http://www.diva-portal.org Postprint This is the accepted version of a paper published in Taxon. This paper has been peer-reviewed but does not include the final publisher proof-corrections or journal pagination. Citation for the original published paper (version of record): Swenson, U., Nylinder, S., Munzinger, J. (2013) Towards a natural classification of Sapotaceae subfamily Chrysophylloideae in Oceania and Southeast Asia based on nuclear sequence data. Taxon, 63: 746-770 Access to the published version may require subscription. N.B. When citing this work, cite the original published paper. Permanent link to this version: http://urn.kb.se/resolve?urn=urn:nbn:se:nrm:diva-55 DiVA (Digitala Vetenskapliga Arkivet) Naturhistoriska riksmuseet http://www.diva-portal.org This is a pre-published version of an article appearing in Taxon. Citation of the original published paper is: Author(s): Swenson, U., S. Nylinder & J. Munzinger Year: 2013 Title: Towards a natural classification of Sapotaceae subfamily Chrysophylloideae in Oceania and Southeast Asia based on nuclear sequence data Journal/Book: Taxon Volume and pages: 63: 746–770. doi: Access to the published version, if not Open Access, requires subscription to the journal. Please visit: http://www.ingentaconnect.com/content/iapt/tax 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, F-34000 France Author of correspondence: Ulf Swenson, ulf.swenson@nrm.se Suggested running title: Chrysophylloideae in Oceania and SE Asia 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 majority-rule 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 monophyletic 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 as Supplementary Data of the online version of this article (http://www.ingentaconnect.com/content/iapt/tax). 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 morphological homoplasy. Various systems of classification recognizing 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 Sarcospermatoideae (Swenson & Anderberg, 2005). Chrysophylloideae are most diverse in South America and Australasia, Sapotoideae in Africa and Asia, while Sarcospermatoideae are restricted 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. All members of Chrysophylloideae in Australasia form a monophyletic group (Swenson & Anderberg, 2005; Bartish & al., 2005; Swenson & al., 2008b), and molecular dating suggests emergence in the area by the Early Eocene, some 52 million 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; Triono & al., 2007; Swenson & al., 2007a) suggest 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 prevented 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 assessed by phylogenetic analyses using nuclear (ETS, ITS) and chloroplast (trnH–psbA, trnS–G) sequence data, as well as morphology (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 opposite (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 latter with several subclades but with weak internal node support. Two morphological characters traditionally used for generic recognition 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 & Munzinger (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 and Morat, 2008; Swenson and Tehler, 2009). As currently circumscribed, Beccariella includes some 20 species and Planchonella about 60 species (Swenson & al., 2007a, 2007b). 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.; Herrmann-Erlee & Lam, 1957), and Pouteria (28 spp.; Herrmann-Erlee & 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 (Swenson & 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 flowers 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 exceptions), a fruit having up to five seeds, and a long seed scar that often is linear and covers 90–100% of the seed length. However, in contention with this, a search (accessed March 2013) in the online World Checklist at the Royal Botanical Garden, Kew (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 genus of Chrysophylloideae in Oceania and Southeast Asia. It has a circum-Pacific 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). Species of this genus exhibit intriguing traits, such as a gynomonoecious 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 occurring in the Pacific. Here we will address phylogenetic relationships 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 hypotheses on its relationships. Swenson & al. (2007a, 2007b) 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 colonized 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 presently found in continental Southeast Asia. However, the generated 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 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 relationships within Chrysophylloideae in Oceania and Southeast Asia, excluding the distantly related Asian Xantolis ?? (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 morphological 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 monophyletic genera. MATERIALS AND METHODS Nomenclature and taxon sampling. — We follow the subfamily classification of Swenson & Anderberg (2005). The checklist of Sapotaceae (Govaerts & al., 2001) follows Pennington's (1991) generic classification and includes a full list of published names, but accepted names are continuously updated 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, including 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 generi-types of two Pouteria segregates: Blabeia Baehni (represented 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 territory 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 Australasian 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; 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, 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 provide a sufficient number of informative characters needs consideration. 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 coding and noncoding cpDNA and nrDNA regions with varying success. Of the cpDNA regions used for phylogenetic estimates (atpβ–rbcL, ndhF, petN–psbM, psbM–trnD, psbB–psbH, rpl20–rps12, trnC–petN, trnH–psbA, trnLtrnF, and trnS-trnG) the average 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 silica gel dried 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, amplification, 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, 18S-ETS (Baldwin and Markos, 1998) and Sap-1 (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 and 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 identified by Swenson & al. (2007b). We decided to exclude these and reduced the clade to three species (P. glauca, P. lauracea, and 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 selected 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 poly-T-regions, followed by the same PCR protocol described above. Purified products were sequenced using an ABI3130xl Automated DNA Sequencer (Applied Biosystems, Foster City, California, USA). Alignment, gap coding, and model testing. — Alignments were performed on each separate sequenced locus in MAFFT v.6.818b (Katoh & al., 2005) using the L-INS-i predefined parameter settings. Resulting matrices were imported into MESQUITE (Maddison & Maddison, 2011) and minor manual adjustments were made to the alignments. Following the method of Simmons and 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 best-fit substitution model using jModelTest (Posada, 2008) under the Bayesian information 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 1.6.2 (part of the BEAST package) as an output file for Bayesian inference in BEAST 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 topology, 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 reviewed in Tracer v1.5 (Rambaut & Drummond, 2009). A proportion of the samples in each run were discarded as burn-in, and the posterior set of trees was summarized in TreeAnnotator 1.6.2 (Drummond & Rambaut, 2007). The resulting tree was then visualized in FigTree v1.3.1 (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 consider 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. The morphological data used here have been gathered from earlier studies (Swenson & Anderberg, 2005; Swenson & al., 2007a, 2007b, 2008a, 2008b), revisions (van Royen, 1957; Herrmann-Erlee & Lam, 1957; Herrmann-Erlee & van Royen, 1957) and herbarium material deposited at L, MO, NOU, P, PAP, and S (abbreviations follow Holmgren & al., 1990). The morphological terminology follows Harris & Harris (1997). We used MacClade 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 venation, 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 surfaces, 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 actinomorphic and partly sympetalous, comprising a tube and free corolla 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 cup-shaped, urn-shaped, narrowly campanulate, or tubular. This character is in fact a combination 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 cup-shaped 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 urn-shaped 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. 