its members are readily identified based on
the presence of white latex, malpighiaceous trichomes, simple
and entire leaves, and flowers in fascicles. Natural groups in
the family have been difficult to circumscribe because of mor
phological homoplasy. Various systems of classification rec
ognizing anything from 53 to 122 genera have been proposed
(Lam, 1939; Aubréville, 1964; Baehni, 1965; Pennington,
1991). Recent phylogenetic analyses using molecular data,
often combined with morphological characters (Anderberg
& Swenson, 2003; Smedmark & al., 2006), have identified
three major lineages, which now are formally recognized as
subfamilies Chrysophylloideae, Sapotoideae, and Sarcosper
matoideae (Swenson & Anderberg, 2005). Chrysophylloideae
are most diverse in South America and Australasia, Sapot
oideae in Africa and Asia, while Sarcospermatoideae are re
stricted to Asia. Many systematic and biogeographic problems
remain unsolved within each subfamily, and this paper focuses
on systematic issues in the Chrysophylloideae of Oceania and
Southeast Asia.
746
All members of Chrysophylloideae in Australasia form a
monophyletic group (Bartish & al., 2005; Swenson & Ander
berg, 2005; Swenson & al., 2008b), and molecular dating sug
gests emergence in the area by the Early Eocene, some 52 mil
lion years ago (Ma), with subsequent radiation, range expansion
and multiple dispersals to New Caledonia (Bartish & al., 2011).
Phylogenetic analyses of the Australasian members (Bartish
& al., 2005; Swenson & al., 2007a; Triono & al., 2007) sug
gest that at least seven lineages can be delineated as genera,
which prompted Swenson & al. (2007a) to recognize Magodendron Vink, Niemeyera F. Muell. and Pichonia Pierre, and
to resurrect the Pouteria Aubl. segregates Beccariella Pierre,
Planchonella Pierre, Sersalisia R. Br. and Van-royena Aubrév.
(Fig. 1). However, the phylogenetic study by Swenson & al.
(2007a) revealed several problems: (i) a large polytomy pre
vented them from conclusive hypotheses about intergeneric
relationships, (ii) Van-royena either grouped with the Australian
species Niemeyera antiloga (F. Muell.) T.D. Penn. or collapsed
into a polytomy, and (iii) generic limits remained unclear within
an Australian–New Caledonian clade, the Niemeyera complex.
Generic limits in the Niemeyera complex have been as
sessed by phylogenetic analyses using nuclear (ETS, ITS) and
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Swenson & al. • Chrysophylloideae in Oceania and SE Asia
Fig. 1. Field images of members of subfamily Chrysophylloideae (Sapotaceae) growing in Oceania. A, Niemeyera chartacea; B, Niemeyera antiloga; C, Van-royena castanosperma; D, Pycnandra belepensis; E, Pichonia deplanchei; F, Sersalisia sericea; G, Planchonella baillonii; H, Planchonella ericiflora; I, Planchonella minutiflora; J, Planchonella tahitensis; K, Planchonella rufocostata; L, Beccariella sp. nov. — Photos: A, C,
F by Hugh Nicholson (Australia); B by Glenn Leiper (Australia); D, I by Ulf Swenson (Sweden); E by JeanLouis Ruiz (New Caledonia); G, H,
K, L by Jérôme Munzinger (France); J by JeanFrançois Butaud (Tahiti).
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Swenson & al. • Chrysophylloideae in Oceania and SE Asia
chloroplast (trnH-psbA, trnS-G) sequence data, as well as mor
phology (Swenson & al., 2008a). The complex included seven
genera in the classification by Aubréville (1964, 1967), a number
that was decreased to three by Pennington (1991). Both authors
recognized Leptostylis Benth., a genus distinguished by oppo
site (rather than alternate) leaves and four sepals (not five or
more). There are two strongly supported clades in the complex,
one confined to Australia and another to New Caledonia, the lat
ter with several subclades but with weak internal node support.
Two morphological characters traditionally used for generic rec
ognition in the Niemeyera complex are the anisomerous flower
and the number of stamens inserted opposite each corolla lobe.
Both characters are highly homoplastic and are not diagnostic
even for small subclades (Swenson & al., 2008a). A generic
framework based on a monophyletic criterion was proposed
that allowed recognition of anywhere from one to five genera,
a framework differing from the concept used by Aubréville or
Pennington. Nonetheless, based on unpublished molecular data
and the discovery of several new species, Swenson & Mun
zinger (2009) found that a narrow generic concept was untenable
and accepted Niemeyera for the species in Australia and Pycnandra Benth. for the lineage confined to New Caledonia. The
New Caledonian subclades were relegated to subgeneric rank.
However, it is still unclear if Leptostylis can be recognized at
generic level or if it is better subsumed in Pycnandra.
Beccariella and Planchonella are usually rainforest trees
or shrubs in maquis vegetation. They are widely distributed in
Oceania and Southeast Asia and have convoluted taxonomic
histories (Swenson & Morat, 2008; Swenson & Tehler, 2009).
As currently circumscribed, Beccariella includes some 20 spe
cies and Planchonella about 60 species (Swenson & al., 2007a,
b). Some outdated revisions are available for Malesia, but at that
time the species were distributed among Planchonella (99 spp.;
Van Royen, 1957), Krausella H.J. Lam (4 spp.; HerrmannErlee
& Lam, 1957), and Pouteria (28 spp.; HerrmannErlee & Van
Royen, 1957). Groups were so poorly delimited that the revision
of Pouteria treated taxa that now are firmly placed in Beccariella, Pichonia, Planchonella, Pycnandra, and Sersalisia (Swen
son & al., 2007a; Swenson & Munzinger, 2010a). Pouteria is
a genus restricted to South America (Swenson & al., 2008b).
There are no unique synapomorphies for either Beccariella or
Planchonella, but they are distinguished based on character
combinations. Common characters of both genera include flow
ers with a corolla with staminodes, styles with stigmatic areas
visible using a hand lens, seeds with foliaceous cotyledons, a
radicle that extends below the cotyledon commissure, and an
endosperm. Species of Beccariella are identified by areolate
leaf venation (tertiaries or quaternaries), stamens inserted in the
middle or near the base of the corolla tube, a fruit with usually
one (or two) seed(s), and a linear and shorter seed scar that often
is 50%–75% of the length of the seed. Species of Planchonella
are recognized by the lack of areolate leaf venation, stamens
that are inserted just below the tube orifice (with a few excep
tions), a fruit having up to five seeds, and a long seed scar that
often is linear and covers 90%–100% of the seed length. How
ever, in contention with this, a search (accessed March 2013) in
the online World Checklist at the Royal Botanical Garden, Kew
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(http://apps.kew.org/wcsp/home.do) for taxa in these genera
(and native to Southeast Asia and Oceania), revealed 17 species
of Beccariella, 107 of Planchonella, and 28 of Pouteria, which
we believe reflects an unnatural classification.
Planchonella is the largest and most widely distributed ge
nus of Chrysophylloideae in Oceania and Southeast Asia. It has
a circumPacific distribution extending from southern China,
Thailand, south to Australia and New Zealand, into the Pacific
to include French Polynesia, Hawaii and back to Taiwan. New
Caledonia is recognized as an important hotspot of biodiversity
(Myers & al., 2000; Lowry & al., 2004) and includes almost
40 species of Planchonella (Munzinger & Swenson, 2009). Spe
cies of this genus exhibit intriguing traits, such as a gynomo
noecious sexual system (Méndez & Munzinger, 2010), as well
as being successful colonizers of the Pacific islands (Bartish
& al., 2005). One New Caledonian endemic species, P. baillonii
(Zahlbr.) Dubard, is the sole member of a lineage that is sister
to the entire genus (Swenson & al., 2007a; Triono & al., 2007).
This sister position was maintained in a phylogenetic analysis
using nine cpDNA and nrDNA loci (Swenson & al., 2008b),
but the study only included seven species of Planchonella. It is
pertinent to address the phylogenetic position of P. baillonii in
a broader context because New Caledonia seems to have been
submerged after rifting from Australia and therefore unavailable
for colonization before 37 Ma (Grandcolas & al., 2008), but see
Heads (2008) for an alternative view.
Apart from the species in New Caledonia, few studies have
addressed problems concerning the Planchonella species oc
curring in the Pacific. Here we will address phylogenetic rela
tionships among taxa found in Hawaii and French Polynesia.
Planchonella sandwicensis (A. Gray) Pierre grows on all main
islands of Hawaii. It is an extremely variable species that has
inspired some scholars to recognize up to six different species
(see Wagner & al., 1990). There are two widely different hy
potheses on its relationships. Swenson & al. (2007a, b) found a
close relationship to taxa in Fiji. In contrast, Triono & al. (2007)
found it as the sister species to subfamily Chrysophylloideae,
i.e., a possible member of Sapotoideae or Sarcospermatoideae.
If the former is correct, the species is most probably derived
from a common ancestor within the Pacific region, which col
onized Hawaii from islands in the west, but if the latter is true,
then the evolutionary lineage ought to be very old and its origin
is more difficult to explain, since the closest relatives are pres
ently found in continental Southeast Asia. However, the gener
ated sequences by Triono & al. (2007) were never submitted to
GenBank (http://www.ncbi.nlm.nih.gov/ last accessed March
2013), making them inaccessible for further testing.
Two Pacific species of Planchonella, P. grayana H. St.
John and P. tahitensis (Nadeaud) Pierre ex Dubard (Fig. 1J), are
at odds with all other species of the genus by having stamens
inserted near the base of the corolla tube, rather than below
the tube orifice as is diagnostic for the genus (Swenson & al.,
2007b). Planchonella grayana occurs from Vanuatu in the west
to Tuamotu (French Polynesia) in the east, while P. tahitensis is restricted to the Society Islands. Distinct morphological
forms have also been described from Tahiti (Fosberg, 1992) or
are possibly novel species, for instance from Raiatea (Society
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Swenson & al. • Chrysophylloideae in Oceania and SE Asia
Islands). The phylogenetic positions of these species are of
interest due to their odd floral morphology, taxonomic status,
and conservation status. The native flora of the Society Islands,
including P. tahitensis (Taputuari & Tchung, 2003; Pouteau
& al., 2012), is under threat from the invasive alien Miconia
calvescens DC. (Melastomataceae; Meyer & Florence, 1996).
The present study aims to estimate phylogenetic relation
ships within Chrysophylloideae in Oceania and Southeast Asia,
excluding the distantly related Asian Xantolis Raf. (Anderberg
& Swenson, 2003; Swenson & Anderberg, 2005), by using
an extended taxon sample and nrDNA sequence data (ETS,
ITS, RPB2), analysed using Bayesian inference and parsimony
jackknifing. Our primary goals are to: (i) resolve the backbone
polytomy; (ii) test the generic concept proposed by Swenson
& al. (2007a); (iii) investigate the relationships of Van-royena
and Niemeyera antiloga; (iv) optimize diagnostic morphologi
cal characters on the phylogeny; (v) solve the specific questions
pertinent to Planchonella in French Polynesia, P. baillonii in
New Caledonia, and P. sandwicensis in Hawaii; and (vi) make
nomenclatural changes as far as possible to obtain monophy
letic genera.
MATERIALS AND METHODS
Nomenclature and taxon sampling. — We follow the sub
family classification of Swenson & Anderberg (2005). The
checklist of Sapotaceae (Govaerts & al., 2001) follows Pen
nington’s (1991) generic classification and includes a full list
of published names, but accepted names are continuously up
dated online at the World Checklist of Selected Plant Families,
Royal Botanical Garden, Kew (http://apps.kew.org/wcsp/home
.do). This online resource follows the amended classification
proposed by Swenson & al. (2007a), which we herein follow
and aim to test.
A total of 168 terminals were selected for this study, in
cluding all type species of the genera recognized to date except
for Leptostylis longiflora Benth. (probably conspecific with
L. filipes Benth.) and Magodendron (Appendix 1) as well as the
generitypes of two Pouteria segregates: Blabeia Baehni (rep
resented by Planchonella endlicheri (Montrouz.) Guillaumin)
and Fontbrunea Pierre (represented by Pouteria malaccensis
(C.B. Clarke) Baehni). We also included two species of Krausella,
one of Beauvisagea Pierre, represented by Pouteria maclayana
(F. Muell.) Baehni (Govaerts & al., 2001), two unplaced species
of Chrysophyllum L., and finally three species of Leptostylis
(Aubréville, 1967). The remaining terminals are names to be
found online at the World Checklist resource (accessed March
2013). A few taxa, however, are either novel or impossible to
determine and are therefore indicated by quotation marks. The
systematic problem of Planchonella sandwicensis is addressed
using three accessions, two from Oahu and one from Kauai. The
problem of French Polynesian Planchonella is approached with
seven accessions collected in Wallis and Futuna (a French terri
tory northeast of Fiji) and from several archipelagos of French
Polynesia (Austral, Society, and Tuamotu Islands). Our sample
includes all accepted species of Beccariella and Niemeyera, 80%
of Pichonia and Sersalisia, 72% of Pycnandra, and 57% of Planchonella. In addition, nine accessions of Pouteria are included
that are predicted to belong to one or the other clade.
When an outgroup is selected it ought to be as closely
related as possible, but not necessarily the sister group to the
ingroup, and it need not necessarily include more than one
taxon (Nixon & Carpenter, 1993). Xantolis and the Australa
sian clade represent Chrysophylloideae in Southeast Asia and
Oceania; the latter has been demonstrated monophyletic in all
recent cladistic analyses using morphology and/or molecular
evidence (Bartish & al., 2005, 2011; Swenson & Anderberg,
2005; Swenson & al., 2007a, 2008b). Xantolis is sister to the
entire subfamily and strongly diverged from the Australasian
taxa, and therefore not an appropriate outgroup. Genera that
form the sister clade to the Australasian taxa are all from South
America, a clade that possibly includes the African genus Aubregrinia Heine. We therefore selected two species of Ecclinusa Mart., a genus suggested to be one of the closest relatives
to the Australasian taxa (Swenson & al., 2008b; Bartish & al.,
2011). If any of the additional taxa included here were not to
be member of this clade, they are expected to fall between the
outgroup and the Australasian clade, not within it.
Molecular data. — Selection of molecular markers to pro
vide a sufficient number of informative characters needs con
sideration. Different cpDNA regions vary in their relative utility
to resolve relationships within angiosperm families (Shaw & al.,
2005, 2007). Broad studies of Sapotaceae have used several cod
ing and noncoding cpDNA and nrDNA regions with varying
success. Of the cpDNA regions used for phylogenetic estimates
(atpB-rbcL, ndhF, petN-psbM, psbM-trnD, psbB-psbH, rpl20rps12, trnC-petN, trnH-psbA, trnL-trnF, and trnS-trnG) the av
erage number of informative substitutions is close to 3%, where
trnS-trnG has 1% (Swenson & al., 2008a) and trnH-psbA has 6%
informative sites (Swenson & al., 2008b). In contrast, the nrDNA
loci ITS and ETS have proven to be much more useful with some
23%–54% informative characters (Bartish & al., 2005; Swenson
& al., 2008b), which is why they were selected for this study.
Plant material for DNA extraction was collected either as sil
ica geldried leaf material or fragments removed from herbarium
specimens. New sequences of ITS1 and ITS2 were obtained from
61 accessions, following the protocol for DNA extraction, ampli
fication, and sequencing described by Bartish & al. (2005), and
added to 106 previously published sequences. All primers are
listed in Table 1. ETS sequences of 122 accessions were obtained
by using two primers, 18SETS (Baldwin & Markos, 1998) and
Sap1 (Swenson & al., 2008a), following the touchdown PCR
protocol: 95°C for 5 min, followed by 4 cycles of 95°C for 30 s,
57°C for 30 s, 72°C for 1 min 15 s, 4 cycles of 95°C for 30 s, 55°C
for 30 s, 72°C for 1 min 15 s, and 32 cycles of 95°C for 30 s, 53°C
for 30 s, 72°C 1 min 15 s followed by 72°C for 8 min. Both ITS
and ETS may occur in multiple copies in a genome, which can
indicate hybrid origin of a taxon (Poczai & Hyvönen, 2010). We
carefully checked the sequences for double peaks in the proof
reading procedure for the presence of multiple copies. Multiple
copies of ETS were detected in eight Planchonella species, all
from a clade confined to New Caledonia, i.e., Clade D3b identi
fied by Swenson & al. (2007b). We decided to exclude these and
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Swenson & al. • Chrysophylloideae in Oceania and SE Asia
reduced the clade to three species (P. glauca, P. lauracea, P. “Ile
Yande”) in order to minimize problems related to concerted
evolution or lineage sorting (Álvarez & Wendel, 2003).
We sought to include an additional nuclear marker and se
lected RPB2, which has proven to be useful for phylogenetic
estimates on high as well as low systematic level (Oxelman
& Bremer, 2000; Oxelman & al., 2004; Eggens & al., 2007). The
RPB2 region for 161 accessions was amplified using the primers
P6F and P7R (Denton & al., 1998), and several herein designed
primers to obtain sequences around polyTregions, followed by
the same PCR protocol described above. Purified products were
sequenced using an ABI3130xl Automated DNA Sequencer
(Applied Biosystems, Foster City, California, U.S.A.).
Alignment, gap coding, and model testing. — Alignments
were performed on each separate sequenced locus in MAFFT
v.6.818b (Katoh & al., 2005) using the LINSi predefined pa
rameter settings. Resulting matrices were imported into MES
QUITE v.2.75 (Maddison & Maddison, 2011) and minor manual
adjustments were made to the alignments. Following the method
of Simmons & Ochoterena (2000), gaps for each locus were
manually scored as binary characters (present/absent) in separate
partitions, disregarding uninformative deletions or inserts. Each
gene partition was tested for the bestfit substitution model using
jModelTest v.0.1.1 (Posada, 2008) under the Bayesian informa
tion criterion (BIC) (Schwartz 1978; Posada & Buckley, 2004)
in order to minimize the number of substitution rate parameters.
Binary data (gaps) were assigned a simple substitution model
allowing unconstrained reversible gains/losses of characters.
Phylogenetic analyses. — Phylogenetic relationships were
estimated with Bayesian inference (Rannala & Yang, 1996;
Yang & Rannala, 1997) and parsimony jackknifing (Farris
& al., 1996). The aligned matrix was prepared in BEAUti v.1.6.2
(part of the BEAST package) as an output file for Bayesian in
ference in BEAST v.1.6.2 (Drummond & Rambaut, 2007). Each
locus was treated as a unique partition. Substitution models
were set by manual modification of the rate parameters. This
study used the BEAST package to primarily derive a tree topol
ogy, not divergence time estimates under a molecular clock
assumption. The molecular clock was therefore unconstrained
and the root was fixed by using a normal prior with an arbitrary
mean (100) and a narrow standard deviation (0.1). The Monte
Carlo Markov chains (MCMC) were set to run five times, each
for 100 million generations, to assure independent convergence
on all parameters (ESS values > 200), sampling trees every
25,000 generations. Convergence and chain mixing were re
viewed in Tracer v.1.5 (Rambaut & Drummond, 2009). A pro
portion of the samples in each run were discarded as burnin,
and the posterior set of trees was summarized in TreeAnnotator
v.1.6.2 (Drummond & Rambaut, 2007). The resulting tree was
then visualized in FigTree v.1.3.1 (Rambaut, 2009).