1 I–J). A campanulate flower is similar to an urn-shaped 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 different 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, and 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 Sapotaceae, which may, however, differ at the apex. It has either visible, 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). Character 6–8. — A number of fruit characters show strong congruence and can be amalgamated to three features: (i) cotyledons foliaceous, having a radicle extending below the cotyledon commissure, endosperm present; (ii) plano-convex cotyledons without a visible radicle, endosperm absent; and (iii) plano-convex 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 represented 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 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 and 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 Figure 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 subgenus 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 accessions 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 majority-rule consensus tree obtained from the BEAST analysis of nuclear sequences of Chrysophylloideae in Oceania and Southeast Asia. They show all high congruence with the generic concept proposed 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 Chrysophylloideae (Bartish & al., 2005; Swenson & al., 2007a, 2007b, 2008a; Triono & al., 2007). Exact affinities of Magodendron and Vanroyena 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 Australasian 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, members currently placed in Chrysophyllum or Pouteria need to be transferred. Leptostylis need 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 well resolved phylogeny brings to light interesting 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 Australasian-Pacific group will be addressed elsewhere. Useful morphological characters. — Morphology has repeatedly 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 number of stamens opposite each corolla lobe, and the ratio between the length of the corolla tube and the corolla lobes (Swenson & al., 2007a, 2008a, 2008b). 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 present 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 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 sepals (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 sometimes difficult according to the definition above. Nevertheless, the corolla can be termed narrowly campanulate in Beccariella, Van-royena, and some Sersalisia (Fig. 3C). A cup-shaped corolla 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 urn-shaped, where the calyx is more or less as long as, or slightly longer than, the corolla tube (Fig. 1I–J). Urn-shaped flowers never have spreading or revolute corolla lobes comparable to the cup-shaped flower. Exceptions to the urn-shaped 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 well-exposed corolla tube (Fig. 1G–H). Stamens are inserted opposite the corolla lobes in Sapotaceae, 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 stamens 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 orifice, character states that are restricted to the other genera under study. Stamen insertion in Van-royena has been reported as basal (Herrmann-Erlee & 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 identify 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 contain strong phylogenetic signal, useful in character combination for circumscribing groups, as well as in the field (Fig. 3E). The distribution of foliaceous or plano-convex cotyledons, absence or presence of an exserted radicle, and absence or presence of an endosperm, are clearly correlated among members 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 cotyledons, an exserted radicle and an endosperm, (ii) Magodendron has planoconvex cotyledons, an exserted radicle and non-endospermous seeds, and (iii) all other genera have seeds with plano-convex cotyledons, an included radicle and no endosperm. 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 homoplasy in Chrysophylloideae, the characters here overlaid in 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 generi-types. The genera were described as close relatives, Niemeyera being distinguished by its berry-like fruit and thin seed coat (testa) in contrast to Amorphospermum with its drupe-like 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, 2010c). 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 circle 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 venation 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 (Baill.) Baehni. — Beccariella is strongly supported as an independent lineage with maximum Bayesian and jackknife support. This relationship is fully congruent with earlier findings (Bartish & al., 2005; Swenson & al., 2007a), and Clade C of Triono & al. (2007). The position within the Australasian 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 20 species and another seven species (Planchonella firma (Miq.) Dubard, P. lamprophylla (K. Krause) H.J. Lam, P. ledermanii (K. Krause) H.J. Lam, P. moluccana (Burck) H.J. Lam, Pouteria gillsonii Vink, P. richardii (F. Muell.) Baehni and 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 became a later homonym and illegitimate already at publication. Subsequent classification systems like Herrmann-Erlee & 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 (Sapotaceae) against the rarely used name Beccariella Ces. (Podoscyphaceae), a proposal the Nomenclatural Committee for Vascular Plants was unable to agree upon (Brummitt, 2011b). Thus, species 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., and Pyriluma Aubrév.) in Australasia are polyphyletic or embedded in Planchonella (Swenson & al., 2007a, 2007b). Some generic names are still in synonymy with Pouteria (Pennington, 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 generi-types 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 remain 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 (Herrmann-Erlee & 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, staminodes 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 3-merous flower, staminodes, and a peculiar fruit (van 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, 1891) and later transferred to a section of Sideroxylon L. (Engler in Engler & Prantl, 1897), was based on Sideroxylon crebrifolium (Baill.) Engl. and has never been used in any classification. In fact, Baehni (1965) simultaneously 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 the International Code of Botanical Nomenclature (McNeill & al., 2006), that Pleioluma is used for this genus since it is the earliest legitimate name available. Indeed, 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 characterized by non-areolate leaf venation, sepals that are generally tomentulose on the outer surface but glabrous inside, flowers with staminodes and stamens inserted just below the tube orifice, 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 60 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 recognize 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 addition our analyses clearly recover it within the genus. Planchonella sandwicensis is distributed across all main islands of Hawaii forming the north-eastern outpost of the generic 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 separated 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, 2007b). 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 similar in leaf venation, but belongs to Sapotoideae, not Chrysophylloideae (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 mesic to wet forests (Butaud & al., 2011; Pouteau & al., 2012). All seven accessions analysed here are recovered in a single, strongly supported clade, with some support for internal resolution, and all have a unique triple-T-insertion 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 x 7–10 cm) in comparison to narrowly elliptic and small (7–11 x 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 applicable. 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 suggested to form a complex of similar species with unclear species limits (Swenson & al., 2007a, 2007b). 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 large-scale 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, 5-celled fruit with one seed that has an exserted radicle. This is fragmented information, but Dubard (1912), as well as Pennington (1991), associated the species with Planchonella (or Pouteria section Oligotheca). Despite the incomplete material, these characters are in full agreement with the character combination of Planchonella. Our careful inspection of the 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 character combination that is a mixture between Pichonia (sepals pubescent inside, stamens in the tube orifice and simple style) and Planchonella (non-areolate 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 (Herrmann-Erlee & 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 & Anderberg, 2005; Swenson & al., 2007a) and strengthen the conclusion that there are nine lineages of Chrysophylloideae in Oceania and Southeast Asia (ten with Xantolis) that warrant formal recognition. Below, we provide a generic key 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 homotypic synonyms are cited for species in order to save space. Type citations follow the text on the label, or the original description if the type material has not been viewed (n.v.). For more complete nomenclature, see Govaerts & al. (2001) and the World Checklist at the Royal Botanical Garden (http://apps.kew.org/wcsp/home.do). On going floristic work in Australia appear in Flora of Australia, but the volume of Sapotaceae is currently unpublished (L. Jessup, pers. comm.). Typifications of Australian species, if needed, will appear in this flora. Typification of Pleioluma species occurring 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 4. Tertiary or quaternary leaf venation areolate ................................................................................................ 5 4. Tertiary or quaternary leaf venation never areolate ...................................................................................... 8 5. Flowers born on trunk .............................................................................................................. Magodendron 5. Flowers axillary ............................................................................................................................................. 6 6. Stamens inserted in lower half of corolla tube; foliaceous cotyledons .......................................... Pleioluma 6. Stamens inserted in or just below corolla tube orifice; cotyledons plano-convex ........................................ 7 7. Flowers cup-shaped; style simple ..................................................................................................... Pichonia 7. Flowers narrowly campanulate or tubular; style with several stigmatic areas ................................ Sersalisia 8. Stamens inserted just below corolla tube orifice; foliaceous cotyledons .................................. Planchonella 8. Stamens inserted near middle of corolla tube; plano-convex 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. illegit. ≡ Sersalisia antiloga (F. Muell.) Domin, Biblioth. Bot. 89: 508 (1928). ≡ Chrysophyllum antilogum (F. Muell.) Vink, 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 non-areolate; sepals glabrous inside; flowers axillary, cup-shaped; stamens inserted in corolla tube orifice; staminodes absent; style simple; cotyledons plano-convex, 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 another, as yet undescribed species (L. Jessup, pers. comm.). Magodendron Vink, Nova Guinea, n.s., 8: 124 (1957). — Type: Magodendron venefici (C.T. White & W.D. Francis) Vink, Nova Guinea, n.s., 8: 125 (1957). ≡ Achradotypus venefici C.T. White & W.D. Francis, Proc. Roy. Soc. Queensland 38: 254 (1927). Diagnostic character combination. — Leaf venation areolate; sepals glabrous inside; flowers born on trunk, cup-shaped; stamens inserted in corolla tube orifice; staminodes present, irregularly incised; style with stigmatic areas; cotyledons plano-convex with a radicle extending below cotyledon commissure; endosperm absent. Two species confined to New Guinea (Vink, 1995). Niemeyera F. Muell., Fragm. 7: 114 (1870), nom. cons. — Type: Niemeyera prunifera (F. Muell.) F. Muell., ≡ Chrysophyllum pruniferum F. Muell. Diagnostic character combination. — Leaves glabrous or glabrescent below, usually translucent; tertiary leaf venation oblique, non-areolate; sepals glabrous inside; flowers axillary, cup-shaped; stamens inserted in corolla tube orifice; staminodes absent; style simple; cotyledons plano-convex, white; radicle included in cotyledons; endosperm absent. 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. Sapot.: 22 (1890). — Type: Pichonia balansana Pierre. ≡ Epiluma Baill., Hist. Pl. 11: 287 (1891). — Type: Epiluma pyriformis Baill. = Rhamnoluma Baill., Hist. Pl. 11: 287 (1891). — Type: Rhamnoluma novocaledonica (Engl.) Baill. ≡ Lucuma novocaledonica Engl. = Wokoia Baehni, Arch. Sci. 17: 78 (1964). — Type: Wokoia rhopalocarpa (P. Royen) Baehni ≡ Pouteria rhopalocarpa P. Royen = Arnanthus Baehni, Arch. Sci. 17: 78 (1964). — Type: Arnanthus balansae Baehni ≡ Chrysophyllum balansae Baill., nom. illeg. Diagnostic character combination. — Tertiary leaf venation laxly reticulate, higher venation areolate; sepals usually pubescent inside in upper part or sometimes glabrous; flowers usually axillary, cup-shaped; stamens inserted in corolla tube orifice; staminodes present, generally entire; style simple; 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, Boissiera 7: 92 (1943). ≡ Pouteria hochreutineri (H.J. Lam) H.J. Lam, Blumea 5: 337 (1943). Holotype: New Guinea, Boridi, c. 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, Nova Guinea, n.s., 10: 134 (1959). ≡ Wokoia rhopalocarpa (P. Royen) Baehni, Arch. Sci 17 (1): 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. Sapot.: 34 (1890), nom. cons. — Type: Planchonella obovata (R. Br.) Pierre ≡ Sersalisia obovata R. Br. = Iteiluma Baill., Bull. Mens. Soc. Linn. Paris 2: 892 (1890), nom. rej. — Type: Iteiluma baillonii (Zahlbr.) Baill. ≡ Lucuma baillonii Zahlbr. ≡ Poissonella Pierre, Not. Bot. Sapot.: 29 (1890). = Peuceluma Baill., Bull. Mens. Soc. Linn. Paris 2: 895 (1890), nom. rej. — Type: Peuceluma pinifolia Baill. = Beauvisagea Pierre, Not. Bot. Sapot.: 15 (1890). — Type: Beauvisagea pomifera Zippel ex Baill. = Bureavella Pierre, Not. Bot. Sapot.: 16 (1890). — Type: Bureavella maclayana (F. Muell.) Pierre. ≡ Bassia maclayana F. Muell. = Fontbrunea Pierre, Not. Bot. Sapot.: 31 (1890). — Type: Fontbrunea malaccensis (C.B. Clarke) Pierre ≡ Sideroxylon malaccense C.B. Clarke = Krausella H.J. Lam, Nova Guinea 14: 566 (1932). — Type: Krausella polyneura (K. Krause) H.J. Lam ≡ Sideroxylon polyneurum K. Krause = Albertisiella Pierre ex Aubrév., Adansonia, n.s., 4: 42 (1964). — Type: Albertisiella novoguineensis (Vink) Aubrév. ≡ Chrysophyllum novoguineense Vink = Blabeia Baehni, Arch. Sci. 17 (1): 77 (1964). — Type: Blabeia endlicheri (Montrouz.) Baehni ≡ Sapota endlicheri Montrouz. = Pyriluma Aubrév., Fl. Nouv.-Caléd. 1: 83 (1967). — Type: Pyriluma sphaerocarpa (Baill.) Aubrév. ≡ Sideroxylon sphaerocarpa Baill. in Bull. Mens. Soc. Linn. Paris 2: 891. 1890. Diagnostic character combination. — Tertiary leaf venation usually reticulate or oblique, higher venation non-areolate; sepals glabrous inside or rarely pubescent to varying degree; flowers usually axillary, urn-shaped 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 Malesia, Australia, New Caledonia, and on to the Pacific Islands, reaching French Polynesia and Hawaii. The highest diversity is in New Caledonia (c. 40 spp.), followed by New Guinea (c. 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). However, 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 accordance with Article 11.5 of the International Code of Botanical Nomenclature (McNeill & al., 2006), 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, Bull. Jard. Bot. Buitenzorg, III, 7: 217 (1925). ≡ Sideroxylon forbesii S. Moore, J. Bot. 61 (Suppl.): 30 [June] (1923). ≡ Pouteria forbesii (S. Moore) Baehni, Candollea 9: 408 (1942). ≡ Krausella forbesii (S. Moore) H.J. Lam, 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, Bot. Jahrb. Syst. 58: 474 [November] (1923). ≡ Krausella patentinervia (K. Krause) Erlee, Blumea 8: 448 (1957). — Lectotype designated by Herrmann-Erlee & Lam (1957): Kaiser-Wilhelmsland [New Guinea], Kani Geberges, 1000 m alt., 31 October 1907, R. Schlechter 16746 (P!). 