Jackknife analysis, implemented in PAUP* 4.0 (Swofford,
2002), was also performed on the dataset to retrieve parsimony
support values. The settings were as follows: 1000 jackknife
replicates with a single random addition sequence, TBR branch
swapping, collapsing branches of zero length, steepest descent
not in effect, and saving a maximum of 1000 trees in each
replicate. The excluded fraction of characters in each replicate
was set to 37%.
Posterior probability (PP) and parsimony jackknife values
(JK) below 0.8 and 50%, respectively, are not reported. We con
sider PP values of 0.95 or more to be strong indicators of node
support, whereas JK values of 50%–74% are weak, 75%–89%
moderate, and 90%–100% are considered strong. Nodes that
receive less than JK 50% and below PP 0.8 posterior support
are collapsed in the phylogeny with one exception (the plausible
position of Van-royena).
Morphological data. — Morphology is highly homoplastic
in Sapotaceae and unique generic synapomorphies are absent
or very rare. Swenson & al. (2007a) proposed that character
state combinations ought to be used for generic recognition.
Table 1. List of primers used.
DNA region
Primer
Sequence 5′–3′
ETS
18SETS
ACT TAC ACA TGC ATG GCT TAA TCT
Baldwin & Markos (1998)
ETS
Sap1
CGT ACT TGA GCG TGT TGG TGT
Swenson & al. (2008a)
ITS
18SF
GAA CCT TAT CGT TTA GAG GAA GG
Rydin & al. (2004)
ITS
26RN
CCG CCA GAT TTT CAC GCT GGG C
Rydin & al. (2004)
RPB2
P6F
TGG GGA ATG ATG TGT CCT GC
Denton & al. (1998)
RPB2
P7R
CCC ATG GCT TGC TTC CCC AT
Denton & al. (1998)
RPB2
270FSap
CCT AGT GTT ACC TTT TAC CCT GAT TG
This study*
RPB2
570FSap
CAT GGC ATT CAA CTA CTG AAG AGT TG
This study*
RPB2
540RSap
CAA CTC TTC AGT AGT TGA ATG CCA TG
This study*
RPB2
435FSap
TAA TAT GTC AGC TTG TGG TGG AGA G
This study*
RPB2
435RSap
CTC TCC ACC ACA AGC TGA CAT ATT A
This study*
RPB2
800FSap
GCA TCT CTG TCC TCT TGA TTA CTT GAA TG
This study*
RPB2
800RSap
CAT TCA AGT AAT CAA GAG GAC AGA GAT GC
This study*
RPB2
1020FSap
GCT ACT TTG AAG CAC TTG CTA GAA TCT
This study*
RPB2
1020RSap
AGA TTC TAG CAA GTG CTT CAA AGT AGC
This study*
*Primers designed by Bodil Cronholm, Swedish Museum of Natural History.
750
Reference
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Swenson & al. • Chrysophylloideae in Oceania and SE Asia
The morphological data used here have been gathered from
earlier studies (Swenson & Anderberg, 2005; Swenson & al.,
2007a, b, 2008a, b), revisions (Van Royen, 1957; HerrmannEr
lee & Lam, 1957; HerrmannErlee & Van Royen, 1957) and her
barium material deposited at L, MO, NOU, P, PAP, and S. The
morphological terminology follows Harris & Harris (1997).
We used MacClade v.4.0 (Maddison & Maddison, 2000) to
optimize the characters on the maximum clade credibility tree
(MCC) obtained from the BEAST analysis. We then condensed
the phylogeny, opting for genera to be in proportion to the
number of the species, and traced eight characters of which
seven have been identified as useful for diagnostic purposes
(Swenson & al., 2007a). One character, pubescence on sepals,
is used here for the first time. The condensed tree is designed
to comprehensibly visualize the characters.
Character 1. – Tertiary leaf venation has been used for
diagnostic purposes with variable success (Van Royen, 1957;
Pennington, 1991; Triono & al., 2007). A fine areolate vena
tion, formed when higher orders of veins anastomose and form
a closed reticulate pattern, easily observed with a hand lens
(Swenson & Munzinger, 2009), is diagnostic for Beccariella,
Pichonia, and Sersalisia (Swenson & al., 2007a).
Character 2. – Malpighiaceous trichomes are usually
present as an indument on sepals on inner, outer, or both sur
faces, but can also be secondarily lost. Revisional work has
demonstrated that the distribution of trichomes can indicate
relationships. For example, sepals of Planchonella and Pycnandra are usually pubescent outside but glabrous inside (Swenson
& al., 2007b; Swenson & Munzinger, 2009), while members of
Pichonia usually have an indument on both sides of the sepals
(Swenson & Munzinger, 2012). Thus, presence or absence of
trichomes on the inner surface may be congruent with clades.
Character 3. – The corolla of Sapotaceae is actinomor
phic and partly sympetalous, comprising a tube and free co
rolla lobes. Depending on the ratio between the length of the
corolla tube and the lobes, and the corolla length versus the
calyx length, the corolla can be cupshaped, urnshaped, nar
rowly campanulate, or tubular. This character is in fact a com
bination of different features and could be reductively coded
for phylogenetic analyses, but it is also homoplastic (Swenson
& Anderberg, 2005; Swenson & al., 2007a, 2008b). There is no
doubt that flower types are frequently difficult to characterize,
but we wanted to investigate their potential diagnostic value as a
character. The calyx of a cupshaped flower is usually less than
50% of the length of the corolla, the corolla tube is shorter than
the lobes, and the lobes are spreading (Fig. 1A, D). The corolla
of an urnshaped flower usually has a tube and corolla lobes
of equal length (or slightly shorter lobes), and the calyx often
extends above the corolla tube orifice (Fig. 1I, J). A campanulate
flower is similar to an urnshaped and/or a tubular flower, but
the corolla exerts slightly above the sepals and forms a small bell
(Fig. 1L). Tubular flowers have a visible corolla tube, longer than
the sepals and clearly longer than the corolla lobes (Fig. 1F–H).
Character 4. – Stamens in Sapotaceae are inserted at dif
ferent levels in the corolla tube. This character was important
for Aubréville (1964) and less so for Pennington (1991), but has
been found to be diagnostic for monophyletic groups (Swenson
& al., 2007a). Hence, stamens are inserted in the tube orifice
(Niemeyera, Pichonia, Pycnandra), just below the tube orifice
(Planchonella), in the middle of the tube or near the corolla
base (Beccariella).
Character 5. – One entire style is always present in Sapot
aceae, which may, however, differ at the apex. It has either vis
ible, small, round stigmatic areas readily identified with a hand
lens (Fig. 1F), a character suggested diagnostic for Beccariella, Planchonella, Sersalisia, and Van-royena (Swenson & al.,
2007a). In contrast, if no stigmatic areas are visible (Fig. 1A),
the style apex is termed simple as in Niemeyera, Pichonia, and
Pycnandra (Pennington, 1991; Swenson & Anderberg, 2005).
Characters 6–8. – A number of fruit characters show strong
congruence and can be amalgamated to three features: (i) cotyle
dons foliaceous, having a radicle extending below the cotyledon
commissure, endosperm present; (ii) planoconvex cotyledons
without a visible radicle, endosperm absent; and (iii) planocon
vex cotyledons with an exserted radicle, endosperm absent.
RESULTS
Data. — The complete matrix contains 2563 characters,
of which 2486 are from aligned nuclear sequences and 77 are
from three partitions of coded gaps (Table 2). ETS is repre
sented by 403 nucleotides of which 198 (49.1%) are parsimony
informative, ITS of 914 nucleotides of which 320 (35%) are
parsimony informative, and RPB2 of 1169 nucleotides of which
151 (12.9%) are parsimony informative.
The model test resulted in the selection of TrN + Γ for both
ETS and ITS, and TPM1 + Γ for RPB2. Initial analyses revealed
nested parameter ranges for ITS and ETS, and the two loci were
therefore set to share the substitution model in order to decrease
the total number of parameters.
Tree topology. — Bayesian analysis and parsimony
jackknifing of the molecular dataset recovered similar tree
topologies. Taxa not formerly analysed in any phylogenetic
Table 2. Characteristics of nuclear sequences in each of the data partitions (excluding the outgroup)
Number of characters
Data
Aligned
Constant
Uninformative
Informative
Gaps
ETS
403
114
91
198 (49.1%)
22
914
418
176
320 (35.0%)
42
RPB2
ITS
1169
483
535
151 (12.9%)
13
Total
2486
1015
802
668 (26.9%)
77
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751
TAXON 62 (4) • August 2013: 746–770
Swenson & al. • Chrysophylloideae in Oceania and SE Asia
context are all recovered with maximum support within the
previously identified Australasian clade. After collapsing nodes
with support below the defined thresholds (0.8 PP, 50% JK),
the tree topologies are close to identical (Fig. 2). Polytomies
are mainly restricted to terminal positions in each genus. One
deep node receives only moderate JK support (83%) and no
Bayesian support. Similar to previous results, the placements of
the Australian Van-royena and the New Guinean Magodendron
are still uncertain, and no other taxon is identified as closely
related to these genera which, in addition, have rather unusual
diagnostic character combinations (Swenson & Anderberg,
2005; Swenson & al., 2007a).
All species included in our analyses were dispersed across
six distinct clades with strong Bayesian support and frequently
with moderate or strong jackknife support. Each of these clades
include one generic type (from top to bottom in Fig. 2): Beccariella sebertii (Pancher) Pierre, Sersalisia sericea (Aiton) R. Br.,
Pichonia balansana Pierre, Niemeyera prunifera (F. Muell.)
F. Muell., Pycnandra benthamii Baill., and Planchonella obovata (R. Br.) Pierre. Embedded in these six clades are species
of Chrysophyllum, Krausella, Leptostylis, Pouteria, and Planchonella (in Beccariella) and the genera can only be rendered
monophyletic following taxonomic transfers. However, Niemeyera is not monophyletic in its present circumscription since
N. antiloga is rendered sister to the two clades corresponding to
Niemeyera and Pycnandra. Both genera are strongly supported,
but subclades relegated to subgeneric rank in the latter genus
(Swenson & Munzinger, 2009) are all recovered in a polytomy.
All species of Leptostylis and the unplaced Chrysophyllum
wagapense Guillaumin form a clade that is sister, with weak
support, to Pycnandra subg. Sebertia (Pierre ex Engl.) Swenson
& Munzinger (Swenson & Munzinger, 2010a).
Planchonella is largely monophyletic with the exception
of four species that are recovered in Beccariella with strong
support. Again, P. baillonii is sister to the entire genus. The
unplaced Chrysophyllum bakhuizenii P. Royen, the Pouteria
segregate Krausella, and some species currently placed in Pouteria, are all embedded in Planchonella. Overall, the clades
identified by Swenson & al. (2007b; Clades D1, D2, and D3),
are all strongly supported in this phylogeny and mutual clade
relationships find good support. Two widely distributed species
in Southeast Asia, Pouteria maclayana and P. malaccensis, are
sisters to Clades D1 and D2. All three accessions of P. sandwicensis from Hawaii group together within Planchonella, a
group with close affinity to taxa from Fiji. All accessions of
P. grayana and P. tahitensis, from Alofi and Futuna in the west
to Tuamotu in the east, are found monophyletic with maximum
support, but mutual molecular differences are small and acces
sions are not reconciled to species or distribution. One clade
circumscribes three accessions of P. linggensis (Burck) Pierre,
one of P. chartacea (F. Muell. ex Benth.) H.J. Lam (Australia),
and one of P. solida P. Royen (New Guinea), in which P. linggensis is polyphyletic.
Optimization of morphology. — Figure 3 shows the eight
morphological characters optimized on the majorityrule con
sensus tree obtained from the BEAST analysis of nuclear se
quences of Chrysophylloideae in Oceania and Southeast Asia.
752
They show all high congruence with the generic concept pro
posed by Swenson & al. (2007a), but some characters still show
a degree of homoplasy. This will be discussed below.
DISCUSSION
Overall resolution. — Our phylogenetic analyses based on
nuclear sequence data yielded improved resolution and a better
understanding of phylogenetic relationships within Chryso
phylloideae (Bartish & al., 2005; Swenson & al., 2007a, b,
2008a; Triono & al., 2007). Exact affinities of Magodendron
and Van-royena remain unclear and both genera, in principle,
fall back to an unresolved polytomy. Nevertheless, our current
analyses find no close relationship between Van-royena and
Niemeyera antiloga, since the latter is strongly supported as
the sister to Pycnandra and a narrowly defined Niemeyera
(Fig. 2). The proposal that Chrysophyllum and Pouteria are not
present in Australasia (Swenson & al., 2007a, 2008b; Swenson
& Munzinger, 2009) finds strong phylogenetic support, because
the included taxa are deeply embedded in different Austral
asian groups. Instead, for this region, the genera Beccariella,
Magodendron, Niemeyera, Pichonia, Planchonella, Pycnandra, Sersalisia, and Van-royena are reconfirmed, and no other
important lineage of the subfamily is identified. Hence, mem
bers currently placed in Chrysophyllum or Pouteria need to
be transferred. Leptostylis needs to be relegated to subgeneric
level within Pycnandra, and a revision is under preparation.
However, two systematic problems remain to be solved: the
polyphyly of Niemeyera and the unresolved nomenclatural
problem concerning the name Beccariella (Swenson & Tehler,
2009; Brummitt, 2011b).
The overall wellresolved phylogeny brings to light in
teresting biogeographic perspectives, such as whether New
Caledonia hosts a relict biota or was submerged about 37 Ma
and later emergent (Grandcolas & al., 2008); and whether the
Fijian flora is derived from immigrants from Asia, Australia,
and/or New Caledonia (Keppel & al., 2009) or evolved in situ
(Heads, 2006). A detailed biogeographic study of this Austral
asianPacific group will be addressed elsewhere.
Useful morphological characters. — Morphology has re
peatedly been demonstrated to be homoplastic in Sapotaceae.
Examples of traditionally used characters believed useful but
shown to be homoplastic include the number of petals, the num
ber of stamens opposite each corolla lobe, and the ratio between
the length of the corolla tube and the corolla lobes (Swenson
& al., 2007a, 2008a, b). Presence of staminodes in Sapotaceae is
a plesiomorphic feature, which has been reduced several times
in Chrysophylloideae (Swenson & Anderberg, 2005; Swenson
& al., 2008b) and Sapotoideae (Smedmark & al., 2006), but can
be diagnostic for less inclusive groups. Staminodes are pres
ent in most Chrysophylloideae in the studied group, but have
been reduced once, in the clade Niemeyera-Pycnandra. Hence,
absence of staminodes is diagnostic for this clade. Another
useful character to distinguish between genera is where the
flowers are born. All genera, except Magodendron and Pycnandra, generally have axillary flowers, whereas the former is
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TAXON 62 (4) • August 2013: 746–770
Swenson & al. • Chrysophylloideae in Oceania and SE Asia
cauliflorous and the latter is usually ramiflorous. Apart from
morphology, geographic distribution is a very strong indicator
of clade affinity, except in the widespread genera Beccariella,
Planchonella, and Sersalisia. In the discussion below of the
eight morphological characters used here, taxon names are
ordered as they appear in Fig. 3 (not alphabetically).
Areolate venation is present in all species of Beccariella,
Sersalisia, Pichonia, and Magodendron, including all species
of Planchonella and Pouteria that are recovered in these clades
(Fig. 3A). In contrast, members of Van-royena, Niemeyera,
Pycnandra and Planchonella always lack areolate venation,
and instead have visible tertiary leaf venation, in agreement
with the findings of Triono & al. (2007).
The sepals of most Chrysophylloideae taxa are pubescent
on the outer surface, except when the indument is secondarily
lost, but presence or absence of indument on the inner surface
seems to contain strong phylogenetic information (Fig. 3B).
In Beccariella and Van-royena, trichomes are always present
on the sepals’ inner surface, often covering the entire area and
forming a tomentose or even woolly indument. Sepal indument
is homoplastic in Sersalisia and Pichonia, and the sepals of
P. balansana, P. deplanchei (Baill.) Swenson & Munzinger,
and P. lecomtei (Guillaumin) T.D. Penn. are glabrous inside.
However, if an indument is present on the inner surface, the
trichomes are usually concentrated in the upper part of the se
pals (Swenson & Munzinger, 2012). Sepals of all other genera
are glabrous inside, except in some species scattered in the
Planchonella phylogeny (P. australis (R. Br.) Pierre, P. chartacea, and P. cyclopensis P. Royen), in which the density of
indument varies, often even between sepals in the same flower.
Classification of floral types is partly subjective and some
times difficult according to the definition above. Nevertheless,
the corolla can be termed narrowly campanulate in Beccariella,
Van-royena, and some Sersalisia (Fig. 3C). A cupshaped co
rolla with spreading lobes is characteristic for Pichonia, Magodendron, Niemeyera, and Pycnandra, and in the two latter the
corolla lobes are frequently revolute (Fig. 1A, D). The type
of corolla in Planchonella is generally urnshaped, where the
calyx is more or less as long as, or slightly longer than, the
corolla tube (Fig. 1I, J). Urnshaped flowers never have spread
ing or revolute corolla lobes comparable to the cupshaped
flower. Exceptions to the urnshaped flower in Planchonella
are found in P. baillonii, P. ericiflora Munzinger & Swenson
and P. myrsinoides (Benth.) S.T. Blake ex Francis, species that
are scattered across the genus and have tubular flowers with a
wellexposed corolla tube (Fig. 1G, H).
Stamens are inserted opposite the corolla lobes in Sapot
aceae, and it has been controversial whether the insertion point
within the corolla tube carries any phylogenetic information.
Both Aubréville (1967) and Van Royen (1957) used the character,
as opposed to Pennington (1991) who could not find significant
correlation between the insertion point of stamens and other
characters, perhaps because his circumscription of Pouteria
was an unnatural amalgamation of different lineages, which
obscured character correlations. However, the phylogenetic
study by Swenson & al. (2007a) found strong correlation between
stamen insertion and monophyletic groups. Our analysis agrees
with these previous findings, but, again, there are exceptions
(Fig. 3D). All species of Beccariella (and Van-royena) have sta
mens inserted either in the middle or near the base of the tube.