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 Herrman-Erlee and 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. Distribution: New Guinea. Planchonella garcinioides (K. Krause) Swenson, comb. nov. ≡ Sideroxylon garcinioides K. Krause, Bot. Jahrb. Syst. 58: 477 (1923). ≡ Lucuma garcinioides (K. Krause) H.J. Lam, Nova Guinea 14: 568 (1932). ≡ Pouteria garcinioides (K. Krause) Baehni, Candollea 9: 338 (1942). — Lectotype designated by Vink (2002): Neu Guinea, Sepik-Gebiet, im alluvial wald am Sepik, beim Larger Malu, alt. 20-40 m, I.1913, Ledermann 10728 (L!) (holotype in B, destroyed). = Planchonella solida P. Royen, Blumea 8: 404, 433 (1957). — Holotype: New Guinea, Yalu, near Lae, 5.III.1950, D. Fryar 3344 (SING!), isotype (L!, LAE n.v.). Note. — Herrmann-Erlee and van Royen (1957), in their revision 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. Simultaneously, 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. Distribution: New Guinea. Planchonella lamii P. Royen, Blumea 8: 398, 432 (1957). — Holotype: Amboina, near Sirimau, 450 m alt., 11.II.1930, NIFS bb 14283 (L!). = Chrysophyllum bakhuizenii P. Royen, 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 material of this species and Planchonella lamii shows that they are conspecific and should be united. Planchonella maclayana (F. Muell.) Swenson, comb. nov. ≡ Bassia maclayana F. Muell., Vict. Chem. & Druggist 7 (April): 93 (1885). ≡ Illipe maclayana (F. Muell.) F. Muell., Descr. Notes Papuan Pl. 2 (6): 12 (1885). ≡ Bureavella maclayana (F. Muell.) Pierre, Not. Bot. Sapot.: 16 (1890). ≡ Lucuma maclayana (F. Muell.) H.J. Lam, Bull. Jard. Bot. Buitenzorg, III, 7: 22 (1925). ≡ Pouteria maclayana (F. Muell.) Baehni, Candollea 9: 307 (1942). — Type: New Guinea, Miklouho-Maclay (MEL!, P!). Distribution: Widespread in Malesia, Solomon Islands. Planchonella macrantha (Merr.) Swenson, comb. nov. ≡ Sideroxylon macranthum Merr., Publ. Bur. Sci. Gov. Lab. 35: 56 (1905 publ. 1906). ≡ Lucuma macrantha (Merr.) H.J. Lam, Bull. Jard. Bot. Buitenzorg, III, 7: 225 (1925). ≡ Pouteria macrantha (Merr.) Baehni, Candollea 9: 328 (1942). ≡ Bureavella macrantha (Merr.) Aubrév., 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, destroyed), isotypes (BO!, K!, SING!, US!). Distribution: Philippines to Maluku. Planchonella malaccensis (C.B. Clarke) Swenson, comb. nov. ≡ Sideroxylon malaccense C.B. Clarke in J.D. Hooker, Fl. Brit. India 3: 537 (1882). ≡ Fontbrunea malaccensis (C.B. Clarke) Pierre, Not. Bot. Sapot.: 31 (1890). ≡ Lucuma malaccensis (C.B. Clarke) Dubard, Ann. Mus. Colon. Marseille, sér. 2, 10: 19 (1912). ≡ Pouteria malaccensis (C.B. Clarke) Baehni, Candollea 9: 302 (1942). ≡ Xantolis malaccensis (C.B. Clarke) Baehni, 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, 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°29S, 142°41’E, 19.VII.1990, W. Takeuchi 6276 (L!), isotype (L!). Distribution: New Guinea. Planchonella orkor (Vink) Swenson, comb. nov. ≡ Pouteria orkor Vink, Blumea 47: 134 (2002). — Holotype: New Guinea, Saidor Subdistrict, Naho-Rawa, 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, Blumea 8: 503 (1957). — Holotype: Indonesia, Sumatra, Riouw District, Indragiri, Moeata 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, Bot. Jahrb. Syst. 58: 475 (1923). ≡ Krausella polyneura (K. Krause) H.J. Lam, Nova Guinea 14: 567 (1932). ≡ Pouteria multinervis T.D. Penn., Gen. Sapotac.: 202 (1991). — Lectotype selected by Herrmann-Erlee & Lam (1957): Deutsch-Neuguinea, Sepikgebiet, Ledermann 9054 (K!), isolectotype (SING!). Distribution: New Guinea. Planchonella pullenii (Vink) Swenson, comb. nov. ≡ Pouteria pullenii Vink, 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, Blumea 47: 136 (2002). — Holotype: Philippines, Mindoro, Mt. Halcon area near Paitan, Dulangan 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, 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., Philipp. J. Sci., C 10: 59 (1915). ≡ Pouteria villamilii (Merr.) Baehni, Candollea 9: 318 (1942). ≡ Bureavella villamilii (Merr.) Aubrév., Adansonia, n.s., 3: 331 (1963). — Holotype: Philippines, Luzon, Province 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, 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, Boissiera 11: 150 (1965). ≡ Sersalisia section Pleioluma Baill., Hist. Pl. 11: 280 (Sept.-Oct. 1891). — Type: Pleioluma crebrifolia (Baill.) Swenson & Munzinger ≡ Lucuma crebrifolia Baill. = Beccariella Pierre, Not. Bot. Sapot.: 30 (1890), nom. illeg., non Beccariella Ces. in Atti. Reale Accad. Sci. Fis. 8: 9 (1879). — Type: Beccariella sebertii (Pancher) Pierre ≡ Chrysophyllum sebertii Pancher Diagnostic character combination. — Leaf venation areolate; 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 several 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, Blumea 8: 308, 428 (1957). ≡ Beccariella azou (P. Royen) Aubrév., 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., Bull. Mens. Soc. Linn. Paris 2: 889 (1890). ≡ Planchonella balansana (Pierre ex Baill.) Pierre ex Dubard, Ann. Mus. Colon. Marseille, sér. 2, 10: 46 (1912). ≡ Pouteria balansana (Pierre ex Baill.) Baehni, Candollea 9: 317 (1942). ≡ Beccariella balansana (Pierre ex Baill.) Aubrév., 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., Mem. Acad. Roy. Sci. Lyon, Sect. Sci. 10: 229 (1860). ≡ Planchonella baueri (Montrouz.) Dubard, Ann. Mus. Colon. Marseille, sér. 2, 10: 53 (1912). ≡ Pouteria baueri (Montrouz.) Baehni, Candollea 9: 329 (1942). Beccariella baueri (Montrouz.) Aubrév., Adansonia, n.s., 2: 193 (1962). — Holotype: Nouvelle-Calédonie, 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., Fragm. 8: 111 (1873). ≡ Sersalisia brownlessiana (F. Muell.) Domin, Biblioth. Bot. 89: 508 (1928). ≡ Pouteria brownlessiana (F. Muell.) Baehni, Candollea 9: 318 (1942). ≡ Planchonella brownlessiana (F. Muell.) P. Royen, Blumea 8: 343 (1957). ≡ Beccariella brownlessiana (F. Muell.) Swenson, Bartish & Munzinger, 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., Bull. Mens. Soc. Linn. Paris 2: 897 (1891). ≡ Sideroxylon crebrifolium (Baill.) Engl. in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam., Nachtr. 1: 277 (1897). ≡ Planchonella crebrifolia (Baill.) Pierre ex Dubard, Ann. Mus. Colon. Marseille, sér. 2, 10: 53 (1912). ≡ Beccariella crebrifolia (Baill.) Aubrév., Adansonia, n.s., 2: 193 (1962). ≡ Pouteria crebrifolia (Baill.) Baehni, Boissiera 11: 59 (1965). — Lectotype designated here: Nouvelle Calédonie, 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 the Paris Herbarium (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)". However, there are four specimens of this collection in Paris, none bearing any sign of being seen by Aubréville. Hence, and in accordance with Article 9.2 of the International Code of Botanical Nomenclature (McNeill & al., 2006), one well-preserved and fertile specimen of Balansa 3154 is here designated as lectotype. Pleioluma densinervia (K. Krause) Swenson, comb. nov. ≡ Sideroxylon densinervium K. Krause, Bot. Jahrb. Syst. 58: 476 (1923). ≡ Planchonella densinervia (K. Krause) H.J. Lam, Nova Guinea 14: 562 (1932). ≡ Pouteria densinervia (K. Krause) Baehni, Candollea 9: 342 (1942). — Lectotype selected 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, Blumea 8: 352, 431 (1957). ≡ Pouteria dies-reginae (P. Royen) Vink, Blumea 47: 103 (2002). — Holotype: New Guinea, 4 km SW of Bernhard 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, Ann. Jard. Bot. Buitenzorg 5: 17 (1885). ≡ Beccariella firma (Miq.) Pierre, Not. Bot.: 30 (1890). ≡ Planchonella firma (Miq.) Dubard, Ann. Mus. Colon. Marseille, sér. 2, 10: 59 (1912). ≡ Pouteria firma (Miq.) Baehni, Candollea 9: 284 (1942). — Type: Teysmann s.n. (BO n.v.). Distribution: Widespread, from Thailand via Malesia to Solomon Islands. Pleioluma foxworthyi (Elmer) Swenson, comb. nov. ≡ Sideroxylon foxworthyi Elmer, Leafl. Philipp. Bot. 5: 1836 (1913). ≡ Planchonella foxworthyi (Elmer) H.J. Lam, Proc. Pacific Sci. Congr. 6 (4): 678 (1940). — Type: Philippine Islands, Island of Palawan, Puerto Princesa, Mt. Pulgar, III.1911, Elmer 12824 (E!, FI n.v., G!, L!, NSW n.v., P!, PNH [destroyed], US!, Z!). Distribution: Philippines. Pleioluma gillisonii (Vink) Swenson, comb. nov. ≡ Pouteria gillisonii Vink, Blumea 47: 104 (2002). — Holotype: Papua New Guinea, Morobe Province, Morobe Subprovince, 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, Bot. Jahrb. Syst. 58: 474 (1923), nom. illeg., non Hillebr., Fl. Hawaiian Isl.: 277 (1888). ≡ Planchonella krausei H.J. Lam, Nova Guinea 14: 561 (1932). ≡ Pouteria krausei (H.J. Lam) Baehni, Candollea 9: 322 (1942). — Lectotype designated by van Royen (1957): Nieuw Guinea, Kaiser Wilhelmsland, Gomadjiji, c. 450 m alt., 23.VIII.1909, R. Schlechter 19915 (L!), isolectotype (BR!, P!). Distribution: New Guinea. Pleioluma lamprophylla (K. Krause) Swenson, comb. nov. ≡ Sideroxylon lamprophyllum K. Krause, Bot. Jahrb. Syst. 58: 481 (1923). ≡ Planchonella lamprophylla (K. Krause) H.J. Lam, Nova Guinea 14: 564 (1932). ≡ Pouteria lamprophylla (K. Krause) Baehni, Candollea 9: 333 (1942). — Lectotype designated by van Royen (1957): Neu-Guinea, Sepik-Gebeit, 1912-1913, Ledermann 10318 (L!), isolectotype (K!). Distribution: New Guinea. Pleioluma lanatifolia (P. Royen) Swenson, comb. nov. ≡ Planchonella lanatifolia P. Royen, Blumea 8: 306, 428 (1957). ≡ Pouteria lanatifolia (P. Royen) Vink, 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., Bull. Mens. Soc. Linn. Paris 2: 887 (1890). ≡ Planchonella lasiantha (Baill.) Dubard, Ann. Mus. Colon. Marseille, sér. 2, 10: 58 (1912). ≡ Pouteria lasiantha (Baill.) Baehni, Candollea 9: 324 (1942). ≡ Beccariella lasiantha (Baill.) Aubrév., Fl. Nouv.-Caléd. 1: 114 (1967). — Holotype: Nouvelle-Calédonie, Mont Mi, 25.III.1869, Balansa 1322 (P!), isotype (L!). Distribution: New Caledonia. Pleioluma laurifolia (A. Rich.) Swenson, comb. nov. ≡ Sersalisia laurifolia A. Rich. in J.S.C. Dumont d’Urville, Voy. Astrolabe 2: 84 (1834). ≡ Achras laurifolia (A. Rich.) F. Muell ex Benth., Fl. Austral. 4: 282 (1868). ≡ Sideroxylon laurifolium (A. Rich.) Engl., Bot. Jahrb. Syst. 12: 517 (1890), nom. illeg. ≡ Planchonella laurifolia (A. Rich.) Pierre, Not. Bot.: 36 (1890). ≡ Beccariella laurifolia (A. Rich.) Aubrév., Adansonia, n.s., 2: 193 (1962). ≡ Sideroxylon richardii F. Muell., Syst. Census Austral. Pl.: 92 (1882). ≡ Pouteria richardii (F. Muell.) Baehni, Candollea 9: 287 (1942). — Type: Voyage de l’Astrolabe 6 (E!, P!). Distribution: Australia. Pleioluma ledermannii (K. Krause) Swenson, comb. nov. ≡ Sideroxylon ledermannii K. Krause, Bot. Jahrb. Syst. 58: 475 (1923). ≡ Planchonella ledermannii (K. Krause) H.J. Lam, Nova Guinea 14: 561 (1932). ≡ Pouteria ledermannii (K. Krause) Baehni, Candollea 9: 341 (1942). — Neotype: designated by van Royen (1957) (original type, Lederman 12248 in B, destroyed): Neu-Guinea, Kaiser Wilhelmsland, Sepik Region, Ledermann 6956 (K!), isotype (E!). Distribution: New Guinea. Pleioluma longipetiolata (Aubrév.) Swenson & Munzinger, comb. nov. — Beccariella longipetiolata Aubrév., Fl. Nouv.-Caléd. 1: 121 (1967). — Holotype: Nouvelle-Calédonia, Prony, XI.1914, I. Franc 1885 (P!), isotype (P!). Distribution: New Caledonia. Pleioluma lucens (P. Royen) Swenson & Munzinger, comb. nov. ≡ Planchonella lucens P. Royen, Blumea 8: 429 (1957). ≡ Pouteria royenii Baehni, Boissiera 11: 55 (1965). ≡ Beccariella lucens (P. Royen) Aubrév., Adansonia, n.s., 2: 193 (1962). Holotype: Nouvella-Calédonie, 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, Blumea 8: 320, 429 (1957). ≡ Pouteria pearsoniorum Jessup, Austrobaileya 6: 163 (2001). ≡ Beccariella macrocarpa (P. Royen) Swenson, Bartish & Munzinger, 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, Nova Guinea 14: 563 (1932). ≡ Pouteria macropoda (H.J. Lam) Baehni, Candollea 9: 410 (1942). Neotype designated by van Royen (1957): New Guinea, Alola, 6000 ft alt., 5.XII.1935, C. E. Carr 13631 (L!), isotypes (BM!, K!, SING!). Distributed: Sulawesi to New Guinea. Pleioluma moluccana (Burck) Swenson, comb. nov. ≡ Sideroxylon moluccanum Burck, Ann. Jard. Bot. Buitenzorg 5: 19 (1885). ≡ Beccariella moluccana (Burck) Pierre, Not. Bot. Sapot.: 30 (1890). ≡ Planchonella moluccana (Burck) H.J. Lam, Bull. Jard. Bot. Buitenzorg, III, 7: 200 (1925). ≡ Pouteria moluccana (Burck) Baehni, Candollea 9: 327 (1942). — Lectotype: Archipel. Ind., Teysmann 7819 (BO), isolectotype (L). Distribution: Widespread, from Java to New Guinea. Pleioluma monticola (K. Krause) Swenson, comb. nov. ≡ Sideroxylon monticolum K. Krause, Bot. Jahrb. Syst. 58: 481 (1923). ≡ Planchonella monticola (K. Krause) H.J. Lam, Nova Guinea 14: 561 (1932). ≡ Pouteria monticola (K. Krause) H .J. Lam, 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!), isotypes (BM!, SING!). Distribution: New Guinea. Pleioluma novocaledonica (Dubard) Swenson & Munzinger, comb. nov. ≡ Planchonella novocaledonica Dubard, Notul. Syst. (Paris) 2: 84 (1911). ≡ Sideroxylon novocaledonicum (Dubard) Baehni, Candollea 9: 428 (1942), nom. illeg. ≡ Beccariella novocaledonica (Dubard) Aubrév., Adansonia, n.s., 2: 193 (1962). ≡ Pouteria egassia Baehni, Boissiera 11: 60 (1965). — Holotype: Nouvelle-Calédonie, Petit 130 (P!), isotypes (G!, L!). Distribution: New Caledonia. Pleioluma papyracea (P. Royen) Swenson, comb. nov. ≡ Planchonella papyracea P. Royen, Blumea 8: 431 (1957). ≡ Beccariella papyracea (P. Royen) Aubrév., Adansonia, n.s., 3: 335 (1963). ≡ Pouteria papyracea (P. Royen) Baehni, Boissiera 11: 59 (1965). — Holotype: Australia, Queensland, Cook District, c. 30 km NE of Atherton, Krauss 102 (BRI n.v.). Distribution: Australia. Pleioluma queenslandica (P. Royen) Swenson, comb. nov. ≡ Planchonella queenslandica P. Royen, Blumea 8: 341, 430 (1957). ≡ Beccariella queenslandica (P. Royen) Aubrév., Adansonia, n.s., 3: 335 (1963). ≡ Pouteria queenslandica (P. Royen) Jessup, Austrobaileya 6: 161 (2001). — Holotype: 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, Bot. Jahrb. Syst. 58: 474 (1923). ≡ Planchonella rigidifolia (K. Krause) H.J. Lam, Nova Guinea 14: 560 (1932). ≡ Pouteria rigidifolia (K. Krause) Baehni, Candollea 9: 319 (1942). — Lectotype designated here: New Guinea, Torricelli Mts., 800 m alt., 22.IX.1909, R. Schlechter 20322 (P00648143), isolectotype (P00648141) (holotype in B destroyed). Distribution: New Guinea. Pleioluma rubicunda (Pierre ex Baill.) Swenson & Munzinger, comb. nov. ≡ Lucuma rubicunda Pierre ex Baill., Bull. Mens. Soc. Linn. Paris 2: 883 (1890). ≡ Beccariella rubicunda (Pierre ex Baill.) Pierre, Not. Bot. Sapot.: 30 (1890). ≡ Planchonella rubicunda (Pierre ex Baill.) Dubard in Guillaumin, Ann. Mus. Colon. Marseille, sér. 2, 9: 287 (1911). ≡ Pouteria rubicunda (Pierre ex Baill.) Baehni, Candollea 9: 31 (1942). — Holotype: 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 J. A. I. Pancher & H. Sebert, Not. Bois Nouv. Caléd.: 194 (1874). ≡ Beccariella sebertii (Pancher) Pierre, Not. Bot. 1: 30 (1890). ≡ Planchonella sebertii (Pancher) Dubard, Ann. Mus. Colon. Marseille, sér. 2, 10: 58 (1912). ≡ Pouteria sebertii (Pancher) Baehni, Candollea 9: 297 (1942). — Type: Nouvelle Calédonie, Sébert & Fournier 49 (P!). Distribution: New Caledonia. Pleioluma singuliflora (C.T. White & W.D. Francis) Swenson, comb. nov. ≡ Sideroxylon singuliflorum C.T. White & W.D. Francis, Proc. Roy. Soc. Queensland 37: 161 (1926[1925]). ≡ Pouteria singuliflora (C.T. White & W.D. Francis) Baehni, Candollea 9: 316 (1942). ≡ Planchonella singuliflora (C.T. White & W.D. Francis) P. Royen, Blumea 8: 345 (1957). ≡ Beccariella singuliflora (C.T. White & W.D. Francis) Swenson, Bartish & Munzinger, Cladistics 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., Bull. Mens. Soc. Linn. Paris 2: 886 (1890). ≡ Planchonella vieillardii (Baill.) Dubard, Ann. Mus. Colon. Marseille, sér. 2, 10: 58 (1912). ≡ Pouteria vieillardii (Baill.) Baehni, Candollea 9: 414 (1942). ≡ Beccariella vieillardii (Baill.) Swenson, Bartish & Munzinger, 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, Blumea 47: 118 (2002). — Holotype: Indonesia, Irian Jaya, sourroundings of Ayawasi, 01°14’S, 132°12E, c. 450 m alt., 16.I.1996, W. Ave 4163 (L!), isotypes (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 xylocarpa (F. Muell. ex Benth.) H.J. Lam, Bull. Jard. Bot. Buitenzorg, III, 7: 218 (1925). ≡ Sersalisia xerocarpa (F. Muell. ex Benth.) Domin, Biblioth. Bot. 89: 508 (1928). ≡ Pouteria xerocarpa (F. Muell. ex Benth.) Baehni, Boissiera 11: 58 (1965). ≡ Beccariella xerocarpa (F. Muell. ex Benth.) Aubrév., Adansonia, n.s., 3: 335 (1963). — Syntypes: Australia, Rockingham Bay, Dallachy s.n. (BRI!, K!, L!, MEL!, P!). Distribution: Australia. Pycnandra Benth., in G. Bentham and J.D. Hooker, Gen. Pl. 2: 658 (1876). — Type: Pycnandra benthamii Baill. = Leptostylis Benth., in G. Bentham and J.D. Hooker, Gen. Pl. 2: 659 (1876). — Lectotype: Leptostylis longiflora Benth., designated by Vink in Nova Guinea, n.s., 8: 87 (1957). = Achradotypus Baill., Bull. Mens. Soc. Linn. Paris 2: 881 (1890). — Type: Achradotypus vieillardii Baill. = Trouettia Pierre ex Baill., Bull. Mens. Soc. Linn. Paris 2: 945 [3 Jun.] (1891). — Type: Trouettia leptoclada Pierre ex Baill. = Chorioluma Baill., Hist. Pl. 11: 287 [Sep.