Swenson & al. (2007a) reported that the species from Australia
have stamens inserted near the base whereas those from New
Caledonia have the stamens inserted in the middle of the corolla
tube. In fact, this statement needs amendment, since there are
species of Beccariella in Australia, New Caledonia, and New
Guinea that have stamens inserted either near the base or in the
middle of the corolla tube. However, it is still valid that no species
of Beccariella have stamens inserted in or just below the tube or
ifice, character states that are restricted to the other genera under
study. Stamen insertion in Van-royena has been reported as basal
(HerrmannErlee & Lam, 1957; Aubréville, 1963; Swenson & al.,
2007a), but the point of insertion is at or just above the middle of
the corolla tube, while the filament is attached to the tube and
run down to the base. In Sersalisia and Pycnandra the character
varies, and exceptions in Pycnandra are P. atrofusca Swenson
& Munzinger, P. benthamii, P. carinocostata Vink, P. fastuosa
(Baill.) Vink and P. kaalaensis Aubrév., species that represent
three subgenera and have stamens inserted just below the tube
orifice. Leptostylis filipes, which will be transferred to Pycnandra, has tubular flowers and stamens inserted in the middle of
the corolla tube. Pichonia, Magodendron, and Niemeyera have
stamens inserted in the tube orifice. Finally, all except for some
species of Planchonella have stamens inserted just below the
tube orifice. Stamens in P. aneityensis (Guillaumin) H.J. Lam
ex P. Royen are inserted in the middle of the corolla tube, and
in P. grayana and P. tahitensis the stamens are inserted at the
corolla base, a unique position within the genus. These three taxa
are all found in the same clade of Pacific species.
Styles with several stigmatic areas at the apex, easy to iden
tify with a hand lens, was suggested as an important character to
differentiate Beccariella, Van-royena, Sersalisia, Magodendron,
and Planchonella from the simple style without such clearly
visible stigmatic areas in Pichonia, Niemeyera, and Pycnandra
(Swenson & al., 2007a). We found no deviation from this pattern
in the sampled species and suggest that this character does con
tain strong phylogenetic signal, useful in character combination
for circumscribing groups, as well as in the field (Fig. 3E).
The distribution of foliaceous or planoconvex cotyledons,
absence or presence of an exserted radicle, and absence or
presence of an endosperm, are clearly correlated among mem
bers of the study group, and are reported in a single figure
(Fig. 3F). Three combinations of the characters are known: (i)
seeds of Beccariella and Planchonella have foliaceous coty
ledons, an exserted radicle and an endosperm, (ii) Magodendron has planoconvex cotyledons, an exserted radicle and
nonendospermous seeds, and (iii) all other genera have seeds
with planoconvex cotyledons, an included radicle and no en
dosperm. The character combination for Magodendron is rare
in Chrysophylloideae, only found in Chromolucuma Ducke
and Pradosia Liais (Swenson & Anderberg, 2005), and in some
species of Pouteria (South America) and Synsepalum (A. DC.)
Daniell (Africa) sensu Pennington (1991).
In summary, despite a high degree of morphological homo
plasy in Chrysophylloideae, the characters here overlaid in
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753
1
97
1
99
1
100
1
100
0.96
1
59
.97
64
1
1
85
1
91
0.96
64
1
100
1
1
62
1
100
1
98
0.72
1
100
1
100
0.95
61
0.95
0.9
1
100
1
84
1
52
1
100
83
1
84
0.92
73
1
100
1
100
0.8
1
86
1
98
1
100
1
100
1
85
1
76
0.9
72
1
95
0.88
61
1
100
0.96
64
1
100
0.99
1
87
1
0.97
100
73
1
0.97
100
0.86
1
100
1
100
0.96
1
51
100
0.93
61
1
78
0.98
60
1
76
1
100
1
87
1
100
1
100
1
97
0.8
0.92
59
1
92
1
84
1
64
1
88
0.98
85
1
67
1
100
1
95
D2
63
1
100
1
76
0.97
52
1
66
1
100
1
100
1
90
1
86
1
1
1
99
D1
0.99
64 0.86
64
1
74
1
100
1
99
1
100
0.85
1
63
1
100
Current classification
Beccariella
Chrysophyllum
Krausella
Leptostylis
Niemeyera
Magodendron
Pichonia
Planchonella
Pycnandra
Pouteria
Sersalisia
Van-royena
0.82
1
88
1
95
D3
0.8
1
90
1
100
1
100
1
100
1
.92
1
73
1
100
1
68
1
100
1
99
0.8
1
97
1
0.81
1
100
1
100
.87
1
96
0.95
0.8
58
1
100
1
100
1
100
0.96
67
1
100
1
99
1
1 100
100
Planchonella moluccana
Beccariella macrocarpa
Beccariella papyracea
Beccariella singuliflora
Beccariella xerocarpa
Beccariella brownlessiana
Beccariella queenslandica
Beccariella “acutifolia”
Beccariella longipetiolata M2264
Beccariella longipetiolata M2911
Beccariella baueri
Beccariella rubicunda
Beccariella Munzinger2581
Planchonella firma
Pouteria wandae
Pouteria gillisonii
Planchonella lamprophylla
Planchonella ledermannii
Pouteria richardii
Beccariella Takeuchi25691
Beccariella sebertii
Beccariella vieillardii
Beccariella novocaledonica
Beccariella lucens
Beccariella lasiantha
Beccariella crebrifolia
Beccariella “belepensis”
Beccariella brevipedicellata
Beccariella balansana
Van-royena castanosperma
Sersalisia sessiliflora
Sersalisia sericea
Pouteria luzoniensis
Pichonia occidentalis
Pichonia deplanchei
Pichonia lecomtei
Pichonia daenikeri
Pichonia balansana
Pichonia grandiflora
Pichonia balansae
Pichonia dubia S918
Pichonia dubia M3065
Magodendron mennyae
Niemeyera antiloga
Niemeyera whitei
Niemeyera Ford2429
Niemeyera prunifera
Niemeyera chartacea
Pycnandra canaliculata
Pycnandra acuminata
Chrysophyllum wagapense
Leptostylis filipes
Leptostylis grandifolia
Leptostylis goroensis
Pycnandra ouaiemensis
Pycnandra comptonii
Pycnandra blaffartii
Pycnandra controversa
Pycnandra vieillardii
Pycnandra griseosepala
Pycnandra neocaledonica
Pycnandra kaalaensis
Pycnandra decandra
Pycnandra belepensis
Pycnandra bracteolata
Pycnandra glabella
Pycnandra sessilifolia
Pycnandra francii
Pycnandra schmidii
Pycnandra pubiflora
Pycnandra caeruleilatex
Pycnandra sessiliflora
Pycnandra sarlinii
Pycnandra deplanchei
Pycnandra fastuosa
Pycnandra carinocostata
Pycnandra viridiflora
Pycnandra blanchonii
Pycnandra gordoniifolia
Pycnandra paucinervia
Pycnandra atrofusca
Pycnandra benthamii
Pycnandra glaberrima
Pycnandra balansae
Pycnandra cylindricarpa
Pycnandra linearifolia
Planchonella baillonii
Pouteria malaccensis
Pouteria maclayana
Planchonella linggensis Vanuatu
Planchonella chartacea Australia
Planchonella linggensis Futuna
Planchonella solida New Guinea
Planchonella linggensis New Guinea
Planchonella xylocarpa
Planchonella xylocarpa
Pouteria stellibacca
Planchonella maingayi
Planchonella sphaerocarpa
Planchonella arnhemica
Planchonella pohlmaniana
Planchonella asterocarpon
Planchonella membranacea
Planchonella dothioensis
Planchonella thiensis
Planchonella laetervirens
Planchonella mandjeliana
Planchonella cauliflora
Planchonella luteocostata
Planchonella amieuana
Planchonella endlicheri
Planchonella ericiflora
Planchonella Pillon150
Planchonella kaalaensis
Planchonella koumaciensis
Planchonella Munzinger6150
Planchonella minutiflora
Planchonella pronyensis
Planchonella rufocostata
Planchonella saligna
Planchonella crassinervia
Planchonella roseoloba
Planchonella skottsbergii
Planchonella kuebiniensis
Planchonella howeana
Planchonella myrsinifolia
Planchonella cotinifolia
Planchonella eerwah
Planchonella costata
Planchonella australis
Planchonella myrsinoides
Planchonella euphlebia
Planchonella Ile Yande
Planchonella lauracea
Planchonella glauca
Chrysophyllum bakhuizenii
Planchonella clemensii
Planchonella obovata
Planchonella mindanaensis
Planchonella Munzinger6514 Fiji
Planchonella vitiensis Fiji
Planchonella aneityensis Vanuatu
Planchonella tahitensis Raiatea
Planchonella grayana Tuamotu
Planchonella tahitensis Tahiti
Planchonella grayana Alofi
Planchonella grayana Futuna
Planchonella grayana Australs
P. grayana var. florencei Tahiti
Pouteria pullenii
Pouteria Armstrong316
Planchonella Munzinger6490
Planchonella smithii
Planchonella umbonata
Planchonella sandwicensis K119
Planchonella sandwicensis Okamoto
Planchonella sandwicensis M3227
Planchonella cyclopensis P1365
Planchonella anteridifera
Planchonella thyrsoidea
Planchonella cyclopensis T25495
Planchonella dulcitan
Planchonella torricellensis Futuna
Planchonella torricellensis Alofi
Krausella patentinervia
Krausella polyneura
B Sepals inner surface
Van-royena
Sersalisia
Van-royena
Sersalisia
Pichonia
Pichonia
Magodendron
Niemeyera antiloga
Magodendron
Niemeyera antiloga
Niemeyera
Niemeyera
Pycnandra
Pycnandra
Planchonella
Planchonella
not areolate
areolate
D Stamens inserted
Van-royena
Sersalisia
Van-royena
Sersalisia
Pichonia
Pichonia
Magodendron
Niemeyera antiloga
Magodendron
Niemeyera antiloga
Niemeyera
Niemeyera
Pycnandra
Pycnandra
Planchonella
Planchonella
cup-shaped
urn-shaped
narrowly campanulate
tubular
F Seed characters
Van-royena
Sersalisia
in tube orifice
just below tube orifice
in middle of the tube
near the base
Beccariella
Beccariella
E Style
pubescent
glabrous
Beccariella
Beccariella
C Flowers
Beccariella
Beccariella
A Higher leaf venation
Van-royena
Sersalisia
Pichonia
Pichonia
Magodendron
Niemeyera antiloga
Magodendron
Niemeyera antiloga
Niemeyera
Niemeyera
Pycnandra
Pycnandra
Planchonella
Planchonella
with stigmatic areas
simple
cotyledons foliaceous,
radicle exserted and
endosperm present
cotyledons plano-convex,
radicle included and
endosperm absent
cotyledons plano-convex,
radicle exserted and
endosperm absent
Fig. 3. Eight morphological characters of diagnostic value mapped on a condensed generic phylogeny of Chrysophylloideae (Sapotaceae) present
in Oceania and Southeast Asia. Three seed characters are mapped in F. Black lines represent an equivocal state. Embedded triangles or lines in
different colours represent taxa with homoplasious characters (see text).
◄ Fig. 2. Maximum clade credibility tree obtained from the BEAST analysis of nuclear sequences of Chrysophylloideae (Sapotaceae) from Ocea
nia and Southeast Asia. Currently accepted names are colourcoded according to the legend. Posterior probabilities (PP; above) and parsimony
jackknifing values (JK; below) are found along the branches. Nodes with support below PP 0.8 and JK 50 are collapsed, except for the estimated
relationship of Van-royena. Three subclades (D1–D3) within Planchonella, identified by Swenson & al. (2007b), are marked with arrows. The
outgroup Ecclinusa is pruned from the figure.
TAXON 62 (4) • August 2013: 746–770
Swenson & al. • Chrysophylloideae in Oceania and SE Asia
the condensed phylogeny show high or full congruence with
the identified groups. We believe that we have now arrived at
a stage where unique character combinations can be used to
identify natural groups within Chrysophylloideae present in
Oceania and Southeast Asia.
Resurrection of Amorphospermum. — Niemeyera and
Amorphospermum F. Muell. were originally described with
one species each, N. prunifera and A. antilogum F. Muell.
(Mueller, 1870), hence, these species are the two generitypes.
The genera were described as close relatives, Niemeyera being
distinguished by its berrylike fruit and thin seed coat (testa) in
contrast to Amorphospermum with its drupelike fruit and thick
seed coat. Both genera were later united with Chrysophyllum
(Vink, 1958), but Aubréville (1964) resurrected them in his
system of classification, accepting two species in each genus.
By contrast, Pennington (1991) accepted Niemeyera but not
Amorphospermum, and considered the seed coat differences
to be merely of specific importance. In fact, Niemeyera sensu
Pennington was an unnatural assemblage of Australian species
(now in Niemeyera) and seven taxa from New Caledonia (now
Pycnandra; Swenson & Munzinger, 2009, 2010a, c).
Niemeyera antiloga, a species confined to the rainforest of
Queensland, was unsatisfactorily accommodated in Niemeyera,
because phylogenetic analyses of nuclear sequences grouped
it with Van-royena, in conflict with combined morphological
and molecular data, which grouped it with other species of Niemeyera (Swenson & al., 2007a). The current phylogenetic study
lends strong support to N. antiloga being the sister species of
Niemeyera plus Pycnandra. Our study demonstrates that the cir
cle is now closed and we are back to the generic notion Mueller
(1870) suggested over 140 years ago. Therefore, we propose
that Amorphospermum is resurrected, presently as a monotypic
genus. Apart from seed coat differences, Amorphospermum is
readily distinguished based on leaf characters. The texture is
never translucent, the upper surface is glossy green while the
lower is paler with a persistent indument, and the tertiary ve
nation is parallel (Fig. 1B). Species of Niemeyera have leaves of
similar colour on both surfaces, are usually glabrescent below,
and have horizontal or oblique tertiary leaf venation (Fig. 1A).
Other floral and fruiting characters of Amorphospermum and
Niemeyera share many similarities.
Resurrection of Pleioluma. — Beccariella is strongly sup
ported as an independent lineage with maximum Bayesian and
jackknife support. This relationship is fully congruent with ear
lier findings (Bartish & al., 2005; Swenson & al., 2007a), and
Clade C of Triono & al. (2007). The position within the Australa
sian Chrysophylloideae is unambiguous but its sister relationship
receives only moderate jackknife support. The clade, however, is
readily identified with a character combination of (i) leaves with
areolate venation, (ii) sepals that are pubescent on both surfaces,
(iii) presence of staminodes, (iv) stamens inserted in the middle
of the corolla tube or near the base, (v) style with stigmatic areas,
and (vi) seeds with foliaceous cotyledons, exserted radicle, and
endosperm (Fig. 3). The genus currently contains about twenty
species and another seven species (Planchonella firma (Miq.)
Dubard, P. lamprophylla (K. Krause) H.J. Lam, P. ledermannii
(K. Krause) H.J. Lam, P. moluccana (Burck) H.J. Lam, Pouteria
756
gillsonii Vink, P. richardii (F. Muell.) Baehni, P. wandae Vink)
are here recovered in this clade and need to be transferred to it,
but the name Beccariella is a later homonym and cannot be used
(Swenson & Tehler, 2009; Brummitt, 2011b).
When Pierre (1890) described Beccariella in Sapotaceae
he was probably unaware of the fact that the name was already
occupied for a fungal genus from Borneo. Hence, Beccariella
Pierre became a later homonym and illegitimate already at pub
lication. Subsequent classification systems like HerrmannErlee
& Lam (1957), Van Royen (1957), Baehni (1965), and Pennington
(1991) have not used Beccariella or identified the assemblage as
a natural group. In contrast, Aubréville (1962) resurrected the
genus and designated B. sebertii (included in this study) as the
generic type, and since then the name has been used in New
Caledonia (Aubréville, 1964, 1967), but not in Australia or New
Guinea. Swenson & Tehler (2009) addressed this nomenclatural
problem and suggested conserving Beccariella Pierre (Sapota
ceae) against the rarely used name Beccariella Ces. (Podoscy
phaceae), a proposal the Nomenclatural Committee for Vascular
Plants was unable to agree upon (Brummitt, 2011b). Thus, spe
cies belonging to Beccariella (or that clade) are currently left in
limbo and need to be accommodated under a valid name.
Several of the proposed Sapotaceae genera (Albertisiella
Pierre ex Aubrév., Bureavella Pierre, Iteiluma Baill., Peuceluma
Baill., Pyriluma Aubrév.) in Australasia are polyphyletic or
embedded in Planchonella (Swenson & al., 2007a, b). Some
generic names are still in synonymy with Pouteria (Penning
ton, 1991; Govaerts & al., 2001) and could be used instead of
Beccariella if the generic type is demonstrated to be a member
of this clade. However, three generitypes are here recovered in
Planchonella and cannot be used: Beauvisagea that is united
with Pouteria maclayana (Govaerts & al., 2001), Fontbrunea
represented by Pouteria malaccensis, and Blabeia represented
by Planchonella endlicheri (Fig. 2). Three generic names re
main to be explored: Krausella, Pleioluma, and Wokoia Baehni.
The genus Krausella was described by Lam (1932) and
believed to include four to six species from New Guinea, but
much of the studied material was incomplete or even sterile
(HerrmannErlee & Lam, 1957). The generic type K. polyneura
(K. Krause) H.J. Lam is currently classified as Pouteria multinervis T.D. Penn. (see Pennington, 1991: 202). After receiving
permission from Kew Gardens, we removed a leaf fragment
from the type collection of K. polyneura and amplified most
of ITS and ETS (unsuccessful with RPB2), as well as all three
markers for K. patentinervia (K. Krause) Erlee. These two
species are embedded within Planchonella, sister to P. torricellensis (K. Schum.) H.J. Lam (Fig. 2). In fact, the sequences
of K. polyneura and K. patentinervia are identical. We are
confident that both species are members of Planchonella and
not Beccariella (or any other genus) since, in addition, the leaf
venation is not areolate, the sepals are glabrous inside, stami
nodes are present, stamens are inserted below the tube orifice,
and the styles possess stigmatic areas. Thus, Krausella is here
united with Planchonella.
Pouteria rhopalocarpa P. Royen was described from New
Guinea, but its status as a new genus was in doubt because
of a 3merous flower, staminodes, and a peculiar fruit (Van
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Swenson & al. • Chrysophylloideae in Oceania and SE Asia
Royen, 1959). In any event, Baehni (1964) described Wokoia
and suggested a close relationship to the Malagasy genus
Tsebona Capuron, a member of tribe Omphalocarpeae sensu
Pennington (1991), which is polyphyletic and subsumed in
Chrysophylloideae (Swenson & Anderberg, 2005). However,
from the morphological description (Van Royen, 1959; Vink,
2002) and the generic framework here proposed, this species
is clearly a member of Pichonia. The leaf venation is areolate,
sepals are glabrous on the inside at the base but tomentose on
the upper part, flowers have staminodes, anthers are inserted
in the tube orifice, styles are simple without clear stigmatic
areas, and the seeds have planoconvex cotyledons, an included
radicle and lack endosperm. We therefore unite Wokoia with
Pichonia, which has nomenclatural priority.
The remaining option to accommodate the species of Beccariella is the genus Pleioluma (Baehni, 1965). Pleioluma, first
described as a section of Sersalisia (Baillon, 1891b) and later
transferred to a section of Sideroxylon L. (Engler, 1897), was
based on Sideroxylon crebrifolium (Baill.) Engl. and has never
been used in any classification. In fact, Baehni (1965) simultane
ously transferred this species to Pouteria (p. 59) and in the same
publication (p. 150) accepted it at generic level as Pleioluma
with uncertain relationship. Regardless of this confusion (his
publication appeared after he had deceased), we demonstrate
that Sideroxylon (Beccariella) crebrifolium is deeply nested in
the clade called Beccariella and we suggest, in accordance with
Article 11.3 of McNeill & al. (2012), that Pleioluma is used for
this genus since it is the earliest legitimate name available. In
deed, it is an appropriate name since “pleio” is of Greek origin
and means “more”, whereas “luma” (of unclear origin) means
“cavity”, and could likely refer to the areolate leaf venation.