-Oct.] (1891). — Type: Chorioluma coriacea (Baill.) Baill. ≡ Sideroxylon coriaceum Baill. = Ochrothallus Pierre ex Baill., Hist. Pl. 11: 298 [Sep.-Oct.] (1891). — Type: Ochrothallus sessilifolius (Pancher & Sebert) Pierre ex Baill. ≡ Chrysophyllum sessilifolium Pancher & Sebert = Sebertia Pierre ex Engl., Nat. Pflanzenfam., Nachtr. 4 (1): 280 (1897). — Type: Sebertia acuminata (Baill.) Engl. ≡ Sersalisia acuminata Baill. = Tropalanthe S. Moore, 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. Diagnostic character combination. — Higher leaf venation non-areolate; sepals glabrous inside; flowers usually born along branches, cup-shaped (rarely tubular); stamens usually inserted in corolla tube orifice (rarely below); staminodes absent; style simple; cotyledons plano-convex, red or rarely pinkish, radicle included in cotyledons; endosperm absent. Some 55 recognized species with another ten waiting to be described, all restricted to New Caledonia. Current classification accepts four subgenera (Swenson & Munzinger, 2009, 2010a, 2010b, 2010c), and Pycnandra subgenus Leptostylis 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. Fl. Nov. Holl.: 529 (1810). — Type Sersalisia sericea (Aiton) R. Br. ≡ Sideroxylon sericeum Aiton 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 plano-convex, 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 species Pouteria unmackiana (F.M. Bailey) Erlee, which is still not analysed but probably belongs to this genus. One widely distributed species, S. luzoniensis. Sersalisia luzoniensis (Merr.) Swenson, comb. nov. ≡ Sideroxylon luzoniense Merr., Philipp. J. Sci. 1 (Suppl.): 222 (1906). ≡ Lucuma luzoniensis (Merr.) H.J. Lam, Bull. Jard. Bot. Buitenzorg, III, 7: 227 (1925). ≡ Pouteria luzoniensis (Merr.) Baehni, Candollea 9: 365 (1942). ≡ Fontbrunea luzoniensis (Merr.) Aubrév., Adansonia, n.s., 3: 333 (1963). — Original type material in PNH is destroyed. Lectotype designated here: Philippines, Luzon, Province of Rizal, I.1906, Foxworthy 127 (K!), isolectotype (US!). Distribution: Widespread, from the Philippine Islands in the north, Borneo, Sulawesi, and New Guinea in the south. Van-royena Aubrév., Adansonia, n.s., 3: 329 (1963). — Type: Van-royena castanosperma (C.T. White) Aubrév. ≡ Chrysophyllum castanospermum C.T. White. Diagnostic character combination. — Tertiary leaf venation parallel and reticulate, non-areolate; 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 plano-convex, 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 Whitfield 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 collected 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 environmental 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), Jean-Yves Meyer (Papeete), and Rava Taputuarai (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. Jean-François Butaud (Tahiti), Glenn Leiper (Australia), Hugh Nicholson (Australia), and Jean-Louis Ruiz (New Caledonia) are 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 "fish-up" 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). LITERATURE CITED Álvarez, I. & Wendel, J.F. 2003. Ribosomal ITS sequences and plant phylogenetic inference. Mol. Phylogen. Evol. 29: 417–434. Anderberg, A.A. & Swenson, U. 2003. 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Bishop Museum, Honolulu. Yang, Z. & Rannala, B. 1997. Bayesian phylogenetic inference using DNA sequences: a Markov Chain Monte Carlo method. Mol. Biol. Evol. 14: 717–724. Figure legends 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 spec. nov. Photos: A, C, F by Hugh Nicholson (Australia); B by Glenn Leiper (Australia); D, I by Ulf Swenson (Sweden); E by Jean-Louis Ruiz (New Caledonia); G, H, K, L by Jérôme Munzinger (France); J by Jean-François Butaud (Tahiti). Fig. 2. Maximum clade credibility tree obtained from the BEAST analysis of nuclear sequences of Chrysophylloideae (Sapotaceae) from Oceania and Southeast Asia. Currently accepted names are colour-coded according to the legend. Posterior probabilities (PP; above) and parsimony jackknifing (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. 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. 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. Type species of genera are indicated in bold. Undescribed species and those not transferred here appear in citation marks. Table 1. List of primers used DNA region Primer Sequence 5'–3' Reference ETS 18S-ETS ACT TAC ACA TGC ATG GCT TAA TCT Baldwin & Markos (1998) ETS Sap-1 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 270F-Sap CCT AGT GTT ACC TTT TAC CCT GAT TG This study* RPB2 570F-Sap CAT GGC ATT CAA CTA CTG AAG AGT TG This study* RPB2 540R-Sap CAA CTC TTC AGT AGT TGA ATG CCA TG This study* RPB2 435F-Sap TAA TAT GTC AGC TTG TGG TGG AGA G This study* RPB2 435R-Sap CTC TCC ACC ACA AGC TGA CAT ATT A This study* This study* RPB2 800F-Sap GCA TCT CTG TCC TCT TGA TTA CTT GAA TG RPB2 800R-Sap CAT TCA AGT AAT CAA GAG GAC AGA GAT GC This study* RPB2 1020F-Sap GCT ACT TTG AAG CAC TTG CTA GAA TCT This study* RPB2 1020R-Sap AGA TTC TAG CAA GTG CTT CAA AGT AGC This study* *Primers designed by Bodil Cronholm, Swedish Museum of Natural History. Table 2. Characteristics of nuclear sequences in each of the data partitions (excluding the outgroup) Data Number of characters Aligned Constant Uninformative Informative Gaps ETS 403 114 91 198 (49.1%) 22 ITS 914 418 176 320 (35.0%) 42 RPB2 1169 483 535 151 (12.9%) 13 Total 2486 1015 802 668 (26.9%) 77 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. Beccariella balansana (Pierre) Aubrév., New Caledonia, Munzinger & al. 4196 (MO, NOU, P, S), HE860127, HE860066, HE995704. Beccariella baueri (Montrouz.) Aubrév., New Caledonia, Munzinger 340 (NOU, P, S), HE860128, AY552113, HE995705. Beccariella "belepensis" Swenson & Munzinger, New Caledonia, Swenson & al. 917 (MO, NOU, P, S), HE860129, HE860067, HE995706. Beccariella brevipedicellata (P. Royen) Aubrév., New Caledonia, Dumontet, Poullain & Zongo 658 (NOU, P, S), HE860130, HE860068, HE995707. Beccariella brownlessiana (F. Muell.) Swenson, Bartish & Munzinger, Australia, Bartish & Ford 17 (BRI, S), HE860131, DQ154063, HE995708. Beccariella crebrifolia (Baill.) Aubrév., New Caledonia, Munzinger, Lowry & Létocart 2036 (NOU, P, S), HE860132, DQ154042, HE995597. Beccariella lasiantha (Baill.) Aubrév., New Caledonia, Munzinger 2308 (NOU, P, S), HE860133, DQ154044, HE995598. Beccariella longipetiolata Aubrév., New Caledonia, Munzinger & Létocart 2264 (NOU, P, S), HE860135, DQ154045, HE995709. Beccariella longipetiolata Aubrév., New Caledonia, Munzinger, Pillon & Butin 2911 (NOU, P, S), HE860136, HE860070, HE995710. Beccariella lucens (P. Royen) Aubrév., New Caledonia, Munzinger & Létocart 2310 (NOU, P, S), HE860137, HE860071, HE995711. Beccariella macrocarpa (P. Royen) Swenson, Bartish & Munzinger, Australia, Bartish & Ford 24 (S), HE860138, DQ154073, HE995712. Beccariella "Munzinger2581", New Caledonia, Munzinger, Labat & Butin 2581 (NOU, P, S), HE860140, HE860073, HE995713. Beccariella novocaledonica (Dubard) Aubrév., New Caledonia, Munzinger 2304 (MO, NOU, S), HE860141, HE860074, HE995714. Beccariella papyracea (P. Royen) Aubrév., Australia, Bartish & Ford 20 (S), HE860142, DQ154046, HE995715. Beccariella queenslandica (P. Royen) Aubrév., Australia, Bartish & Jessup 8 (S), HE860143, DQ154043, HE995659. Beccariella rubicunda (Pierre ex Baill.) Pierre, New Caledonia, Swenson, McPherson & Mouly 596 (NOU, P, S), HE860144, AY552160, HE995716. Beccariella sebertii (Pancher) Pierre, New Caledonia, Munzinger, Pignal, Dagostini & Hopkins 1975 (S), HE860145, DQ154047, HE995717. Beccariella singuliflora (C.T. White & W.D. Francis) Swenson, Bartish & Munzinger, Australia, Ford & Metcalfe 4547 (QRS, S), HE860146, DQ246699, HE995718. Beccariella "Takeuchi25691", New Guinea, Takeuchi 25691 (LAE, S), HE860147, HE860075, HE995599. Beccariella vieillardii (Baill.) Swenson, Bartish & Munzinger, New Caledonia, Munzinger 2282 (S), HE860148, DQ154087, HE995719. Beccariella 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. Chrysophyllum 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. Ecclinusa 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. Krausella polyneura (K. Krause) H.J. Lam, Papua New Guinea, Ledermann 9054 (K), HF678108, HF912279, —. Leptostylis filipes Benth., New Caledonia, Webster & Hildreth 14665 (P), EU661382, AY552135, HE995603. Leptostylis goroensis Aubrév., New Caledonia, Munzinger 2288 (NOU, P, S), EU661383, DQ154052, HE995604. Leptostylis 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. Niemeyera chartacea (F.M.Bailey) C.T. White, Australia, Bartish & Jessup 5 (S), HE860157, DQ154057, HE995606. Niemeyera "Ford2429", Australia, Andrew Ford 2429 (S), EU661389, EF025089, HE995607. Niemeyera prunifera (F. Muell.) F. Muell., Australia, Jessup 5238 (S), HE860158, DQ154058, HE995608. Niemeyera 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. Pichonia balansana Pierre, New Caledonia, Veillon 7990 (P), HE860160, AY552109, HE995610. Pichonia daenikeri (Aubrév.) Swenson, Bartish & Munzinger, New Caledonia, Jaffré & Rigault 3038 (NOU, P), HE860161, AY552108, HE995690. Pichonia deplanchei (Baill.) Swenson & Munzinger, New Caledonia, Veillon 377 (NOU, P), HE860162, AY552103, HE995699. Pichonia dubia (Guillaumin) Swenson & Munzinger, New Caledonia, Grande Terre, Munzinger & Swenson 3065 (NOU, S), HE860163, HE860079, HE995721. Pichonia dubia (Guillaumin) Swenson & Munzinger, New Caledonia, Ile Art, Swenson & al. 918 (NOU, P, S), HE860164, HE860080, HE995722. Pichonia grandiflora Swenson & Munzinger, New Caledonia, Swenson & Munzinger 930 (NOU, P, S), HE860165, HE860081, HE995611. Pichonia lecomtei (Guillaumin) T.D. Penn., New Caledonia, Munzinger & al. 2170 (MO, NOU, P, S), HE860166, DQ154061, HE995723. Pichonia 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. Planchonella aneityensis (Guillaumin) H.J. Lam ex P.Royen, Vanuatu, Munzinger, Lowry & Tuiwawa 3665 (NOU, P, S, SUVA), HE860169, HE860083, HE995724. Planchonella anteridifera (C.T. White & W.D. Francis ex Lane-Poole) H.J. Lam, New Guinea, Takeuchi & Ama 17902 (LAE, S), HE860170, EF025109, HE995750. Planchonella arnhemica (F. Muell. ex Benth.) P. Royen, Australia, Harwood 1170 (S), HE860171, AY552107, HE995663. Planchonella asterocarpon (P. Royen) Swenson, Bartish & Munzinger, Australia, Bartish & Ford 25 (BRI, S), HE860172, DQ154078, HE995664. Planchonella australis (R. Br.) Pierre, Australia, Floyd s.n. (S), HE860173, AY552148, HE995613. Planchonella baillonii (Zahlbr.) Dubard, New Caledonia, Munzinger & Dagostini 2119 (MO, NOU, P, S), HE860174, HE860084, HE995725. Planchonella cauliflora Munzinger & Swenson, New Caledonia, Munzinger & al. 3495 (K, MO, NOU, NSW, P, S), HE860175, HE860085, HE995666. Planchonella chartacea (F. Muell. ex Benth.) H.J. Lam, Australia, Bartish & Jessup 1 (BRI, S), HE860176, HE862231, HE995665. Planchonella clemensii (Lecomte) P. Royen, China, Hainan, How 73783 (S), HE860177, HE860086, HE995692. Planchonella costata (Endl.) Pierre, New Zealand (cultivated in USA), Peter W. Fritsch 1770 (CAS), —, AF396230, —. Planchonella costata (Endl.) Pierre, New Zealand (cultivated in Edinburgh), Robertson 9 (E), HE860178, —, HE995726. Planchonella cotinifolia (A.DC.) Dubard, Australia, Bartish & Jessup 11 (BRI, S), HE860179, DQ154066, HE995701. Planchonella crassinervia Dubard, New Caledonia, Munzinger 2275 (NOU, P, S), HE860180, DQ154067, HE995667. Planchonella cyclopensis P. Royen, New Guinea, Polak 1365 (L), HE860181, HE860087, HE995693. Planchonella cyclopensis P. Royen, New Guinea, Takeuchi, Ama & Gambia 25495 (LAE, S), HE860182, HE860088, HE995751. Planchonella dothioensis (Aubrév.) Swenson, Bartish & Munzinger, New Caledonia, Munzinger 995 (MO, NOU, P, S), HE860183, AY552138, HE995668. Planchonella duclitan (Blanco) Bakh.f., New Guinea, Fuentes & Fernando 37140 (L), HE860184, HE860089, —. Planchonella eerwah (F.M. Bailey) P. Royen, Australia, Floyd s.n. (S), EU661400, AY552147, HE995669. Planchonella endlicheri (Montrouz.) Guillaumin, New Caledonia, Munzinger, Lowry & Létocart 2038 (NOU, P, S), HE860185, DQ154068, HE995614. Planchonella ericiflora Munzinger & Swenson, New Caledonia, Munzinger & al. 4197 (MO, NOU, P, S), HE860186, HE860090, HE995660. Planchonella euphlebia (F. Muell.) Francis, Australia, Bartish & Ford 18 (BRI, MO, S), HE860187, DQ154069, HE995727. Planchonella firma (Miq.) Dubard, New Guinea, Takeuchi, Ama & Jisaka 21437 (LAE, S), HE860188, HE860091, HE995615. Planchonella glauca Swenson & Munzinger, New Caledonia, Swenson, McPherson & Mouly 625 (S), HE860189, AY552104, HE995728. Planchonella grayana H.St.John, French Polynesia, Australs, Butaud 192 (PAP), HE860191, HE860092, HE995616. Planchonella grayana var. florencei Fosberg, French Polynesia, Tahiti, Florence 3967 (BISH, PAP, S), —, HE860093, —. Planchonella grayana H. St.John, French Polynesia, Tuamotus, Butaud & Lagouy 1173 (PAP), HE860193, HE860094, HE995617. Planchonella grayana H. St.John, Wallis and Futuna, Alofi, Munzinger 5479 (NOU, P, S), HE860190, HE860095, HE995729. Planchonella grayana H. St.John, Wallis and Futuna, Futuna, Munzinger, Meyer & Jourdan 5282 (P, S), HE860192, HE860096, HE995730. Planchonella howeana (F. Muell.) Pierre, Lord Howe Island, Le Cussan 1210 (BRI), HE860194, EF025094, HE995681. Planchonella "Ile Yande", New Caledonia, Swenson & Munzinger 715 (S), HE860195, EF025091, —. Planchonella kaalaensis Aubrév., New Caledonia, Swenson & Munzinger 706 (S), HE860196, HE860097, HE995694. Planchonella koumaciensis Aubrév., New Caledonia, Munzinger 2665 (NOU), HE860197, EF025095, HE995682. Planchonella kuebiniensis Aubrév., New Caledonia, Munzinger & Létocart 2057 (S), HE860198, DQ154070, HE995670. Planchonella laetevirens (Baill.) Pierre ex Dubard, New Caledonia, Munzinger, Pignal & Lowry 2001 (MO, NOU, P, S), HE860199, DQ154071, HE995618. Planchonella lamprophylla (K. Krause) H.J. Lam, New Guinea, Takeuchi, Ama & Gambia 25573 (LAE, S), HE860200, HE860098, HE995619. Planchonella lauracea (Baill.) Dubard, New Caledonia, McPherson & Munzinger 18070 (MO, S), HE860201, AY552145, HE995731. Planchonella ledermannii (K. Krause) H.J. Lam, New Guinea, Takeuchi, Towati, Jisaka & Ama 17700 (LAE, S), HE860202, HE860099, HE995732. Planchonella linggensis (Burck) Pierre, Wallis and Futuna, Futuna, Munzinger 5330 (NOU, P, S), HE860203, HE860100, HE995733. Planchonella linggensis (Burck) Pierre, New Guinea, Takeuchi, Jisaka, Towati & Ama 21108 (LAE, S), HE860204, HE860101, HE995695. Planchonella linggensis (Burck) Pierre, Vanuatu, Wheatley 271 (K), —, DQ154062, —. Planchonella luteocostata Munzinger & Swenson, New Caledonia, Munzinger, Jaffré & Roumagnac 2375 (NOU, S), EU661401, EF025099, HE995755. Planchonella maingayi (C.B. Clarke) P. Royen, Singapore, Wilkie & Gwee 507 (E), HE860205, HE860102, HE995620. Planchonella mandjeliana Munzinger & Swenson, New Caledonia, Munzinger & Pillon 2861 (NOU, P, S), HE860206, EF025100, HE995680. Planchonella membranacea H.J. Lam, Fiji, Smith 4609 (S), HE860207, DQ154074, HE995621. Planchonella mindanaensis H.J. Lam, Philippines, Clemens 1015 (PNH), HE860208, HE860103, HE995622. Planchonella minutiflora Munzinger & Swenson, New Caledonia, MacKee 16639 (NOU, P, S), HE860209, HE860104, HE995734. Planchonella moluccana (Burck) H.J. Lam, New Guinea, Takeuchi, Ama & Gambia 25550 (S), HE860139, HE860072, HE995671. Planchonella "Munzinger 6150", New Caledonia, Munzinger 6150 (NOU) , HE860210, HE860105, HE995735. Planchonella "Munzinger 6490", Fiji, Munzinger 6490 (NOU, SUVA) , HE860211, HE860106, HE995736. Planchonella "Munzinger 6514", Fiji, Munzinger 6514 (NOU, SUVA) , HE860212, HE860107, HE995737. Planchonella myrsinifolia (F. Muell.) Swenson, Bartish & Munzinger, Australia, Floyd s.n. (S), HE860213, AY552143, HE995702. Planchonella myrsinoides (Benth.) S.T. Blake ex Francis, Australia, McDonald 3365 (QRS, S), HE860214, EF025092, HE995738. Planchonella obovata (R. Br.) Pierre, Taiwan, Chung & Anderberg 1166 (HAST, S), EU661402, DQ154076, HE995739. Planchonella "Pillon 150", New Caledonia, Pillon, Barrabé & Rigault 150 (NOU, P, S), HE860215, HE860108, HE995696. Planchonella pohlmaniana (F. Muell.) Pierre ex Dubard, Australia, Bartish & Ford 22 (BRI, S), HE860216, DQ154079, HE995672. Planchonella pronyensis Guillaumin, New Caledonia, Munzinger 2051 (NOU), HE860217, DQ154080, HE995623. Planchonella roseoloba Munzinger & Swenson, New Caledonia, Munzinger 2311 (NOU, P, S), HE860218, DQ154090, HE995673. Planchonella rufocostata Munzinger & Swenson, New Caledonia, Munzinger, Labat & Butin 2583 (NOU, S), HE860219, DQ154089, HE995684. Planchonella saligna S. Moore, New Caledonia, Munzinger, Létocart & Gâteblé 2218 (NOU, P, S), HE860220, DQ154083, HE995624. Planchonella sandwicensis (A. Gray) Pierre, Hawaii, Ohao, Koolan 119d (GB), HE860221, DQ154084, HE995740. Planchonella sandwicensis (A. Gray) Pierre, Hawaii, Kauai, Merello, Bess & Johnson 3227 (MO, S) , HE860222, HE860109, HE995741. Planchonella sandwicensis (A. Gray) Pierre, Hawaii, Ohau, Doug Okamoto s.n. (S), HE860223, HE860110, HE995742. Planchonella skottsbergii Guillaumin, New Caledonia, Munzinger, Gâteblé & Amice 2391 (NOU, P, S), HE860224, DQ154085, HE995674. Planchonella smithii (P. Royen) A.C. Sm., Fiji, Munzinger 6495 (NOU, S, SUVA), HE860225, HE860111, HE995743. Planchonella solida P. Royen, New Guinea, Takeuchi, Towati & Ama 17286 (LAE, S), HE860226, EF025104, HE995675. Planchonella sphaerocarpa (Baill.) Dubard, New Caledonia, Tronchet, Munzinger & Oddi 389 (MO, P), EU661403, AY552139, HE995625. Planchonella tahitensis (Nadeaud) Pierre ex Dubard, French Polynesia, Raiatea, Meyer & Taputuarai 3013 (NOU, PAP), HE860227, EF025108, HE995626. Planchonella tahitensis (Nadeaud) Pierre ex Dubard, French Polynesia, Tahiti, Meyer & Taputuarai 3051 (PAP), HE860228, EF025105, HE995744. Planchonella thiensis Aubrév., New Caledonia, Munzinger, Dagostini, Rigault & Kurpisz 2625 (S), HE860229, EF025106, HE995676. Planchonella thyrsoidea C.T. White, New Guinea, Whitfeld PA-2E-0072 (LAE, MIN, S), HE860230, HE860112, HE995745. Planchonella torricellensis (K. Schum.) H.J. Lam, Wallis and Futuna, Alofi, Munzinger 5473 (NOU, S), HE860231, HE860113, HE995746. Planchonella torricellensis (K. Schum.) H.J. Lam, Wallis and Futuna, Futuna, Munzinger 5284 (NOU), HE860232, HE860114, HE995747. Planchonella umbonata (P. Royen) A.C. Sm., Fiji, Smith 8298 (K), HE860233, DQ154086, HE995627. Planchonella vitiensis Gillespie, Fiji, Smith 7700 (S), HE860234, DQ154088, HE995628. Planchonella xylocarpa (C.T. White) Swenson, Bartish & Munzinger, Australia, Bartish & Ford 31 (BRI, S), HE860235, DQ154049, HE995629. Planchonella 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, —. Pouteria gillisonii Vink, New Guinea, Takeuchi & Ama 19050B (LAE, S), HE860237, HE860116, HE995748. Pouteria luzoniensis (Merr.) Baehni, New Guinea, Conn, Damas, Fazang, Paul & Kuria 5063 (L), HE860238, HE860117, HE995697. Pouteria maclayana (F. Muell.) Baehni, Papua New Guinea, Regaldo & Katik 1102 (L), HE860239, HE860118, HE995698. Pouteria malaccensis (C.B. Clarke) Baehni, Singapore, Swenson, Karim & Fadli 1017 (S, SING), HE860240, HE860119, HE995679. Pouteria pullenii Vink, New Guinea, Takeuchi, Jisaka, Towati & Ama 21157 (LAE, S), —, HE860120, HE995630. Pouteria richardii (F. Muell.) Baehni, Australia, Dixon & Leach 1032 (BRI, DNA, S), HE860134, HE860069, HE995749. Pouteria stellibacca J.F. Maxwell, Thailand, Boonkongchart & Chongko 146 (L), HE860241, HE860121, HE995700. Pouteria 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. Pycnandra atrofusca Swenson & Munzinger, New Caledonia, Munzinger & al. 2618 (NOU, P, S), EU661419, EU661443, HE995632. Pycnandra balansae (Baill.) Swenson & Munzinger, New Caledonia, Munzinger & al. 1451 (S), EU661387, AY552123, HE995754. Pycnandra belepensis Swenson & Munzinger, New Caledonia, Ile Art, Swenson, Munzinger & Barrabé 913 (S), HE860243, HE860123, HE995687. Pycnandra benthamii Baill., New Caledonia, Munzinger, Létocart & Gâteblé 2228 (NOU, P, S), EU661404, EU661436, HE995633. Pycnandra blaffartii Swenson & Munzinger, New Caledonia, Swenson, McPherson & Mouly 597 (NOU, S), EU661423, AY552127, HE995634. Pycnandra blanchonii (Aubrév.) Swenson & Munzinger, New Caledonia, Munzinger, Labat, Leveque & Mandaoué 2576 (S), EU661390, DQ154059, HE995635. Pycnandra bracteolata Swenson & Munzinger, New Caledonia, Munzinger, Pillon & Butin 2885 (NOU, P, S), EU661421, EU661445, HE995636. Pycnandra caeruleilatex Swenson & Munzinger, New Caledonia, Munzinger & al. 2622 (MO, NOU, P, S), EU661426, EU661448, HE995637. Pycnandra canaliculata Swenson & Munzinger, New Caledonia, Munzinger & al. 2067 (MO, NOU, P, S), EU661431, DQ154092, HE995638. Pycnandra carinocostata Vink, New Caledonia, McPherson & Munzinger 18091 (MO, NOU, P, S), EU661405, AY552132, HE995639. Pycnandra comptonii (S. Moore) Vink, New Caledonia, Lowry, McPherson & Le Borgne 5780A (MO, S), EU661407, AY552131, HE995640. Pycnandra controversa (Guillaumin) Vink, New Caledonia, Lowry, McPherson & Le Borgne 5787 (MO, S), EU661408, AY552126, HE995641. Pycnandra cylindricarpa Swenson & Munzinger, New Caledonia, Swenson, McPherson & Mouly 615 (MO, NOU, S), EU661429, AY552110, HE995757. Pycnandra decandra (Montrouz.) Vink, New Caledonia, Ile Art, Swenson, Munzinger & Barrabé 920 (S), HE860244, HE860124, HE995688. Pycnandra deplanchei (Baill.) Swenson & Munzinger, New Caledonia, Munzinger 978 (MO, NOU, P, S), EU661380, AY552120, HE995642. Pycnandra fastuosa (Baill.) Vink, New Caledonia, Munzinger & Swenson 2993 (NOU, S), EU661394, EU661434, HE995643. Pycnandra francii (Guillaumin & Dubard) Swenson & Munzinger, New Caledonia, Munzinger 965 (MO, NOU, P), EU661391, AY552117, HE995644. Pycnandra glabella Swenson & Munzinger, New Caledonia, Munzinger & al. 2615 (NOU, P, S), EU661418, EU661442, HE995645. Pycnandra glaberrima Swenson & Munzinger, New Caledonia, Munzinger & al. 1394 (MO, NOU, P, S), EU661399, AY552133, HE995646. Pycnandra gordoniifolia (S.Moore) Swenson & Munzinger, New Caledonia, Swenson & Munzinger 726a (BRI, MO, NOU, P, S), EU661392, EU661433, HE995647. Pycnandra griseosepala Vink, New Caledonia, Swenson, McPherson & Mouly 627 (MO, NOU, S), EU661414, AY552128, HE995648. Pycnandra kaalaensis Aubrév., New Caledonia, Munzinger & Labat 2599 (NOU, S), EU661415, EU661440, HE995756. Pycnandra linearifolia Swenson & Munzinger, New Caledonia, Munzinger & Blaffart 2786 (NOU, P, S), EU661427, EU661440, HE995753. Pycnandra neocaledonica (S.Moore) Vink, New Caledonia, Tronchet, Munzinger & Oddi 426 (MO, NOU, P, S), EU661416, AY552129, HE995649. Pycnandra ouaiemensis Swenson & Munzinger, New Caledonia, Munzinger, Lowry, Blaffart & Brown 3135 (NOU, S), EU661422, EU661446, HE995689. Pycnandra paucinervia Swenson & Munzinger, New Caledonia, Munzinger & al. 1438 (NOU, P, S), EU661424, AY552159, HE995650. Pycnandra pubiflora Swenson & Munzinger, New Caledonia, Munzinger & al. 2624 (NOU, P, S), EU661420, EU661444, HE995651. Pycnandra sarlinii (Aubrév.) Swenson & Munzinger, New Caledonia, Munzinger 1860 (NOU, P, S), EU661395, EU661435, HE995652. Pycnandra schmidii (Aubrév.) Swenson & Munzinger, New Caledonia, McPherson & Munzinger 18106 (MO, NOU, P, S), EU661396, AY552116, HE995653. Pycnandra sessiliflora Swenson & Munzinger, New Caledonia, Munzinger & McPherson 696 (BRI, MO, NOU, P, S), EU661398, AY552161, HE995691. Pycnandra sessilifolia (Pancher & Sebert) Swenson & Munzinger, New Caledonia, McPherson & Munzinger 18176 (MO, P), EU661397, AY552118, HE995654. Pycnandra vieillardii (Baill.) Vink, New Caledonia, Dumontet, Zongo & Maituku s.n. (S), EU661417, EU661441, HE995655. Pycnandra viridiflora Swenson & Munzinger, New Caledonia, Munzinger, McPherson & Tuiwawa 4195 (NOU, S), HE860245, HE860125, HE995656. Sersalisia sericea (Aiton) R. Br., Australia, Harwood 1172 (S), HE860246, AY552112, HE995657. Sersalisia 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. A B D E F I Figure 1. C G J H K L 1 97 1 99 1 100 1 100 .97 64 0.96 1 59 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 1 87 1 0.97 100 73 1 0.97 100 0.86 1 100 1 100 0.96 1 51 100 0.99 0.93 61 1 78 0.98 60 1 76 1 100 1 87 1 100 1 100 1 97 0.8 1 84 1 64 1 88 0.92 59 63 0.98 85 1 92 1 95 D2 1 67 1 100 1 100 0.97 52 1 76 1 66 1 100 1 100 1 90 Current classification Beccariella Chrysophyllum Krausella Leptostylis Niemeyera Magodendron Pichonia Planchonella Pycnandra Pouteria Sersalisia Van-royena 1 86 1 99 D1 1 1 74 1 0.99 100 64 0.86 64 1 99 1 100 0.85 1 63 1 100 0.82 1 88 1 95 D3 0.8 1 90 1 100 1 100 1 .92 1 73 1 100 1 100 1 68 1 100 1 99 0.95 0.8 0.8 58 1 97 1 0.81 1 100 1 100 .87 1 96 Figure 2. 1 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 Pichonia Magodendron Niemeyera antiloga Magodendron Niemeyera antiloga Niemeyera Niemeyera Pycnandra Pycnandra Planchonella Planchonella cup-shaped urn-shaped narrowly campanulate tubular in tube orifice just below tube orifice in middle of the tube near the base Beccariella Beccariella F Seed characters Van-royena Sersalisia Van-royena Sersalisia Pichonia Pichonia Magodendron Niemeyera antiloga Magodendron Niemeyera antiloga Niemeyera Niemeyera Pycnandra Pycnandra with stigmatic areas simple Planchonella Planchonella Figure 3. Van-royena Sersalisia Pichonia E Style pubescent glabrous Beccariella Beccariella C Flowers Beccariella Beccariella A Higher leaf venation cotyledons foliaceous, radicle exserted and endosperm present cotyledons plano-convex, radicle included and endosperm absent cotyledons plano-convex, radicle exserted and endosperm absent 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 Figure 4. Planchonella F 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