Problems in Planchonella. — Planchonella is strongly
supported as a monophyletic group. The genus can be charac
terized by nonareolate leaf venation, sepals that are generally
tomentulose on the outer surface but glabrous inside, flowers
with staminodes and stamens inserted just below the tube ori
fice, a style with stigmatic areas, and seeds that have foliaceous
cotyledons, an exserted radicle and an endosperm. Exceptions
to this character combination are few. Swenson & al. (2007b)
accepted sixty species in Planchonella and another six taxa are
here recovered within the genus. Based on the above diagnostic
character combination and available herbarium material, it is
safe to transfer several species that have not been available for
molecular analysis (see below). Hence, Planchonella contains
approximately 110 known species, of which some are still to
be described, especially from New Caledonia.
Planchonella baillonii with its tubular, rather than urn
shaped, flowers is again found as sister to the rest of the genus
with strong to moderate support (PP 1; JK 88). We believe the
species represents an old evolutionary lineage, the sole extant
member confined to New Caledonia. Planchonella baillonii
is also the type species of Iteiluma, a name rejected in favour
of Planchonella (Swenson & Morat, 2008; Brummitt, 2011a),
but considering its phylogenetic position it is possible to recog
nize a monotypic genus. However, we reject this solution since
P. baillonii possesses the entire set of characters diagnostic for
Planchonella.
Chrysophyllum bakhuizenii was described from New
Guinea by Van Royen (in Vink, 1958) as a close relative of
C. gordoniifolium S. Moore (now in Pycnandra) from New
Caledonia, because of similar morphology and alleged lack of
staminodes. However, examination of one bud of the original
material reveals presence of staminodes and the remaining
morphology falls within the concept of Planchonella. In addi
tion our analyses clearly recover it within the genus.
Planchonella sandwicensis is distributed across all main
islands of Hawaii forming the northeastern outpost of the ge
neric distribution in the Pacific. Triono & al. (2007) used two
accessions that were recovered as sisters, and found sister to the
entire subfamily Chrysophylloideae, casting doubt on whether
this species is a member of Planchonella or even the subfamily.
They further proposed that P. sandwicensis should be sepa
rated and recognized on generic level. Our analysis of three
accessions from Kauai and Oahu found maximum support of
monophyly and the species is retained inside Planchonella, with
close affinity to taxa from Fiji, i.e., P. smithii (P. Royen) A.C. Sm.
and P. umbonata (P. Royen) A.C. Sm. This result is consistent
with earlier findings using nrDNA and morphology (Swenson
& al., 2007a, b). The only reasonable explanation for Triono
& al. (2007) reporting a relationship outside the subfamily is that
they possibly sequenced material of Sideroxylon polynesicum
(Hillebr.) Smedmark & Anderb., a species that is somewhat sim
ilar in leaf venation, but belongs to Sapotoideae, not Chrysophyl
loideae (Smedmark & al., 2006; Smedmark & Anderberg, 2007).
Scattered over the Pacific islands is a polymorphic complex
of Planchonella that has puzzled earlier botanists. Lam (1942)
accepted two varieties of Planchonella costata (Endl.) Pierre, a
classification that Van Royen (1957) amended and extended to
include another two varieties. Smith (1981) in his Flora of Fiji
accounted for nine species of Planchonella, in which he treated
all former varieties as separate species (P. grayana, P. smithii,
P. umbonata) and relegated P. costata to be restricted to New
Zealand and Norfolk Island. The present sample includes six
accessions from Fiji (P. membranacea H.J. Lam, P. smithii,
P. umbonata, P. vitiensis Gillespie, P. “Munzinger 6490”,
and P. “Munzinger 6514”), as well as P. costata from New
Zealand and seven accessions of P. grayana and P. tahitensis
gathered from Futuna in the west to French Polynesia in the
east (but not from Fiji). All of these taxa, except for P. grayana
and P. tahitensis, are scattered throughout the Planchonella
phylogeny and their closest relatives are usually from another
area. Hence, Smith’s (1981) species concept is easy to reconcile
with the molecular phylogeny.
The two species Planchonella grayana and P. tahitensis have
caused confusion as to whether one, two, or several taxa should
be recognized (Grant & al., 1974; Smith, 1981; J.F. Butaud, pers.
comm.). In Fiji, P. grayana occurs from the coast to about 400 m
altitude and usually forms trees 6–14 m tall, but can be stunted to
no more than one meter tall if growing on exposed sites (Smith,
1981). In French Polynesia, P. tahitensis occurs as a small tree
or up to 20 m tall, from sea level to 700 m altitude in Raiatea
and Tahiti, on calcareous or various volcanic soils, often in me
sic to wet forests (Butaud & al., 2011; Pouteau & al., 2012). All
seven accessions analysed here are recovered in a single, strongly
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Swenson & al. • Chrysophylloideae in Oceania and SE Asia
supported clade, with some support for internal resolution, and
all have a unique tripleTinsertion in the ITS sequence (positions
788–790). The clades, however, are incongruent with species
or area (archipelago) of origin. No qualitative morphological
characters differ between the suspected taxa. Both differ from
all congeners in that the stamens are inserted near the base of
the corolla tube (not below the tube orifice). The small greenish
(Fig. 1J) or somewhat whitish flowers are bisexual or female,
which possibly renders the species gynomonoecious, a sexual
system found in Planchonella (Méndez & Munzinger, 2010).
Variable characters are restricted to the size of the foliage and
the fruit. For example, leaves of the sample from Futuna are
elliptic and large (16–25 × 7–10 cm) in comparison to narrowly
elliptic and small (7–11 × 2–3 cm) in the closely related sample
from Tahiti (P. grayana var. florencei Fosberg). Hence, we are
confident that this is one variable, widely distributed species,
adapted to different soils, humidity and altitudes, but we do not
exclude the possibility that a subspecies concept could be appli
cable. In any event, the correct name to be used is P. tahitensis.
Planchonella linggensis is a widely distributed species
in Malesia and many islands in the Pacific. It grows in fairly
different habitats, such as low altitudinal forests on calcareous
soil in Alofi (Wallis and Futuna) and moist rainforests at about
800 m altitude on ultramafic substrate in New Guinea. Previous
phylogenetic analyses have rendered this species paraphyletic,
possibly conspecific with P. chartacea, and it has been sug
gested to form a complex of similar species with unclear spe
cies limits (Swenson & al., 2007a, b). Here, we included three
accessions of P. linggensis, one each from Futuna, New Guinea
and Vanuatu. All accessions group together with maximum
support but are intermingled with P. chartacea and P. solida
(Fig. 2, Clade D2). A cursory inspection of the accessions used
reveals morphological similarities but also differences, such
as presence or absence of indument on different organs. A
molecular study of a wider sample from different areas, soil
types and altitudes, together with a close examination of the
morphology, ought to reveal species limits within this clade.
Unplaced taxa. — Our present findings make significant
progress towards an understanding of the natural groups within
subfamily Chrysophylloideae in Oceania and Southeast Asia,
but there are several unplaced taxa that are still unavailable for
molecular analysis and are of special interest. We acknowledge
that some of these problems may remain unresolved because
the plants are known from few old collections, often treated
with poison such as mercuric chloride. Because of largescale
deforestation, the relationships among these taxa may never be
known. Here, we mention three examples.
Boerlagella spectabilis (Miq.) H.J. Lam, the generic type
of the family Boerlagellaceae (Lam, 1925), is still an enigmatic
taxon. It was originally described from only one leaf and its vein
characters, accompanied by a discussion of the axillary, 5celled
fruit with one seed that has an exserted radicle. This is frag
mented information, but Dubard (1912), as well as Pennington
(1991), associated the species with Planchonella (or Pouteria
sect. Oligotheca (A. DC.) Baehni). Despite the incomplete ma
terial, these characters are in full agreement with the character
combination of Planchonella. Our careful inspection of the
758
available leaf and fruit (type material) reveals the absence of
areolate venation and an overall venation pattern that is similar
to P. torricellensis, a species we predict it is closely related to.
One species from New Guinea, currently classified as
Planchonella kaernbachiana (Engl.) H.J. Lam, has a charac
ter combination that is a mixture between Pichonia (sepals
pubescent inside, stamens in the tube orifice and simple style)
and Planchonella (nonareolate venation, foliaceous cotyledons
and an endosperm). It is unclear if this taxon is simply another
example of the homoplasy typical in Sapotaceae, a taxon of
mixed collections, or if it represents yet another evolutionary
lineage in the area.
Pouteria celebica Erlee from Sulawesi (Celebes), collected
in rainforest at 800 m altitude, is known only from incomplete
type material (floral buds). The overall morphology is very
similar to the frequently cultivated Neotropical P. multiflora
(A. DC.) Eyma. However, the latter has four sepals (Pennington,
1990) rather than five as in P. celebica (HerrmannErlee & Van
Royen, 1957). Future research will tell if this species represents
an early introduction or an extreme case of parallel evolution.
Towards a natural classification. — The present analyses
reconcile earlier findings (Bartish & al., 2005; Swenson & An
derberg, 2005; Swenson & al., 2007a) and strengthen the conclu
sion that there are nine lineages of Chrysophylloideae in Oceania
and Southeast Asia (ten with Xantolis) that warrant formal rec
ognition. Below, we provide a key to the genera using fertile and
leaf material. In addition, each recognized genus is enumerated
with its synonyms, diagnostic character combination, number
of recognized species, distribution, and necessary combinations
that render each genus monophyletic. Homotypic and heterotypic
synonyms are cited for genera, but only the basionym and ho
motypic synonyms are cited for species in order to save space.
Type citations follow the text on the label, including geographic
coordinates if given, or the original description if the type mate
rial has not been viewed (n.v.). For more complete nomenclature,
see Govaerts & al. (2001) and the World Checklist at the Royal
Botanical Gardens, Kew (http://apps.kew.org/wcsp/home.do).
Ongoing revisional work in Australia appears in Flora of Australia, but the volume of Sapotaceae is currently unpublished
(L. Jessup, pers. comm.). Typification of Australian species,
if needed, will appear in this Flora. Typification of Pleioluma
species in New Caledonia will appear in an upcoming revision
we aim to publish in a near future. A summary of the proposed
classification, including new combinations, is shown in Fig. 4.
Key to the genera of Chrysophylloideae in Southeast
Asia and Oceania (excluding Xantolis)
1. Staminodes absent ............................................ 2
1. Staminodes present ........................................... 4
2. Flowers usually along the branches; cotyledons red (or
pinkish); New Caledonia ....................... Pycnandra
2. Flowers axillary; cotyledons white; Australia ........... 3
3. Leaves below pubescent, not translucent; leaf venation
brochidodromous ...................... Amorphospermum
3. Leaves below glabrous or glabrescent (except for veins), ±
translucent; leaf venation eucamptodromous .. Niemeyera
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Swenson & al. • Chrysophylloideae in Oceania and SE Asia
Pleioluma moluccana
Pleioluma macrocarpa
Pleioluma papyracea
Pleioluma singuliflora
Pleioluma xerocarpa
Pleioluma brownlessiana
Pleioluma queenslandica
“Pleioluma acutifolia”
Pleioluma longipetiolata M2264
Pleioluma longipetiolata M2911
Pleioluma baueri
Pleioluma rubicunda
Pleioluma Munzinger2581
Pleioluma firma
Pleioluma wandae
Pleioluma gillisonii
Pleioluma lamprophylla
Pleioluma ledermannii
Pleioluma laurifolia
Pleioluma Takeuchi25691
Pleioluma sebertii
Pleioluma vieillardii
Pleioluma novocaledonica
Pleioluma lucens
Pleioluma lasiantha
Pleioluma crebrifolia
“Pleioluma belepensis”
Pleioluma balansana
Pleioluma balansana
Van-royena castanosperma
Sersalisia sessiliflora
Sersalisia sericea
Sersalisia luzoniensis
Pichonia occidentalis
Pichonia deplanchei
Pichonia lecomtei
Pichonia daenikeri
Pichonia balansana
Pichonia grandiflora
Pichonia balansae
Pichonia dubia S918
Pichonia dubia M3065
Magodendron mennyae
Amorphospermum antilogum
Niemeyera whitei
Niemeyera Ford2429
Niemeyera prunifera
Niemeyera chartacea
Pycnandra canaliculata
Pycnandra acuminata
“Pycnandra wagapensis”
“Pycnandra filipes”
“Pycnandra grandifolia”
“Pycnandra goroensis”
Pycnandra ouaiemensis
Pycnandra comptonii
Pycnandra blaffartii
Pycnandra controversa
Pycnandra vieillardii
Pycnandra griseosepala
Pycnandra neocaledonica
Pycnandra kaalaensis
Pycnandra decandra
Pycnandra belepensis
Pycnandra bracteolata
Pycnandra glabella
Pycnandra sessilifolia
Pycnandra francii
Pycnandra schmidii
Pycnandra pubiflora
Pycnandra caeruleilatex
Pycnandra sessiliflora
Pycnandra sarlinii
Pycnandra deplanchei
Pycnandra fastuosa
Pycnandra carinocostata
Pycnandra viridiflora
Pycnandra blanchonii
Pycnandra gordoniifolia
Pycnandra paucinervia
Pycnandra atrofusca
Pycnandra benthamii
Pycnandra glaberrima
Pycnandra balansae
Pycnandra cylindricarpa
Pycnandra linearifolia
A
B
C
D
E
F
F
Planchonella
Planchonella baillonii
Planchonella malaccensis
Planchonella maclayana
Planchonella linggensis Vanuatu
Planchonella chartacea Australia
Planchonella linggensis Futuna
Planchonella garcinioides New Guinea
Planchonella linggensis New Guinea
Planchonella xylocarpa
Planchonella xylocarpa
Planchonella stellibacca
Planchonella maingayi
Planchonella sphaerocarpa
Planchonella arnhemica
Planchonella pohlmaniana
Planchonella asterocarpon
Planchonella membranacea
Planchonella dothioensis
Planchonella thiensis
Planchonella laetervirens
Planchonella mandjeliana
Planchonella cauliflora
Planchonella luteocostata
Planchonella amieuana
Planchonella endlicheri
Planchonella ericiflora
Planchonella Pillon150
Planchonella kaalaensis
Planchonella koumaciensis
Planchonella Munzinger6150
Planchonella minutiflora
Planchonella pronyensis
Planchonella rufocostata
Planchonella saligna
Planchonella crassinervia
Planchonella roseoloba
Planchonella skottsbergii
Planchonella kuebiniensis
Planchonella howeana
Planchonella myrsinifolia
Planchonella cotinifolia
Planchonella eerwah
Planchonella costata
Planchonella australis
Planchonella myrsinoides
Planchonella euphlebia
Planchonella Ile Yande
Planchonella lauracea
Planchonella glauca
Planchonella lamii
Planchonella clemensii
Planchonella obovata
Planchonella mindanaensis
Planchonella Munzinger6514 Fiji
Planchonella vitiensis Fiji
Planchonella aneityensis Vanuatu
Planchonella tahitensis Raiatea
Planchonella tahitensis Tuamotu
Planchonella tahitensis Tahiti
Planchonella tahitensis Alofi
Planchonella tahitensis Futuna
Planchonella tahitensis Australs
Planchonella tahitensis Tahiti
Planchonella pullenii
Planchonella Armstrong316
Planchonella Munzinger6490
Planchonella smithii
Planchonella umbonata
Planchonella sandwicensis K119
Planchonella sandwicensis Okamoto
Planchonella sandwicensis M3227
Planchonella cyclopensis P1365
Planchonella anteridifera
Planchonella thyrsoidea
Planchonella cyclopensis T25495
Planchonella dulcitan
Planchonella torricellensis Futuna
Planchonella torricellensis Alofi
Planchonella forbesii
Planchonella polyneura
Fig. 4. Summary of proposed classification and necessary combinations of Oceanian and Southeast Asian Chrysophylloideae (Sapotaceae) based
on BEAST and jackknife analyses of nrDNA sequences. Accepted genera are (A) Pleioluma, (B) Sersalisia, (C) Pichonia, (D) Niemeyera, (E)
Pycnandra, (F) Planchonella, and the small genera Amorphospermum, Magodendron and Van-royena. Types of generic names are indicated in
bold. Recently discovered species appear with collector and number, whereas species under revision and are not yet published appear in quota
tion marks.
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4.
4.
5.
5.
6.
Tertiary or quaternary leaf venation areolate ............ 5
Tertiary or quaternary leaf venation never areolate ..... 8
Flowers born on trunk ...................... Magodendron
Flowers axillary ............................................... 6
Stamens inserted in lower half of corolla tube; foliaceous
cotyledons .......................................... Pleioluma
6. Stamens inserted in or just below corolla tube orifice; cot
yledons planoconvex ........................................ 7
7. Flowers cupshaped; style simple ................ Pichonia
7. Flowers narrowly campanulate or tubular; style with sev
eral stigmatic areas ................................ Sersalisia
8. Stamens inserted just below corolla tube orifice; foliaceous
cotyledons ...................................... Planchonella
8. Stamens inserted near middle of corolla tube; planoconvex
cotyledons ........................................ Van-royena
Amorphospermum F. Muell., Fragm. 7: 112. 1870 – Type:
Amorphospermum antilogum F. Muell., Fragm. 7: 113.
1870 ≡ Lucuma amorphosperma F.M. Bailey, Queensl.
Fl. 3: 955. 1900, nom. illeg. ≡ Sersalisia antiloga (F. Muell.)
Domin in Biblioth. Bot. 89: 508. 1928 ≡ Chrysophyllum
antilogum (F. Muell.) Vink in Blumea 9: 65. 1958 ≡ Niemeyera antiloga (F. Muell.) T.D. Penn., Gen. Sapotac.: 235.
1991 – Lectotype (designated by Vink, 1958): Australie,
Queensland, Thozet s.n., 1870 (MEL n.v.; isolectotype, P!).
Diagnostic character combination. – Leaves pubescent
below, not translucent; tertiary leaf venation parallel, higher
venation nonareolate; sepals glabrous inside; flowers axillary,
cupshaped; stamens inserted in corolla tube orifice; stami
nodes absent; style simple; cotyledons planoconvex, white;
radicle included in cotyledons; endosperm absent.
A single species in forests of eastern Australia, from New
South Wales to Papua New Guinea. There seems to exist an
other, as yet undescribed species (L. Jessup, pers. comm.).
Magodendron Vink in Nova Guinea, n.s., 8: 124. 1957 – Type:
Magodendron venefici (C.T. White & W.D. Francis) Vink
in Nova Guinea, n.s., 8: 125. 1957 ≡ Achradotypus venefici
C.T. White & W.D. Francis in Proc. Roy. Soc. Queensland
38: 254. 1927.
Diagnostic character combination. – Leaf venation areo
late; sepals glabrous inside; flowers born on trunk, cupshaped;
stamens inserted in corolla tube orifice; staminodes present,
irregularly incised; style with stigmatic areas; cotyledons
planoconvex with a radicle extending below cotyledon com
missure; endosperm absent.
Two species confined to New Guinea (Vink, 1995).
Niemeyera F. Muell., Fragm. 7: 114. 1870, nom. cons., Taxon 9:
78. 1960 – Type: Niemeyera prunifera (F. Muell.) F. Muell.,
≡ Chrysophyllum pruniferum F. Muell., Fragm. 6: 26. 1867.
Diagnostic character combination. – Leaves glabrous or
glabrescent below, usually translucent; tertiary leaf venation
oblique, nonareolate; sepals glabrous inside; flowers axillary,
cupshaped; stamens inserted in corolla tube orifice; stami
nodes absent; style simple; cotyledons planoconvex, white;
radicle included in cotyledons; endosperm absent.
760
Three described species and one undescribed species
(L. Jessup, pers. comm.), all confined to Australia. No modern
treatment is available, but will appear in Flora of Australia.
Pichonia Pierre, Not. Bot.: 22. 1890 ≡ Epiluma Baill., Hist. Pl.
11: 287. 1891, nom. illeg. (superfl.) – Type: Pichonia balansana Pierre, Not. Bot.: 23. 1890 (≡ Epiluma pyriformis
Baill., nom. illeg. (superfl.)).
= Rhamnoluma Baill., Hist. Pl. 11: 287. 1891 – Type: Rhamnoluma novocaledonica (Engl.) Baill., Hist. Pl. 11: 288.
1891 ≡ Lucuma novocaledonica Engl. in Bot. Jahrb. Syst.
12: 516. 1890.
= Wokoia Baehni in Arch. Sci. 17: 78. 1964 – Type: Wokoia
rhopalocarpa (P. Royen) Baehni ≡ Pouteria rhopalocarpa
P. Royen in Nova Guinea, n.s., 10: 134. 1959.
= Arnanthus Baehni in Arch. Sci. 17: 78. 1964 – Type: Arnanthus
balansae (Baill.) Baehni ≡ Chrysophyllum balansae Baill.
in Bull. Mens. Soc. Linn. Paris 2: 901. 1891, nom. illeg.
Diagnostic character combination. – Tertiary leaf vena
tion laxly reticulate, higher venation areolate; sepals usually
pubescent inside in upper part or sometimes glabrous; flow
ers usually axillary, cupshaped; stamens inserted in corolla
tube orifice; staminodes present, generally entire; style sim
ple; cotyledons planoconvex, radicle included in cotyledons;
endosperm absent.
Twelve known species, of which seven are endemic to
New Caledonia (Swenson & Munzinger, 2012) and five to
New Guinea, one reaching as far east as the Solomon Islands.
Pichonia hochreutineri (H.J. Lam) Swenson, comb. nov. ≡
Planchonella hochreutineri H.J. Lam in Boissiera 7: 92.
1943 ≡ Pouteria hochreutineri (H.J. Lam) H.J. Lam in
Blumea 5: 337. 1943 – Holotype: New Guinea, Boridi, ca.
1400 ft alt., 9.IX.1935, C.E. Carr 13023 (L!; isotypes BM!,
K!, SING!).
Distribution: New Guinea.
Pichonia rhopalocarpa (P. Royen) Swenson, comb. nov. ≡
Pouteria rhopalocarpa P. Royen in Nova Guinea, n.s., 10:
134. 1959 ≡ Wokoia rhopalocarpa (P. Royen) Baehni in
Arch. Sci 17: 78. 1964 – Holotype: West New Guinea,
Manokwari District, Oransbari, alt. 50 m, 11.X.1955,
R.P. Mangold 51 (L!; isotype MAN n.v.).
Distribution: New Guinea.
Planchonella Pierre, Not. Bot.: 34. 1890, nom. cons., Taxon 57:
1013. 2008 – Type: Planchonella obovata (R. Br.) Pierre,
Not. Bot.: 36. 1890 ≡ Sersalisia obovata R. Br., Prodr.:
529. 1810.
= Iteiluma Baill. in Bull. Mens. Soc. Linn. Paris 2: 892. 1890,
nom. rej., Taxon 57: 1013. 2008 – Type: Iteiluma baillonii
(Zahlbr.) Baill., Hist. Pl. 11: 286. 1891 ≡ Lucuma baillonii
Zahlbr. in Oesterr. Bot. Z. 39: 287. 1889 ≡ Poissonella
Pierre, Not. Bot.: 29. 1890.
= Peuceluma Baill. in Bull. Mens. Soc. Linn. Paris 2: 895.
1890, nom. rej., Taxon 57: 1013. 2008 – Type: Peuceluma
pinifolia Baill.
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Swenson & al. • Chrysophylloideae in Oceania and SE Asia
= Beauvisagea Pierre, Not. Bot.: 15. 1890 – Type: Beauvisagea
pomifera Zippel ex Baill., Hist. Pl. 11: 288. 1891.
= Bureavella Pierre, Not. Bot.: 16. 1890 – Type: Bureavella maclayana (F. Muell.) Pierre ≡ Bassia maclayana F. Muell.,
Vict. Chem. & Druggist 7 (Apr.): 93. 1885.
= Fontbrunea Pierre, Not. Bot.: 31. 1890 – Type: Fontbrunea
malaccensis (C.B. Clarke) Pierre, Not. Bot.: 32. 1890 ≡
Sideroxylon malaccense C.B. Clarke in Hooker, Fl. Brit.
India 3: 537. 1882.
= Krausella H.J. Lam in Nova Guinea 14: 566. 1932 – Type:
Krausella polyneura (K. Krause) H.J. Lam in Nova Guinea
14: 567. 1932 ≡ Sideroxylon polyneurum K. Krause in Bot.
Jahrb. Syst. 58: 475. 1923.
= Albertisiella Pierre ex Aubrév. in Adansonia, n.s., 4: 42. 1964
– Type: Albertisiella novoguineensis (Vink) Aubrév. ≡
Chrysophyllum novoguineense Vink in Blumea 9: 67. 1958.
= Blabeia Baehni in Arch. Sci. 17: 77. 1964 – Type: Blabeia endlicheri (Montrouz.) Baehni ≡ Sapota endlicheri Montrouz.
in Mém. Acad. Roy. Sci. Lyon, Sect. Sci. 10: 228. 1860.
= Pyriluma Aubrév., Fl. Nouv.Caléd. 1: 83. 1967 – Type: Pyriluma sphaerocarpa (Baill.) Aubrév., Fl. Nouv.Caléd. 1:
84. 1967 ≡ Sideroxylon sphaerocarpa Baill. in Bull. Mens.
Soc. Linn. Paris 2: 891. 1890.
Diagnostic character combination. – Tertiary leaf vena
tion usually reticulate or oblique, higher venation nonareolate;
sepals glabrous inside or rarely pubescent to varying degree;
flowers usually axillary, urnshaped or rarely tubular; stamens
inserted just below corolla tube orifice, rarely in middle of tube
or near base; staminodes present, usually entire; style with
stigmatic areas; cotyledons foliaceous with a radicle extending
below cotyledon commissure; endosperm present.
Planchonella is the largest genus of Chrysophylloideae in
the Old World with approximately 110 known species. Several
species in New Caledonia and the west Pacific Islands remain
to be described (or resurrected). Planchonella is distributed
from Thailand and southern China in the north, through Male
sia, Australia, New Caledonia, and on to the Pacific Islands,
reaching French Polynesia and Hawaii. The highest diversity
is in New Caledonia (ca. 40 spp.), followed by New Guinea
(ca. 30 spp.), and Australia (12 spp.). No modern revision is
available, but see Aubréville (1967) for New Caledonia and
Swenson & al. (2007b) for an amended generic description.
Note. – Planchonella is conserved against Iteiluma and
Peuceluma (Swenson & Morat, 2008; Brummitt, 2011a). How
ever, Pierre (1890) described Planchonella, Beauvisagea,
Bureavella, and Fontbrunea in the same publication, i.e., four
competing names with equal priority. Swenson & al. (2007a)
found Bureavella to be embedded in Planchonella and chose the
latter over the former. Since no choice has been made between
Planchonella, Beauvisagea, and Fontbrunea, we choose, in ac
cordance with Article 11.5 in McNeill & al. (2012), Planchonella
over Beauvisagea and Fontbrunea, as the latter two have never
(or very rarely) been used, and therefore the choice requires
fewer new combinations, retaining nomenclatural stability.
Planchonella forbesii (S. Moore) H.J. Lam in Bull. Jard. Bot.
Buitenzorg, ser. 3, 7: 217. 1925 ≡ Sideroxylon forbesii
S. Moore in J. Bot. 61(Suppl.): 30. Jun 1923 ≡ Pouteria
forbesii (S. Moore) Baehni in Candollea 9: 408. 1942 ≡
Krausella forbesii (S. Moore) H.J. Lam in Boissiera 7: 92.
1943 – Holotype: New Guinea, Sogeri Region, Mt. Wori
Wori, alt. 5000 ft, 1885–6, H.O. Forbes 756 (MEL n.v.;
isotypes E!, FI n.v, L!, K!).
= Sideroxylon patentinervium K. Krause in Bot. Jahrb. Syst. 58:
474. Nov 1923 ≡ Krausella patentinervia (K. Krause) Erlee
in Blumea 8: 448. 1957 – Lectotype (designated by Herr
mannErlee & Lam, 1957): KaiserWilhelmsland [New
Guinea], Kani Geberges, 1000 m alt., 31 October 1907,
R. Schlechter 16746 (P!).
Distribution: New Guinea.
Note. – We used an accession determined as Krausella patentinervia. A careful examination of the types and additional
material reveals that the leaf characters used by HerrmanErlee
& Lam (1957) to distinguish the above species are overlapping
and the two are better conceived as conspecific, but different
from the generic type K. polyneura. The name Sideroxylon
forbesii was published a few months before S. patentinervium
and has priority. We mention this here in order to avoid an
unnecessary combination.
Planchonella garcinioides (K. Krause) Swenson, comb. nov.
≡ Sideroxylon garcinioides K. Krause in Bot. Jahrb. Syst.
58: 477. 1923 ≡ Lucuma garcinioides (K. Krause) H.J. Lam
in Nova Guinea 14: 568. 1932 ≡ Pouteria garcinioides
(K. Krause) Baehni in Candollea 9: 338. 1942 – Lectotype
(designated by Vink, 2002): Neu Guinea, SepikGebiet, im
Alluvialwald am Sepik, beim Lager Malu, alt. 20–40 m,
I.1913, Ledermann 10728 (L!) (holotype B †).
= Planchonella solida P. Royen in Blumea 8: 404, 433. 1957
– Holotype: New Guinea, Yalu, near Lae, 5.III.1950,
D. Fryar 3344 (SING!; isotypes L!, LAE n.v.).
Distribution: New Guinea.
Note. – HerrmannErlee & Van Royen (1957), in their re
vision of Pouteria, repeated the original Latin description of
Pouteria garcinioides since no material was available and the
type in Berlin was destroyed during World War II. Simultane
ously, Van Royen (1957) described Planchonella solida, but
Wim Vink (Leiden) united these two species in an unpublished
record. We have used an accession of P. solida, but this species
should be united with Sideroxylon garcinioides, a name that has
priority but needs to be combined with Planchonella.
Planchonella lamii P. Royen in Blumea 8: 398, 432. 1957 –
Holotype: Amboina, near Sirimau, 450 m alt., 11.II.1930,
NIFS bb 14283 (L!).
= Chrysophyllum bakhuizenii P. Royen in Blumea 9: 74. 1958
– Holotype: New Guinea, Alola, alt. 2000 m, C.E. Carr
14159 (L!; isotype BM n.v.).
Distribution: Maluku Islands, New Guinea.
Note. – We sequenced an accession determined as Chrysophyllum bakhuizenii, but an examination of the available mate
rial of this species and Planchonella lamii shows that they are
conspecific and should be united.
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761
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Planchonella maclayana (F. Muell.) Swenson, comb. nov. ≡
Bassia maclayana F. Muell. in Vict. Chem. & Druggist 7
(Apr.): 93. 1885 ≡ Illipe maclayana (F. Muell.) F. Muell.,
Descr. Notes Papuan Pl. 2(6): 12. 1885 ≡ Bureavella maclayana (F. Muell.) Pierre, Not. Bot.: 16. 1890 ≡ Lucuma
maclayana (F. Muell.) H.J. Lam in Bull. Jard. Bot. Buiten
zorg, ser. 3, 7: 22. 1925 ≡ Pouteria maclayana (F. Muell.)
Baehni in Candollea 9: 307. 1942 – Type: New Guinea,
MiklouhoMaclay (MEL!, P!).
Distribution: Widespread in Malesia, Solomon Islands.
Note. – Lectotypifying this species is here avoided since the
type material in the Melbourne and Paris herbaria are of seed
fragments only. Ongoing revisional work for the Flora of Australia may identify additional material that is in better condition.
Planchonella macrantha (Merr.) Swenson, comb. nov. ≡
Sideroxylon macranthum Merr. in Publ. Bur. Sci. Gov.
Lab. 35: 56. 1906 (“1905”) ≡ Lucuma macrantha (Merr.)
H.J. Lam in Bull. Jard. Bot. Buitenzorg, ser. 3, 7: 225.
1925 ≡ Pouteria macrantha (Merr.) Baehni in Candol
lea 9: 328. 1942 ≡ Bureavella macrantha (Merr.) Aubrév.
in Adansonia, n.s., 3: 331. 1963 – Holotype: Philippines,
Luzon, Province Bataan, Lamoa River, Mt. Mariveles,
March 1905, T.E. Borden 21736 (= Forestry Bureau 2741)
(PNH †; isotypes BO!, K!, SING!, US!).
Distribution: Philippines to Maluku.
Planchonella malaccensis (C.B. Clarke) Swenson, comb. nov.
≡ Sideroxylon malaccense C.B. Clarke in Hooker, Fl. Brit.
India 3: 537. 1882 ≡ Fontbrunea malaccensis (C.B. Clarke)
Pierre, Not. Bot.: 31. 1890 ≡ Lucuma malaccensis
(C.B. Clarke) Dubard in Ann. Mus. Colon. Marseille, sér. 2,
10: 19. 1912 ≡ Pouteria malaccensis (C.B. Clarke) Baehni in
Candollea 9: 302. 1942 ≡ Xantolis malaccensis (C.B. Clarke)
Baehni in Boissiera 11: 23. 1965 – Holotype: Malaya, 1871,
A.C. Maingay 994 (SING!; isotypes K!, L!, P!).
Widely distributed: Thailand, Malay Peninsula, Sumatra,
Sulawesi, Borneo and New Guinea.
Planchonella menait (Vink) Swenson, comb. nov. ≡ Pouteria
menait Vink in Blumea 47: 131. 2002 – Holotype: Papua
New Guinea, East Sepik Province, Hunstein Range (Mt.
Samsai), at the site “Camp 3” on slopes above the main
streamcourse, 450 m alt., 04°29′ S, 142°41′ E, 19.VII.1990,
W. Takeuchi 6276 (L!; isotype L!).
Distribution: New Guinea.
Planchonella orkor (Vink) Swenson, comb. nov. ≡ Pouteria
orkor Vink in Blumea 47: 134. 2002 – Holotype: New
Guinea, Saidor Subdistrict, NahoRawa, Budemu, 4150 ft
alt., 24.X.1964, C.D. Sayers NGF 21329 (L!; isotypes BM!,
LAE n.v.).
Distribution: New Guinea.
Planchonella paucinervia (Erlee) Swenson, comb. nov. ≡ Pouteria paucinervia Erlee in Blumea 8: 503. 1957 – Holotype:
Indonesia, Sumatra, Riouw District, Indragiri, Moeata
762
Pedjanki, 11.IV.1939, P. Buwalda 6503 (L!; isotypes A n.v.,
BO n.v., PNH!, SING!).
Distribution: Malay Peninsula, Sumatra.
Planchonella polyneura (K. Krause) Swenson, comb. nov. ≡ Sideroxylon polyneurum K. Krause in Bot. Jahrb. Syst. 58: 475.
1923 ≡ Krausella polyneura (K. Krause) H.J. Lam in Nova
Guinea 14: 567. 1932 ≡ Pouteria multinervis T.D. Penn.,
Gen. Sapotac.: 202. 1991 – Lectotype (designated by Her
rmannErlee & Lam, 1957): DeutschNeuguinea, Sepikge
biet, Ledermann 9054 (K!; isolectotype SING!).
Distribution: New Guinea.
Planchonella pullenii (Vink) Swenson, comb. nov. ≡ Pouteria
pullenii Vink in Blumea 47: 98. 2002 – Holotype: New
Guinea, Milne Bay District, Baniara Subdistrict, south
of Opanabu village, 149°43′ E, 10°01′ S, 600–700 m alt.,
16.VII.1969, A. Kanis 1243 (L!; isotypes A n.v., BRI n.v.,
CANB n.v., CHR n.v., K n.v., LAE n.v.).
Distribution: New Guinea.
Planchonella ridsdalei (Vink) Swenson, comb. nov. ≡ Pouteria ridsdalei Vink in Blumea 47: 136. 2002 – Holotype:
Philippines, Mindoro, Mt. Halcon area near Paitan, Dulan
gan Range, 13°12′ N, 121°12′ E, 8.V.1986, C.E. Ridsdale
1695 (L!; isotype K!).
Distribution: Philippines.
Planchonella stellibacca (J.F. Maxwell) Swenson, comb. nov.
≡ Pouteria stellibacca J.F. Maxwell in Nat. Hist. Bull
Siam Soc. 50: 90. 2002 – Holotype: Thailand, Nakhon
Nayok Province, Muang District, Khao Yai National Park,
14°24.5′ N, 101°23′ E, Klong Sai area, 760 m alt., 8.III.2001,
P. Charoenchai 1024 (CMU n.v.; isotypes A n.v., BHF n.v.,
CAS n.v., L!).
Distribution: Borneo, Thailand.
Planchonella villamilii (Merr.) Swenson, comb. nov. ≡ Sideroxylon villamilii Merr. in Philipp. J. Sci., C 10: 59. 1915
≡ Pouteria villamilii (Merr.) Baehni in Candollea 9: 318.
1942 ≡ Bureavella villamilii (Merr.) Aubrév. in Adansonia,
n.s., 3: 331. 1963 – Holotype: Philippines, Luzon, Prov
ince of Laguna, VI.1913, A. Villamil, Forest Bureau 19762
(PNH, not found; isotype US!).
Distribution: Philippines.
Planchonella whitmorei (Vink) Swenson, comb. nov. ≡ Pouteria whitmorei Vink in Blumea 47: 142. 2002 – Holotype:
Solomon Islands, south Vella Lavella Island, Oula River
area, 14.VIII.1968, C. Kotali & al. BSIP 11182 (L!).
Distribution: Solomon Islands.
Pleioluma (Baill.) Baehni in Boissiera 11: 150. 1965 ≡ Sersalisia sect. Pleioluma Baill., Hist. Pl. 11: 280. Sep–Oct 1891 –
Type: Pleioluma crebrifolia (Baill.) Swenson & Munzinger
≡ Lucuma crebrifolia Baill. in Bull. Mens. Soc. Linn. Paris
2: 897. 1891.
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= Beccariella Pierre, Not. Bot.: 30. 1890, nom. illeg., non
Beccariella Ces. in Atti. Reale Accad. Sci. Fis. 8: 9. 1879
– Type: Beccariella sebertii (Pancher) Pierre ≡ Chrysophyllum sebertii Pancher in Sebert & Pancher, Not. Bois
Nouv. Caléd.: 194. 1874.
Diagnostic character combination. – Leaf venation areo
late; sepals pubescent inside; flowers usually axillary, narrowly
campanulate; stamens inserted in middle or at base of corolla
tube; staminodes present, usually entire; style with stigmatic
areas; cotyledons foliaceous with a radicle extending below
cotyledon commissure; endosperm present.
About 30 species are here attributed to Pleioluma, but sev
eral species are to be described from New Caledonia and New
Guinea. The highest diversity is in New Guinea (14 spp.) and
New Caledonia (13 spp.), extending to Malesia and Southeast
Asia. The only available treatment, then under Beccariella, is
in the Flora of New Caledonia by Aubréville (1967).
Pleioluma azou (P. Royen) Swenson & Munzinger, comb. nov.
≡ Planchonella azou P. Royen in Blumea 8: 308, 428. 1957
≡ Beccariella azou (P. Royen) Aubrév. in Adansonia, n.s.,
2: 193. 1962 – Holotype: NouvelleCalédonie, Port Boisé,
zone maritime, X.1903, Cribs 1424 (P!; isotype L!).
Distribution: New Caledonia.
Pleioluma balansana (Pierre ex Baill.) Swenson & Munzinger,
comb. nov. ≡ Sideroxylon balansanum Pierre ex Baill. in
Bull. Mens. Soc. Linn. Paris 2: 889. 1890 ≡ Planchonella
balansana (Pierre ex Baill.) Pierre ex Dubard in Ann. Mus.
Colon. Marseille, sér. 2, 10: 46. 1912 ≡ Pouteria balansana
(Pierre ex Baill.) Baehni in Candollea 9: 317. 1942 ≡ Beccariella balansana (Pierre ex Baill.) Aubrév. in Adansonia,
n.s., 2: 193. 1962 – Lectotype (designated by Van Royen,
1957): Nouvelle Calédonie, 11.IV.1869, Balansa 1327a (P!).
Distribution: New Caledonia. Beccariella brevipedicellata
sensu Aubréville (1967), a later described taxon, is possibly
conspecific with P. balansana but is not put into synonymy
here; instead we await future results.
Pleioluma baueri (Montrouz.) Swenson & Munzinger, comb.
nov. ≡ Sapota baueri Montrouz. in Mem. Acad. Roy. Sci.
Lyon, Sect. Sci. 10: 229. 1860 ≡ Planchonella baueri (Mon
trouz.) Dubard in Ann. Mus. Colon. Marseille, sér. 2, 10: 53.
1912 ≡ Pouteria baueri (Montrouz.) Baehni in Candollea 9:
329. 1942 ≡ Beccariella baueri (Montrouz.) Aubrév. in Adan
sonia, n.s., 2: 193. 1962 – Holotype: NouvelleCalédonie, Ile
Art, Montrouzier 133 (P; isotype G).
Distribution: New Caledonia.
Pleioluma brownlessiana (F. Muell.) Swenson & Munzinger,
comb. nov. ≡ Achras brownlessiana F. Muell., Fragm. 7: 111.
1870 ≡ Sideroxylon brownlessianum (F. Muell.) F. Muell.,
Syst. Census Austral. Pl.: 92. 1882 ≡ Sersalisia brownlessiana (F. Muell.) Domin in Biblioth. Bot. 89: 508. 1928 ≡ Pouteria brownlessiana (F. Muell.) Baehni in Candollea 9: 318.
1942 ≡ Planchonella brownlessiana (F. Muell.) P. Royen in
Blumea 8: 343. 1957 ≡ Beccariella brownlessiana (F. Muell.)
Swenson & al. in Cladistics 23: 221. 2007 – Type: Australia,
Queensland, Rockingham Bay, Dallachy s.n. (MEL!).
Distribution: Australia.
Pleioluma crebrifolia (Baill.) Swenson & Munzinger, comb.
nov. ≡ Lucuma crebrifolia Baill. in Bull. Mens. Soc. Linn.
Paris 2: 897. 1891 ≡ Sideroxylon crebrifolium (Baill.) Engl.
in Engler & Prantl, Nat. Pflanzenfam., Nachtr. 1: 277. 1897
≡ Planchonella crebrifolia (Baill.) Pierre ex Dubard in
Ann. Mus. Colon. Marseille, sér. 2, 10: 53. 1912 ≡ Beccariella crebrifolia (Baill.) Aubrév. in Adansonia, n.s., 2: 193.
1962 ≡ Pouteria crebrifolia (Baill.) Baehni in Boissiera 11:
59. 1965 – Lectotype (designated here): Nouvelle Calédo
nie, Collines éruptives entre Canala et Couaoua, IV.1871,
Balansa 3154 (P00282377!; isolectotypes P00282375!,
P00282376!, P00282378!).
Distribution: New Caledonia.
Note. – Baillon (1891a) described Lucuma crebrifolia based
on three collections, viz. Vieillard 191, Vieillard 2906, and
Balansa 3154, all deposited in P. Later that year, Baillon (1891b)
described section Pleioluma of the genus Sersalisia, referring
to his earlier work, but did not assign any type collection. Lucuma crebrifolia remained untypified until Aubréville (1967)
stated, in his Flora of New Caledonia, “Holotype: Balansa 3154
(P)”. Effectively, according to Article 9.9 (McNeill & al., 2012),
Aubréville implicitly made a lectotypification. However, there
are four different specimens of equal priority of Balansa 3154
in Paris, none being annotated as holo or lectotype. Hence,
in accordance with Article 9.17 (McNeill & al., 2012), one
wellpreserved and fertile specimen of Balansa 3154 is here
designated as lectotype.
Pleioluma densinervia (K. Krause) Swenson, comb. nov. ≡
Sideroxylon densinervium K. Krause in Bot. Jahrb. Syst. 58:
476. 1923 ≡ Planchonella densinervia (K. Krause) H.J. Lam
in Nova Guinea 14: 562. 1932 ≡ Pouteria densinervia
(K. Krause) Baehni in Candollea 9: 342. 1942 – Lectotype
(designated by Van Royen, 1957): Neuguinea, Sepikgebiet,
1912/13, Ledermann 12698 (L!; isolectotypes BM!, K!).
Distribution: New Guinea.
Pleioluma dies-reginae (P. Royen) Swenson, comb. nov. ≡
Planchonella dies-reginae P. Royen in Blumea 8: 352, 431.
1957 ≡ Pouteria dies-reginae (P. Royen) Vink in Blumea
47: 103. 2002 – Holotype: New Guinea, 4 km SW of Bern
hard Camp, Idenburg River, III.1939, L.J. Brass & C. Versteegh 13150 (L!; isotype A n.v.).
Distribution: New Guinea.
Pleioluma firma (Miq.) Swenson, comb. nov. ≡ Chrysophyllum firmum Miq., Fl. Ned. Ind., Eerste Bijv.: 579. 1861 ≡
Sideroxylon firmum (Miq.) Pierre ex Burck in Ann. Jard.
Bot. Buitenzorg 5: 17. 1885 ≡ Beccariella firma (Miq.)
Pierre, Not. Bot.: 30. 1890 ≡ Planchonella firma (Miq.)
Dubard in Ann. Mus. Colon. Marseille, sér. 2, 10: 59. 1912
≡ Pouteria firma (Miq.) Baehni in Candollea 9: 284. 1942
– Holotype: Teysmann s.n. (BO n.v.).
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Swenson & al. • Chrysophylloideae in Oceania and SE Asia
Distribution: Widespread, from Thailand via Malesia to
Solomon Islands.
Pleioluma foxworthyi (Elmer) Swenson, comb. nov. ≡ Sideroxylon foxworthyi Elmer in Leafl. Philipp. Bot. 5: 1836.
1913 ≡ Planchonella foxworthyi (Elmer) H.J. Lam in Proc.
Pacific Sci. Congr. 6(4): 678. 1940 – Lectotype (designated here): Philippine Islands, Island of Palawan, Puerto
Princesa, Mt. Pulgar, III.1911, Elmer 12824 (P00648085!;
isolectotypes E!, FI n.v., G!, L!, NSW n.v., P!, US!, Z!)
(holotype PNH †).
Distribution: Philippines.
Pleioluma gillisonii (Vink) Swenson, comb. nov. ≡ Pouteria gillisonii Vink in Blumea 47: 104. 2002 – Holotype:
Papua New Guinea, Morobe Province, Morobe Subprov
ince, Waiu Bay, 7°30′ S, 147°15′ E, 300 m alt., 14.VIII.1968,
Gillison & Kairo NGF 25627 (L!; isotypes A n.v., BRI n.v.,
CANB n.v., CHR n.v., LAE n.v.).
Distribution: New Guinea.
Pleioluma krausei (H.J. Lam) Swenson, comb. nov. ≡ Sideroxylon spathulatum K. Krause in Bot. Jahrb. Syst. 58: 474.
1923, nom. illeg., non Hillebr., Fl. Hawaiian Isl.: 277. 1888 ≡
Planchonella krausei H.J. Lam in Nova Guinea 14: 561. 1932
≡ Pouteria krausei (H.J. Lam) Baehni in Candollea 9: 322.
1942 – Lectotype (designated by Van Royen, 1957): Nieuw
Guinea, Kaiser Wilhelmsland, Gomadjiji, ca. 450 m alt.,
23.VIII.1909, R. Schlechter 19915 (L!; isolectotypes BR!, P!).
Distribution: New Guinea.
Pleioluma lamprophylla (K. Krause) Swenson, comb. nov. ≡
Sideroxylon lamprophyllum K. Krause in Bot. Jahrb. Syst.
58: 481. 1923 ≡ Planchonella lamprophylla (K. Krause)
H.J. Lam in Nova Guinea 14: 564. 1932 ≡ Pouteria lamprophylla (K. Krause) Baehni in Candollea 9: 333. 1942 –
Lectotype (designated by Van Royen, 1957): NeuGuinea,
SepikGebeit, 1912–1913, Ledermann 10318 (L!; isolecto
type K!).
Distribution: New Guinea.
Pleioluma lanatifolia (P. Royen) Swenson, comb. nov. ≡ Planchonella lanatifolia P. Royen in Blumea 8: 306, 428. 1957 ≡
Pouteria lanatifolia (P. Royen) Vink in Blumea 47: 107. 2002
– Holotype: Dutch New Guinea, Angi, Arfak Mts., 2200 m
alt., 9.IV.1940, R. Kanehira & S. Hatusima 13891 (A!).
Distribution: New Guinea.
Pleioluma lasiantha (Baill.) Swenson & Munzinger, comb.
nov. ≡ Sideroxylon lasianthum Baill. in Bull. Mens. Soc.
Linn. Paris 2: 887. 1890 ≡ Planchonella lasiantha (Baill.)
Dubard in Ann. Mus. Colon. Marseille, sér. 2, 10: 58. 1912
≡ Pouteria lasiantha (Baill.) Baehni in Candollea 9: 324.
1942 ≡ Beccariella lasiantha (Baill.) Aubrév., Fl. Nouv.
Caléd. 1: 114. 1967 – Holotype: NouvelleCalédonie, Mont
Mi, 25.III.1869, Balansa 1322 (P!; isotype L!).
Distribution: New Caledonia.
764
Pleioluma laurifolia (A. Rich.) Swenson, comb. nov. ≡ Sersalisia laurifolia A. Rich. in Dumont d’Urville, Voy. Astrolabe
2: 84. 1834 ≡ Achras laurifolia (A. Rich.) F. Muell. ex
Benth., Fl. Austral. 4: 282. 1868 ≡ Sideroxylon richardii
F. Muell., Syst. Census Austral. Pl.: 92. 1882 ≡ Sideroxylon laurifolium (A. Rich.) Engl. in Bot. Jahrb. Syst. 12:
517. 1890, nom. illeg. ≡ Planchonella laurifolia (A. Rich.)
Pierre, Not. Bot.: 36. 1890 ≡ Pouteria richardii (F. Muell.)
Baehni in Candollea 9: 287. 1942≡ Beccariella laurifolia
(A. Rich.) Aubrév. in Adansonia, n.s., 2: 193. 1962 – Syn
types: Voyage de l’Astrolabe 6 (E!, P!).
Distribution: Australia.
Pleioluma ledermannii (K. Krause) Swenson, comb. nov. ≡
Sideroxylon ledermannii K. Krause in Bot. Jahrb. Syst.
58: 475. 1923 ≡ Planchonella ledermannii (K. Krause)
H.J. Lam in Nova Guinea 14: 561. 1932 ≡ Pouteria ledermannii (K. Krause) Baehni in Candollea 9: 341. 1942 –
Neotype (designated by Van Royen, 1957): NeuGuinea,
Kaiser Wilhelmsland, Sepik Region, Ledermann 6956 (K!;
isoneotype E!) (original type, Ledermann 12248, B †).
Distribution: New Guinea.
Pleioluma longipetiolata (Aubrév.) Swenson & Munzinger,
comb. nov. ≡ Beccariella longipetiolata Aubrév., Fl.
Nouv.Caléd. 1: 121. 1967 – Holotype: NouvelleCalédonie,
Prony, XI.1914, I. Franc 1885 (P!; isotype P!).
Distribution: New Caledonia.
Pleioluma lucens (P. Royen) Swenson & Munzinger, comb.
nov. ≡ Planchonella lucens P. Royen in Blumea 8: 429.
1957 ≡ Pouteria royenii Baehni in Boissiera 11: 55. 1965 ≡
Beccariella lucens (P. Royen) Aubrév. in Adansonia, n.s.,
2: 193. 1962 – Holotype: NouvelleCalédonie, 19.X.1909,
Mr. & Mrs. Le Rat 763 (P!; isotype L!).
Distribution: New Caledonia.
Pleioluma macrocarpa (P. Royen) Swenson, comb. nov. ≡
Planchonella macrocarpa P. Royen in Blumea 8: 320, 429.
1957 ≡ Pouteria pearsoniorum Jessup in Austrobaileya 6:
163. 2001 ≡ Beccariella macrocarpa (P. Royen) Swenson
& al. in Cladistics 23: 221. 2007 – Holotype: Australia,
Queensland, Cook District, Kaban, Pearson Brothers s.n.
(BRI!; isotype BRI!).
Distribution: Australia.
Pleioluma macropoda (H.J. Lam) Swenson, comb. nov. ≡
Planchonella macropoda H.J. Lam in Nova Guinea 14:
563. 1932 ≡ Pouteria macropoda (H.J. Lam) Baehni in
Candollea 9: 410. 1942 – Neotype (designated by Van
Royen, 1957): New Guinea, Alola, 6000 ft alt., 5.XII.1935,
C.E. Carr 13631 (L!; isoneotypes BM!, K!, SING!).
Distributed: Sulawesi to New Guinea.
Pleioluma moluccana (Burck) Swenson, comb. nov. ≡ Sideroxylon moluccanum Burck in Ann. Jard. Bot. Buitenzorg 5:
19. 1885 ≡ Beccariella moluccana (Burck) Pierre, Not. Bot.:
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Swenson & al. • Chrysophylloideae in Oceania and SE Asia
30. 1890 ≡ Planchonella moluccana (Burck) H.J. Lam in
Bull. Jard. Bot. Buitenzorg, ser. 3, 7: 200. 1925 ≡ Pouteria
moluccana (Burck) Baehni in Candollea 9: 327. 1942 – Holo
type: Archipel. Ind., Teysmann 7819 (BO n.v.; isotype L!).
Distribution: Widespread, from Java to New Guinea.
Pleioluma monticola (K. Krause) Swenson, comb. nov. ≡
Sideroxylon monticolum K. Krause in Bot. Jahrb. Syst. 58:
481. 1923 ≡ Planchonella monticola (K. Krause) H.J. Lam in
Nova Guinea 14: 561. 1932 ≡ Pouteria monticola (K. Krause)
H.J. Lam in Blumea 5: 337. 1943 – Neotype (designated by
Van Royen, 1957): New Guinea, Uniri River, 2200 m alt.,
18.I.1936, C.E. Carr 15189 (L!; isoneotypes BM!, SING!).
Distribution: New Guinea.
Pleioluma novocaledonica (Dubard) Swenson & Munzinger,
comb. nov. ≡ Planchonella novocaledonica Dubard in
Notul. Syst. (Paris) 2: 84. 1911 ≡ Sideroxylon novocaledonicum (Dubard) Baehni in Candollea 9: 428. 1942, nom.
illeg. ≡ Beccariella novocaledonica (Dubard) Aubrév. in
Adansonia, n.s., 2: 193. 1962 ≡ Pouteria egassia Baehni
in Boissiera 11: 60. 1965 – Holotype: NouvelleCalédonie,
Petit 130 (P!; isotypes G!, L!).
Distribution: New Caledonia.
Pleioluma papyracea (P. Royen) Swenson, comb. nov. ≡ Planchonella papyracea P. Royen in Blumea 8: 431. 1957 ≡ Beccariella papyracea (P. Royen) Aubrév. in Adansonia, n.s., 3:
335. 1963 ≡ Pouteria papyracea (P. Royen) Baehni in Bois
siera 11: 59. 1965 – Holotype: Australia, Queensland, Cook
District, ca. 30 km NE of Atherton, Krauss 102 (BRI n.v.).
Distribution: Australia.
Pleioluma queenslandica (P. Royen) Swenson, comb. nov. ≡
Planchonella queenslandica P. Royen in Blumea 8: 341,
430. 1957 ≡ Beccariella queenslandica (P. Royen) Aubrév.
in Adansonia, n.s., 3: 335. 1963 ≡ Pouteria queenslandica
(P. Royen) Jessup in Austrobaileya 6: 161. 2001 – Holo
type: Australia, Queensland, Eungella Mts, 31.III.1937,
H.H. Haines 136Q (K!).
Distribution: Australia.
Pleioluma rigidifolia (K. Krause) Swenson, comb. nov. ≡
Sideroxylon rigidifolium K. Krause in Bot. Jahrb. Syst.
58: 474. 1923 ≡ Planchonella rigidifolia (K. Krause)
H.J. Lam in Nova Guinea 14: 560. 1932 ≡ Pouteria rigidifolia (K. Krause) Baehni in Candollea 9: 319. 1942 – Lectotype (designated here): New Guinea, Torricelli Mts.,
800 m alt., 22.IX.1909, R. Schlechter 20322 (P00648143!;
isolectotypes P00648141!, P00648142!) (holotype B †).
Distribution: New Guinea.
Pleioluma rubicunda (Pierre ex Baill.) Swenson & Munzinger,
comb. nov. ≡ Lucuma rubicunda Pierre ex Baill. in Bull.
Mens. Soc. Linn. Paris 2: 883. 1890 ≡ Beccariella rubicunda (Pierre ex Baill.) Pierre, Not. Bot.: 30. 1890 ≡ Planchonella rubicunda (Pierre ex Baill.) Dubard in Ann. Mus.
Colon. Marseille, sér. 2, 9: 287. 1911 ≡ Pouteria rubicunda
(Pierre ex Baill.) Baehni in Candollea 9: 31. 1942 – Holo
type: Nouvelle Calédonie, forêt situées au sud de Canala,
vers 900 m alt., 20.XI.1869, Balansa 1825 (P00282282!;
isotypes P00282281!, P00282283!).
Distribution: New Caledonia.
Pleioluma sebertii (Pancher) Swenson & Munzinger, comb.
nov. ≡ Chrysophyllum sebertii Pancher in Sebert & Pancher,
Not. Bois Nouv. Caléd.: 194. 1874 ≡ Beccariella sebertii
(Pancher) Pierre, Not. Bot. 1: 30. 1890 ≡ Planchonella
sebertii (Pancher) Dubard in Ann. Mus. Colon. Marseille,
sér. 2, 10: 58. 1912 ≡ Pouteria sebertii (Pancher) Baehni in
Candollea 9: 297. 1942 – Type: Nouvelle Calédonie, Sébert
& Fournier 49 (P!).
Distribution: New Caledonia.
Pleioluma singuliflora (C.T. White & W.D. Francis) Swen
son, comb. nov. ≡ Sideroxylon singuliflorum C.T. White
& W.D. Francis in Proc. Roy. Soc. Queensland 37: 161.
1926 (“1925”) ≡ Pouteria singuliflora (C.T. White
& W.D. Francis) Baehni in Candollea 9: 316. 1942 ≡
Planchonella singuliflora (C.T. White & W.D. Francis)
P. Royen in Blumea 8: 345. 1957 ≡ Beccariella singuliflora
(C.T. White & W.D. Francis) Swenson & al. in Cladis
tics 23: 221. 2007 – Syntypes: Australia, Bellenden Ker,
near summit of Central Peak, North Queensland, south of
Cairns, I.1923, C.T. White s.n. (BRI!, K!).
Distribution: Australia.
Pleioluma vieillardii (Baill.) Swenson & Munzinger, comb.
nov. ≡ Sideroxylon vieillardii Baill. in Bull. Mens. Soc.
Linn. Paris 2: 886. 1890 ≡ Planchonella vieillardii (Baill.)
Dubard in Ann. Mus. Colon. Marseille, sér. 2, 10: 58. 1912
≡ Pouteria vieillardii (Baill.) Baehni in Candollea 9: 414.
1942 ≡ Beccariella vieillardii (Baill.) Swenson & al. in
Cladistics 23: 221. 2007 – Holotype: Nouvelle Calédonie,
Gatope, 1861–67, Vieillard 2889 (P!; isotypes K!, L!).
Distribution: New Caledonia.
Pleioluma wandae (Vink) Swenson, comb. nov. ≡ Pouteria wandae Vink in Blumea 47: 118. 2002 – Holotype:
Indonesia, Irian Jaya, surroundings of Ayawasi, 01°14′ S,
132°12′ E, ca. 450 m alt., 16.I.1996, W. Ave 4163 (L!; iso
types BO n.v., CANB n.v., MAN n.v.).
Distribution: New Guinea.
Pleioluma xerocarpa (F. Muell. ex Benth.) Swenson, comb.
nov. ≡ Achras xerocarpa F. Muell. ex Benth., Fl. Austral. 4:
281. 1868 ≡ Sideroxylon xerocarpum (F. Muell. ex Benth.)
Benth. & Hook f. ex F. Muell., Syst. Census Austral. Pl. 1:
91. 1882 ≡ Planchonella xerocarpa (F. Muell. ex Benth.)
H.J. Lam, Bull. Jard. Bot. Buitenzorg, ser. 3, 7: 218. 1925
≡ Sersalisia xerocarpa (F. Muell. ex Benth.) Domin, Bib
lioth. Bot. 89: 508. 1928 ≡ Pouteria xerocarpa (F. Muell.
ex Benth.) Baehni in Boissiera 11: 58. 1965 ≡ Beccariella
xerocarpa (F. Muell. ex Benth.) Aubrév. in Adansonia,
Version of Record (identical to print version).
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Swenson & al. • Chrysophylloideae in Oceania and SE Asia
n.s., 3: 335. 1963 – Syntypes: Australia, Rockingham Bay,
Dallachy s.n. (BRI!, K!, L!, MEL!, P!).
Distribution: Australia.
Pycnandra Benth. in Bentham & Hooker, Gen. Pl. 2: 658. 1876
– Type: Pycnandra benthamii Baill., Hist. Pl. 11: 299. 1891.
= Leptostylis Benth. in Bentham & Hooker, Gen. Pl. 2: 659.
1876 – Type (designated by Vink in Nova Guinea, n.s., 8:
87. 1957): Leptostylis longiflora Benth.
= Achradotypus Baill. in Bull. Mens. Soc. Linn. Paris 2: 881.
1890 – Type: Achradotypus vieillardii Baill.
= Trouettia Pierre ex Baill. in Bull. Mens. Soc. Linn. Paris
2: 945. 3 Jun 1891 – Type: Trouettia leptoclada Pierre ex
Baill. in Bull. Mens. Soc. Linn. Paris 2: 945. 1891.
= Chorioluma Baill., Hist. Pl. 11: 287. Sep–Oct 1891 – Type:
Chorioluma coriacea (Baill.) Baill. ≡ Sideroxylon coriaceum Baill. in Bull. Mens. Soc. Linn. Paris 2: 892. 1890.
= Ochrothallus Pierre ex Baill., Hist. Pl. 11: 298. Sep–Oct 1891
– Type: Ochrothallus sessilifolius (Pancher & Sebert) Pierre
ex Baill. ≡ Chrysophyllum sessilifolium Pancher & Sebert
in Sebert & Pancher, Not. Bois Nouv. Caléd.: 195. 1874.
= Sebertia Pierre ex Engl., Nat. Pflanzenfam., Nachtr. 4(1):
280. 1897 – Type: Sebertia acuminata (Baill.) Engl. ≡ Sersalisia acuminata Baill. in Bull. Mens. Soc. Linn. Paris
2: 945. 1891.
= Tropalanthe S. Moore in J. Linn Soc., Bot. 45: 354. 1921 –
Type: Tropalanthe comptonii S. Moore.
= Corbassona Aubrév., Fl. Nouv.Caléd. 1: 72. 1967 – Type: Corbassona deplanchei (Baill.) Aubrév. ≡ Chrysophyllum deplanchei Baill. in Bull. Mens. Soc. Linn. Paris 2: 899. 1891.
Diagnostic character combination. – Higher leaf venation
nonareolate; sepals glabrous inside; flowers usually born along
branches, cupshaped (rarely tubular); stamens usually inserted
in corolla tube orifice (rarely below); staminodes absent; style
simple; cotyledons planoconvex, red or rarely pinkish, radicle
included in cotyledons; endosperm absent.
Some 55 recognized species with another 10 waiting to be
described, all restricted to New Caledonia. Current classification
accepts four subgenera (Swenson & Munzinger, 2009, 2010a, b,
c), and a fifth based on Leptostylis Benth. is under revision. Most
species are restricted to either ultramafic or noneultramafic
substrates, and a few to calcareous soils, which means many
species are threatened to the point of extinction due to habitat
destruction from mining, logging and deliberately set fires.
Sersalisia R. Br., Prodr.: 529. 1810 – Type: Sersalisia sericea
(Aiton) R. Br. ≡ Sideroxylon sericeum Aiton, Hort. Kew.
1: 262. 1789.
Diagnostic character combination. – Higher leaf venation
areolate; sepals pubescent or glabrous inside; flowers axillary,
narrowly campanulate or tubular; stamens inserted in or just
below corolla tube orifice; staminodes present, entire; style
with stigmatic areas; cotyledons planoconvex, radicle included
in cotyledons; endosperm absent.
Four to six species, depending on the future status of some
undescribed taxa and the relationships of the Australian spe
cies Pouteria unmackiana (F.M. Bailey) Erlee, which is still
766
not analysed but probably belongs to this genus. One widely
distributed species, S. luzoniensis.
Sersalisia luzoniensis (Merr.) Swenson, comb. nov. ≡ Sideroxylon luzoniense Merr. in Philipp. J. Sci. 1(Suppl.): 222.
1906 ≡ Lucuma luzoniensis (Merr.) H.J. Lam in Bull. Jard.
Bot. Buitenzorg, ser. 3, 7: 227. 1925 ≡ Pouteria luzoniensis
(Merr.) Baehni in Candollea 9: 365. 1942 ≡ Fontbrunea
luzoniensis (Merr.) Aubrév. in Adansonia, n.s., 3: 333. 1963
– Lectotype (designated here): Philippines, Luzon, Prov
ince of Rizal, I.1906, Foxworthy 127 (K!; isolectotype US!)
(original type PNH †).
Distribution: Widespread, from the Philippine Islands in
the north, Borneo, Sulawesi, and New Guinea in the south.
Van-royena Aubrév. in Adansonia, n.s., 3: 329. 1963 – Type:
Van-royena castanosperma (C.T. White) Aubrév. ≡
Chrysophyllum castanospermum C.T. White. in Bot. Bull.
Dept. Agric. Queensland 21: 12. 1919.
Diagnostic character combination. – Tertiary leaf venation
parallel and reticulate, nonareolate; sepals pubescent inside;
flowers axillary, narrowly campanulate; stamens inserted near
middle (or just above) of the corolla tube; staminodes present,
entire; style with stigmatic areas; cotyledons planoconvex,
radicle included in cotyledons; endosperm absent.
A single species restricted to northeast Australia, poorly
monographed, but Van Royen (1957) provided a treatment with
an acceptable illustration.
ACKNOWLEDGMENTS
We are very grateful to Steve Wagstaff, two anonymous reviewers,
and Gail Stride for constructive comments on this manuscript. Mats
Thulin and Jens Klackenberg are thanked for fruitful discussions on
nomenclature. Wayne Takeuchi, George Weiblen, and Timothy Whit
field kindly provided fresh plant material from New Guinea, especially
Wayne, who has sent material to Stockholm for a long period of time.
Nellie Sugii, Doug Okamoto and Mary Merello are acknowledged for
additional samples of Planchonella from Hawaii, and Peter Wilkie and
Nura Abdul Karim who sent material from Singapore. Kate Armstrong
kindly made one sequence available. Material from Vanuatu was col
lected during the Santo 2006 expedition, funded, among others, by
the Stavros Niarchos Foundation, the Total Foundation and the French
Fonds Pacifique, and samples from Futuna were collected as part of an
inventory of exotic species conducted for the Wallis and Futuna envi
ronmental service. Marika Tuiwawa and the SUVA team are thanked
for their help in the field and for getting the collecting permit in Fiji.
Jacques Florence (Paris), JeanYves Meyer (Papeete), and Rava Tapu
tuarai (Papeete) helped with various issues in Tahiti. The herbaria G,
K, L, MO, NOU, P, PAP, and PNH provided access to their collections.
Alison Moore, Rogier deKok, Felix Forest, and Laszlo Csiba (K) are all
acknowledged, who made an exception by extracting DNA from a loose
fragment of a type specimen. Laurence Jessup (Brisbane) and Gerard
Thijsse (Leiden) helped with scanning images of type collections and
various issues. JeanFrançois Butaud (Tahiti), Glenn Leiper (Australia),
Hugh Nicholson (Australia), and JeanLouis Ruiz (New Caledonia) are
Version of Record (identical to print version).
TAXON 62 (4) • August 2013: 746–770
Swenson & al. • Chrysophylloideae in Oceania and SE Asia
thanked for their contribution of field images. We are indebted to Bodil
Cronholm at the molecular laboratory, Swedish Museum of Natural
History, who successfully designed primers for this study and was able
to “fishup” Krausella polyneura from a really messy aliquot! Funds to
Ulf Swenson were provided from the Swedish Research Council and
Regnells Gåvomedel (Royal Swedish Academy of Sciences).
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Appendix 1. Voucher information in the following order: Taxon name with authority, country, collector(s), collector number, (herbarium abbreviation), and
GenBank accessions (ETS, ITS, RPB2). Informal or unpublished names are given in quotation marks and sequences published here have the prefix HE or HF.
BeccAriellA “acutifolia” Swenson & Munzinger, New Caledonia, Munzinger & al. 5759 (NOU, P, S), HE860126, HE860065, HE995703. B. balansana
(Pierre) Aubrév., New Caledonia, Munzinger & al. 4196 (MO, NOU, P, S), HE860127, HE860066, HE995704. B. baueri (Montrouz.) Aubrév., New Caledonia,
Munzinger 340 (NOU, P, S), HE860128, AY552113, HE995705. B. “belepensis” Swenson & Munzinger, New Caledonia, Swenson & al. 917 (MO, NOU, P, S),
HE860129, HE860067, HE995706. B. brevipedicellata (P. Royen) Aubrév., New Caledonia, Dumontet, Poullain & Zongo 658 (NOU, P, S), HE860130, HE860068,
HE995707. B. brownlessiana (F. Muell.) Swenson, Bartish & Munzinger, Australia, Bartish & Ford 17 (BRI, S), HE860131, DQ154063, HE995708. B. crebrifolia (Baill.) Aubrév., New Caledonia, Munzinger, Lowry & Létocart 2036 (NOU, P, S), HE860132, DQ154042, HE995597. B. lasiantha (Baill.) Aubrév., New
Caledonia, Munzinger 2308 (NOU, P, S), HE860133, DQ154044, HE995598. B. longipetiolata Aubrév., New Caledonia, Munzinger & Létocart 2264 (NOU, P,
S), HE860135, DQ154045, HE995709. B. longipetiolata Aubrév., New Caledonia, Munzinger, Pillon & Butin 2911 (NOU, P, S), HE860136, HE860070, HE995710.
B. lucens (P. Royen) Aubrév., New Caledonia, Munzinger & Létocart 2310 (NOU, P, S), HE860137, HE860071, HE995711. B. macrocarpa (P. Royen) Swenson,
Bartish & Munzinger, Australia, Bartish & Ford 24 (S), HE860138, DQ154073, HE995712. B. Munzinger2581, New Caledonia, Munzinger, Labat & Butin 2581
(NOU, P, S), HE860140, HE860073, HE995713. B. novocaledonica (Dubard) Aubrév., New Caledonia, Munzinger 2304 (MO, NOU, S), HE860141, HE860074,
HE995714. B. papyracea (P. Royen) Aubrév., Australia, Bartish & Ford 20 (S), HE860142, DQ154046, HE995715. B. queenslandica (P. Royen) Aubrév., Aus
tralia, Bartish & Jessup 8 (S), HE860143, DQ154043, HE995659. B. rubicunda (Pierre ex Baill.) Pierre, New Caledonia, Swenson, McPherson & Mouly 596
(NOU, P, S), HE860144, AY552160, HE995716. B. sebertii (Pancher) Pierre, New Caledonia, Munzinger, Pignal, Dagostini & Hopkins 1975 (S), HE860145,
DQ154047, HE995717. B. singuliflora (C.T. White & W.D. Francis) Swenson, Bartish & Munzinger, Australia, Ford & Metcalfe 4547 (QRS, S), HE860146,
DQ246699, HE995718. B. Takeuchi25691, New Guinea, Takeuchi 25691 (LAE, S), HE860147, HE860075, HE995599. B. vieillardii (Baill.) Swenson, Bartish
& Munzinger, New Caledonia, Munzinger 2282 (S), HE860148, DQ154087, HE995719. B. xerocarpa (F. Muell. ex Benth.) Aubrév., Australia, Bartish & Ford
28 (BRI, S), HE860149, DQ154048, HE995683. chrySoPhylluM bakhuizenii P. Royen, New Guinea, Takeuchi 18997 (LAE, S), HE860150, HE860076,
HE995600. c. wagapense Guillaumin, New Caledonia, Munzinger, D. & I. Létocart 5634 (MO, NOU, P, S), HE860151, HE860077, HE995601. eccliNuSA
guianensis Eyma, South America, Brazil, Ducke Reserve 05–906 (K), HE860152, DQ246677, HE995661. e. ramiflora Mart., South America, Surinam, Irwing
& al. 55081 (S), HE860153, DQ246678, HE995602. KrAuSellA patentinervia (K. Krause) Erlee, New Guinea, Armstrong 317 (E, S), HE860154, HE860078,
HE995686. K. polyneura (K. Krause) H.J. Lam, Papua New Guinea, Ledermann 9054 (K), HF678108, HF912279, —. lePtoStyliS filipes Benth., New Cale
donia, Webster & Hildreth 14665 (P), EU661382, AY552135, HE995603. l. goroensis Aubrév., New Caledonia, Munzinger 2288 (NOU, P, S), EU661383, DQ154052,
HE995604. l. grandifolia Vink, New Caledonia, Munzinger & Oddi 2121 (MO, NOU, P, S), EU661384, DQ154053, HE995605. MAgodeNdroN mennyae
Vink, New Guinea, Takeuchi, Ama & Siga 16570 (S), HE860155, AY552114, HE995685. NieMeyerA antiloga (F. Muell.) T.D. Penn., Australia, Bartish &
Jessup 4 (S), HE860156, DQ154055, HE995662. N. chartacea (F.M.Bailey) C.T. White, Australia, Bartish & Jessup 5 (S), HE860157, DQ154057, HE995606.
N. Ford2429, Australia, Andrew Ford 2429 (S), EU661389, EF025089, HE995607. N. prunifera (F. Muell.) F. Muell., Australia, Jessup 5238 (S), HE860158,
DQ154058, HE995608. N. whitei (Aubrév.) L.W. Jessup, Australia, Floyd s.n. (S), EU661388, AY552137, HE995609. PichoNiA balansae (Baehni) Swenson &
Munzinger, New Caledonia, Munzinger 975 (MO, NOU, P), HE860159, AY552102, HE995720. P. balansana Pierre, New Caledonia, Veillon 7990 (P), HE860160,
AY552109, HE995610. P. daenikeri (Aubrév.) Swenson, Bartish & Munzinger, New Caledonia, Jaffré & Rigault 3038 (NOU, P), HE860161, AY552108, HE995690.
P. deplanchei (Baill.) Swenson & Munzinger, New Caledonia, Veillon 377 (NOU, P), HE860162, AY552103, HE995699. P. dubia (Guillaumin) Swenson &
Munzinger, New Caledonia, Grande Terre, Munzinger & Swenson 3065 (NOU, S), HE860163, HE860079, HE995721. P. dubia (Guillaumin) Swenson & Munz
inger, New Caledonia, Ile Art, Swenson & al. 918 (NOU, P, S), HE860164, HE860080, HE995722. P. grandiflora Swenson & Munzinger, New Caledonia, Swenson & Munzinger 930 (NOU, P, S), HE860165, HE860081, HE995611. P. lecomtei (Guillaumin) T.D. Penn., New Caledonia, Munzinger & al. 2170 (MO, NOU,
P, S), HE860166, DQ154061, HE995723. P. occidentalis (H.J. Lam) Aubrév., New Guinea, Takeuchi & Ama 22233 (LAE, S), HE860167, HE860082, HE995750.
PlANchoNellA amieuana (Guillaumin) Aubrév., New Caledonia, Dumontet, Zongo & Maituku 510 (NOU, P, S), HE860168, EF025090, HE995612. P. aneityensis (Guillaumin) H.J. Lam ex P.Royen, Vanuatu, Munzinger, Lowry & Tuiwawa 3665 (NOU, P, S, SUVA), HE860169, HE860083, HE995724. P. anteridifera
(C.T. White & W.D. Francis ex LanePoole) H.J. Lam, New Guinea, Takeuchi & Ama 17902 (LAE, S), HE860170, EF025109, HE995750. P. arnhemica (F. Muell.
ex Benth.) P. Royen, Australia, Harwood 1170 (S), HE860171, AY552107, HE995663. P. asterocarpon (P. Royen) Swenson, Bartish & Munzinger, Australia,
Bartish & Ford 25 (BRI, S), HE860172, DQ154078, HE995664. P. australis (R. Br.) Pierre, Australia, Floyd s.n. (S), HE860173, AY552148, HE995613. P. baillonii (Zahlbr.) Dubard, New Caledonia, Munzinger & Dagostini 2119 (MO, NOU, P, S), HE860174, HE860084, HE995725. P. cauliflora Munzinger & Swenson,
New Caledonia, Munzinger & al. 3495 (K, MO, NOU, NSW, P, S), HE860175, HE860085, HE995666. P. chartacea (F. Muell. ex Benth.) H.J. Lam, Australia,
Bartish & Jessup 1 (BRI, S), HE860176, HE862231, HE995665. P. clemensii (Lecomte) P. Royen, China, Hainan, How 73783 (S), HE860177, HE860086, HE995692.
P. costata (Endl.) Pierre, New Zealand (cultivated in USA), Peter W. Fritsch 1770 (CAS), —, AF396230, —. P. costata (Endl.) Pierre, New Zealand (cultivated
in Edinburgh), Robertson 9 (E), HE860178, —, HE995726. P. cotinifolia (A. DC.) Dubard, Australia, Bartish & Jessup 11 (BRI, S), HE860179, DQ154066,
HE995701. P. crassinervia Dubard, New Caledonia, Munzinger 2275 (NOU, P, S), HE860180, DQ154067, HE995667. P. cyclopensis P. Royen, New Guinea,
Polak 1365 (L), HE860181, HE860087, HE995693. P. cyclopensis P. Royen, New Guinea, Takeuchi, Ama & Gambia 25495 (LAE, S), HE860182, HE860088,
HE995751. P. dothioensis (Aubrév.) Swenson, Bartish & Munzinger, New Caledonia, Munzinger 995 (MO, NOU, P, S), HE860183, AY552138, HE995668.
P. duclitan (Blanco) Bakh.f., New Guinea, Fuentes & Fernando 37140 (L), HE860184, HE860089, —. P. eerwah (F.M. Bailey) P. Royen, Australia, Floyd s.n.
(S), EU661400, AY552147, HE995669. P. endlicheri (Montrouz.) Guillaumin, New Caledonia, Munzinger, Lowry & Létocart 2038 (NOU, P, S), HE860185,
DQ154068, HE995614. P. ericiflora Munzinger & Swenson, New Caledonia, Munzinger & al. 4197 (MO, NOU, P, S), HE860186, HE860090, HE995660.
P. euphlebia (F. Muell.) Francis, Australia, Bartish & Ford 18 (BRI, MO, S), HE860187, DQ154069, HE995727. P. firma (Miq.) Dubard, New Guinea, Takeuchi,
Ama & Jisaka 21437 (LAE, S), HE860188, HE860091, HE995615. P. glauca Swenson & Munzinger, New Caledonia, Swenson, McPherson & Mouly 625 (S),
HE860189, AY552104, HE995728. P. grayana H.St.John, French Polynesia, Australs, Butaud 192 (PAP), HE860191, HE860092, HE995616. P. grayana var.
florencei Fosberg, French Polynesia, Tahiti, Florence 3967 (BISH, PAP, S), —, HE860093, —. P. grayana H. St.John, French Polynesia, Tuamotus, Butaud &
Lagouy 1173 (PAP), HE860193, HE860094, HE995617. P. grayana H. St.John, Wallis and Futuna, Alofi, Munzinger 5479 (NOU, P, S), HE860190, HE860095,
HE995729. P. grayana H. St.John, Wallis and Futuna, Futuna, Munzinger, Meyer & Jourdan 5282 (P, S), HE860192, HE860096, HE995730. P. howeana (F. Muell.)
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TAXON 62 (4) • August 2013: 746–770
Appendix 1. Continued.
Pierre, Lord Howe Island, Le Cussan 1210 (BRI), HE860194, EF025094, HE995681. P. Ile Yande, New Caledonia, Swenson & Munzinger 715 (S), HE860195,
EF025091, —. P. kaalaensis Aubrév., New Caledonia, Swenson & Munzinger 706 (S), HE860196, HE860097, HE995694. P. koumaciensis Aubrév., New Cale
donia, Munzinger 2665 (NOU), HE860197, EF025095, HE995682. P. kuebiniensis Aubrév., New Caledonia, Munzinger & Létocart 2057 (S), HE860198, DQ154070,
HE995670. P. laetevirens (Baill.) Pierre ex Dubard, New Caledonia, Munzinger, Pignal & Lowry 2001 (MO, NOU, P, S), HE860199, DQ154071, HE995618.
P. lamprophylla (K. Krause) H.J. Lam, New Guinea, Takeuchi, Ama & Gambia 25573 (LAE, S), HE860200, HE860098, HE995619. P. lauracea (Baill.) Dubard,
New Caledonia, McPherson & Munzinger 18070 (MO, S), HE860201, AY552145, HE995731. P. ledermannii (K. Krause) H.J. Lam, New Guinea, Takeuchi,
Towati, Jisaka & Ama 17700 (LAE, S), HE860202, HE860099, HE995732. P. linggensis (Burck) Pierre, Wallis and Futuna, Futuna, Munzinger 5330 (NOU, P,
S), HE860203, HE860100, HE995733. P. linggensis (Burck) Pierre, New Guinea, Takeuchi, Jisaka, Towati & Ama 21108 (LAE, S), HE860204, HE860101,
HE995695. P. linggensis (Burck) Pierre, Vanuatu, Wheatley 271 (K), —, DQ154062, —. P. luteocostata Munzinger & Swenson, New Caledonia, Munzinger,
Jaffré & Roumagnac 2375 (NOU, S), EU661401, EF025099, HE995755. P. maingayi (C.B. Clarke) P. Royen, Singapore, Wilkie & Gwee 507 (E), HE860205,
HE860102, HE995620. P. mandjeliana Munzinger & Swenson, New Caledonia, Munzinger & Pillon 2861 (NOU, P, S), HE860206, EF025100, HE995680.
P. membranacea H.J. Lam, Fiji, Smith 4609 (S), HE860207, DQ154074, HE995621. P. mindanaensis H.J. Lam, Philippines, Clemens 1015 (PNH), HE860208,
HE860103, HE995622. P. minutiflora Munzinger & Swenson, New Caledonia, MacKee 16639 (NOU, P, S), HE860209, HE860104, HE995734. P. moluccana
(Burck) H.J. Lam, New Guinea, Takeuchi, Ama & Gambia 25550 (S), HE860139, HE860072, HE995671. P. Munzinger 6150, New Caledonia, Munzinger 6150
(NOU), HE860210, HE860105, HE995735. P. Munzinger 6490, Fiji, Munzinger 6490 (NOU, SUVA), HE860211, HE860106, HE995736. P. Munzinger 6514,
Fiji, Munzinger 6514 (NOU, SUVA), HE860212, HE860107, HE995737. P. myrsinifolia (F. Muell.) Swenson, Bartish & Munzinger, Australia, Floyd s.n. (S),
HE860213, AY552143, HE995702. P. myrsinoides (Benth.) S.T. Blake ex Francis, Australia, McDonald 3365 (QRS, S), HE860214, EF025092, HE995738.
P. obovata (R. Br.) Pierre, Taiwan, Chung & Anderberg 1166 (HAST, S), EU661402, DQ154076, HE995739. P. Pillon 150, New Caledonia, Pillon, Barrabé &
Rigault 150 (NOU, P, S), HE860215, HE860108, HE995696. P. pohlmaniana (F. Muell.) Pierre ex Dubard, Australia, Bartish & Ford 22 (BRI, S), HE860216,
DQ154079, HE995672. P. pronyensis Guillaumin, New Caledonia, Munzinger 2051 (NOU), HE860217, DQ154080, HE995623. P. roseoloba Munzinger &
Swenson, New Caledonia, Munzinger 2311 (NOU, P, S), HE860218, DQ154090, HE995673. P. rufocostata Munzinger & Swenson, New Caledonia, Munzinger,
Labat & Butin 2583 (NOU, S), HE860219, DQ154089, HE995684. P. saligna S. Moore, New Caledonia, Munzinger, Létocart & Gâteblé 2218 (NOU, P, S),
HE860220, DQ154083, HE995624. P. sandwicensis (A. Gray) Pierre, Hawaii, Ohao, Koolan 119d (GB), HE860221, DQ154084, HE995740. P. sandwicensis (A.
Gray) Pierre, Hawaii, Kauai, Merello, Bess & Johnson 3227 (MO, S), HE860222, HE860109, HE995741. P. sandwicensis (A. Gray) Pierre, Hawaii, Ohau, Doug
Okamoto s.n. (S), HE860223, HE860110, HE995742. P. skottsbergii Guillaumin, New Caledonia, Munzinger, Gâteblé & Amice 2391 (NOU, P, S), HE860224,
DQ154085, HE995674. P. smithii (P. Royen) A.C. Sm., Fiji, Munzinger 6495 (NOU, S, SUVA), HE860225, HE860111, HE995743. P. solida P. Royen, New Guinea,
Takeuchi, Towati & Ama 17286 (LAE, S), HE860226, EF025104, HE995675. P. sphaerocarpa (Baill.) Dubard, New Caledonia, Tronchet, Munzinger & Oddi 389
(MO, P), EU661403, AY552139, HE995625. P. tahitensis (Nadeaud) Pierre ex Dubard, French Polynesia, Raiatea, Meyer & Taputuarai 3013 (NOU, PAP),
HE860227, EF025108, HE995626. P. tahitensis (Nadeaud) Pierre ex Dubard, French Polynesia, Tahiti, Meyer & Taputuarai 3051 (PAP), HE860228, EF025105,
HE995744. P. thiensis Aubrév., New Caledonia, Munzinger, Dagostini, Rigault & Kurpisz 2625 (S), HE860229, EF025106, HE995676. P. thyrsoidea C.T. White,
New Guinea, Whitfeld PA-2E-0072 (LAE, MIN, S), HE860230, HE860112, HE995745. P. torricellensis (K. Schum.) H.J. Lam, Wallis and Futuna, Alofi, Munzinger 5473 (NOU, S), HE860231, HE860113, HE995746. P. torricellensis (K. Schum.) H.J. Lam, Wallis and Futuna, Futuna, Munzinger 5284 (NOU), HE860232,
HE860114, HE995747. P. umbonata (P. Royen) A.C. Sm., Fiji, Smith 8298 (K), HE860233, DQ154086, HE995627. P. vitiensis Gillespie, Fiji, Smith 7700 (S),
HE860234, DQ154088, HE995628. P. xylocarpa (C.T. White) Swenson, Bartish & Munzinger, Australia, Bartish & Ford 31 (BRI, S), HE860235, DQ154049,
HE995629. P. xylocarpa (C.T. White) Swenson, Bartish & Munzinger, Papua New Guinea, Takeuchi, Towati & Ama 17284 (LAE, S), HE860236, EF025107,
HE995677. PouteriA Armstrong316, New Guinea, Armstrong 316 (E), —, HE860115, —. P. gillisonii Vink, New Guinea, Takeuchi & Ama 19050B (LAE,
S), HE860237, HE860116, HE995748. P. luzoniensis (Merr.) Baehni, New Guinea, Conn, Damas, Fazang, Paul & Kuria 5063 (L), HE860238, HE860117,
HE995697. P. maclayana (F. Muell.) Baehni, Papua New Guinea, Regaldo & Katik 1102 (L), HE860239, HE860118, HE995698. P. malaccensis (C.B. Clarke)
Baehni, Singapore, Swenson, Karim & Fadli 1017 (S, SING), HE860240, HE860119, HE995679. P. pullenii Vink, New Guinea, Takeuchi, Jisaka, Towati & Ama
21157 (LAE, S), —, HE860120, HE995630. P. richardii (F. Muell.) Baehni, Australia, Dixon & Leach 1032 (BRI, DNA, S), HE860134, HE860069, HE995749.
P. stellibacca J.F. Maxwell, Thailand, Boonkongchart & Chongko 146 (L), HE860241, HE860121, HE995700. P. wandae Vink, New Guinea, Armstrong 305 (E,
S), HE860242, HE860122, —. PycNANdrA acuminata (Pierre ex Baill.) Swenson & Munzinger, New Caledonia, Munzinger 1006 (MO, NOU, P), EU661430,
AY552124, HE995631. P. atrofusca Swenson & Munzinger, New Caledonia, Munzinger & al. 2618 (NOU, P, S), EU661419, EU661443, HE995632. P. balansae
(Baill.) Swenson & Munzinger, New Caledonia, Munzinger & al. 1451 (S), EU661387, AY552123, HE995754. P. belepensis Swenson & Munzinger, New Cale
donia, Ile Art, Swenson, Munzinger & Barrabé 913 (S), HE860243, HE860123, HE995687. P. benthamii Baill., New Caledonia, Munzinger, Létocart & Gâteblé
2228 (NOU, P, S), EU661404, EU661436, HE995633. P. blaffartii Swenson & Munzinger, New Caledonia, Swenson, McPherson & Mouly 597 (NOU, S), EU661423,
AY552127, HE995634. P. blanchonii (Aubrév.) Swenson & Munzinger, New Caledonia, Munzinger, Labat, Leveque & Mandaoué 2576 (S), EU661390, DQ154059,
HE995635. P. bracteolata Swenson & Munzinger, New Caledonia, Munzinger, Pillon & Butin 2885 (NOU, P, S), EU661421, EU661445, HE995636. P. caeruleilatex Swenson & Munzinger, New Caledonia, Munzinger & al. 2622 (MO, NOU, P, S), EU661426, EU661448, HE995637. P. canaliculata Swenson & Munz
inger, New Caledonia, Munzinger & al. 2067 (MO, NOU, P, S), EU661431, DQ154092, HE995638. P. carinocostata Vink, New Caledonia, McPherson &
Munzinger 18091 (MO, NOU, P, S), EU661405, AY552132, HE995639. P. comptonii (S. Moore) Vink, New Caledonia, Lowry, McPherson & Le Borgne 5780A
(MO, S), EU661407, AY552131, HE995640. P. controversa (Guillaumin) Vink, New Caledonia, Lowry, McPherson & Le Borgne 5787 (MO, S), EU661408,
AY552126, HE995641. P. cylindricarpa Swenson & Munzinger, New Caledonia, Swenson, McPherson & Mouly 615 (MO, NOU, S), EU661429, AY552110,
HE995757. P. decandra (Montrouz.) Vink, New Caledonia, Ile Art, Swenson, Munzinger & Barrabé 920 (S), HE860244, HE860124, HE995688. P. deplanchei
(Baill.) Swenson & Munzinger, New Caledonia, Munzinger 978 (MO, NOU, P, S), EU661380, AY552120, HE995642. P. fastuosa (Baill.) Vink, New Caledonia,
Munzinger & Swenson 2993 (NOU, S), EU661394, EU661434, HE995643. P. francii (Guillaumin & Dubard) Swenson & Munzinger, New Caledonia, Munzinger
965 (MO, NOU, P), EU661391, AY552117, HE995644. P. glabella Swenson & Munzinger, New Caledonia, Munzinger & al. 2615 (NOU, P, S), EU661418, EU661442,
HE995645. P. glaberrima Swenson & Munzinger, New Caledonia, Munzinger & al. 1394 (MO, NOU, P, S), EU661399, AY552133, HE995646. P. gordoniifolia
(S.Moore) Swenson & Munzinger, New Caledonia, Swenson & Munzinger 726a (BRI, MO, NOU, P, S), EU661392, EU661433, HE995647. P. griseosepala Vink,
New Caledonia, Swenson, McPherson & Mouly 627 (MO, NOU, S), EU661414, AY552128, HE995648. P. kaalaensis Aubrév., New Caledonia, Munzinger &
Labat 2599 (NOU, S), EU661415, EU661440, HE995756. P. linearifolia Swenson & Munzinger, New Caledonia, Munzinger & Blaffart 2786 (NOU, P, S), EU661427,
EU661440, HE995753. P. neocaledonica (S.Moore) Vink, New Caledonia, Tronchet, Munzinger & Oddi 426 (MO, NOU, P, S), EU661416, AY552129, HE995649.
P. ouaiemensis Swenson & Munzinger, New Caledonia, Munzinger, Lowry, Blaffart & Brown 3135 (NOU, S), EU661422, EU661446, HE995689. P. paucinervia
Swenson & Munzinger, New Caledonia, Munzinger & al. 1438 (NOU, P, S), EU661424, AY552159, HE995650. P. pubiflora Swenson & Munzinger, New Cale
donia, Munzinger & al. 2624 (NOU, P, S), EU661420, EU661444, HE995651. P. sarlinii (Aubrév.) Swenson & Munzinger, New Caledonia, Munzinger 1860 (NOU,
P, S), EU661395, EU661435, HE995652. P. schmidii (Aubrév.) Swenson & Munzinger, New Caledonia, McPherson & Munzinger 18106 (MO, NOU, P, S),
EU661396, AY552116, HE995653. P. sessiliflora Swenson & Munzinger, New Caledonia, Munzinger & McPherson 696 (BRI, MO, NOU, P, S), EU661398,
AY552161, HE995691. P. sessilifolia (Pancher & Sebert) Swenson & Munzinger, New Caledonia, McPherson & Munzinger 18176 (MO, P), EU661397, AY552118,
HE995654. P. vieillardii (Baill.) Vink, New Caledonia, Dumontet, Zongo & Maituku s.n. (S), EU661417, EU661441, HE995655. P. viridiflora Swenson & Munz
inger, New Caledonia, Munzinger, McPherson & Tuiwawa 4195 (NOU, S), HE860245, HE860125, HE995656. SerSAliSiA sericea (Aiton) R. Br., Australia,
Harwood 1172 (S), HE860246, AY552112, HE995657. S. sessiliflora (C.T. White) Aubrév., Australia, Bartish & Ford 33 (BRI, MO, P, S), HE860247, DQ154094,
HE995678. VAN-royeNA castanosperma (C.T. White) Aubrév., Australia, Bartish & Ford 26 (S), HE860248, DQ154096, HE995658.
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