A New Subfamily Classification of the Palm Family (Arecaceae)

Blackwell Publishing LtdOxford, UKBOJBotanical Journal of the Linnean Society0024-4074The Linnean Society of London, 2006? 2006 151? 1538 Original Article

PALM FAMILY PHYLOGENY C. B. ASMUSSEN ET AL. Botanical Journal of the Linnean Society, 2006, 151, 15–38. With 3 figures

The Palms Guest edited by William J. Baker and Scott Zona

A new subfamily classiﬁcation of the palm family (Arecaceae): evidence from plastid DNA phylogeny

CONNY B. ASMUSSEN1*, JOHN DRANSFIELD2, VINNIE DEICKMANN1, ANDERS S. BARFOD3, JEAN-CHRISTOPHE PINTAUD4 and WILLIAM J. BAKER2

1Department of Ecology, Royal Veterinary and Agricultural University Copenhagen, Rolighedsvej 21, DK- 1958 Frederiksberg, Denmark 2Herbarium, Royal Botanic Gardens, Kew, Surrey TW9 3AE, UK 3Department of Biological Sciences, University of Aarhus, Ny Munkegade bygn. 541, DK-8000 Århus C, Denmark 4Institut de Recherche pour le Développement, UMR DGPC/DYNADIV, 911 Avenue Agropolis BP 64501, 34394 Montpellier cedex 5, France

Received June 2005; accepted for publication November 2005

Published phylogeny reconstructions of the palm family (Arecaceae) are based on plastid DNA sequences or restriction fragment length polymorphisms (RFLPs), nuclear DNA sequences, morphological characters or a combination thereof, and include between 33 and 90 palm species. The present study represents all previously recognized subfamilies, tribes and subtribes of palms and 161 of the 189 genera. The plastid DNA region matK was sequenced for 178 palm species and ten commelinid monocot outgroup species, and was combined with new and previously published plastid DNA sequences of trnL–trnF, rps16 intron and rbcL. The addition of matK sequences and more taxa resulted in a highly resolved and largely well-supported phylogeny. Most importantly, critical basal nodes are now fully resolved and, in most cases, strongly supported. On the basis of this phylogeny, we have established a new sub- familial classification of the palms, in which five subfamilies are recognized, rather than the six that were included in the previous classification. The circumscriptions of the subfamilies Calamoideae and Nypoideae were corroborated. The phylogeny supported a new circumscription for the subfamily Coryphoideae, including all taxa previously recognized in Coryphoideae with the addition of the tribe Caryoteae, formerly of the subfamily Arecoideae. The phylogenetic analysis also supported a new delimitation for the subfamily Ceroxyloideae that contains the tribes Cyclospatheae and Ceroxyleae, and all genera formerly included in the subfamily Phytelephantoideae, but excludes the tribe Hyophorbeae. Finally, the subfamily Arecoideae was modified to exclude the tribe Caryoteae and to include the tribe Hyophorbeae. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 15–38.

ADDITIONAL KEYWORDS: matK – Palmae – rbcL – rps16 intron – trnL–trnF.

INTRODUCTION stantial progress has been made in the understanding of the relationships within the fam- The palm family (Arecaceae, Palmae) is resolved as a ily. Many estimates of palm phylogeny have been monophyletic group in all higher-level molecular published at various taxonomic levels. Nevertheless, studies of monocots (e.g. Chase et al., 2000; Asmus- numerous ambiguities have persisted, hindering any sen & Chase, 2001). During the last 10 years, sub- attempt to rearrange the formal classiﬁcation of the family, such as, for example, the placements of the *Corresponding author. E-mail: [email protected] tribes Cyclospatheae and Phoeniceae, and of the sub-

16 C. B. ASMUSSEN ET AL. family Phytelephantoideae. In this paper, which is sister to the remaining members of Arecoideae, with focused strictly on the circumscription of palm sub- the Calamoideae sister to all palms excluding Nypa families, we shall refer primarily to those phylogeny (Uhl et al., 1995). However, this result was probably reconstructions that explore the systematics of the influenced by the use of only one taxon as an outgroup, family at the highest level (Uhl et al., 1995; Baker Dioscorea (Dioscoreaceae), which is only distantly et al., 1999; Asmussen, Baker & Dransfield, 2000; related to palms and commelinid monocots as a whole Asmussen & Chase, 2001; Hahn, 2002a; Lewis & (Chase et al., 2000), thereby increasing the potential Doyle, 2002). We use the formal subfamily, tribal for a spurious rooting. In subsequent studies, Baker and subtribal names in the sense of Dransfield & et al. (1999) and Asmussen et al. (2000) did not include Uhl (1998), who divided the family into six subfami- nonpalm outgroups, due to alignment problems, root- lies varying in size from one genus (subfamily ing their phylogenies internally on Nypa following Uhl Nypoideae) to 112 genera (subfamily Arecoideae; see et al. (1995). Although their methods were explicit, Appendix). For the subfamily Calamoideae, however, the results are prone to misinterpretation. However, we use the classification of Baker, Dransfield & Hed- another study, which included nonpalm outgroups, derson (2000a). This study provides part of the justi- supports Nypa as sister to all other palms (Lewis & fication for a forthcoming new classification of palms Doyle, 2001). Unfortunately, in none of these studies based on phylogenetic data (Dransfield et al., 2005); are the relative positions of Nypa or the Calamoideae we make references to the new classification, where strongly supported by bootstrap analysis, rendering appropriate, within the figures and in the discussion the results effectively equivocal. section below. Subfamily Coryphoideae More than half the phylogenetic analyses of the CURRENT STATUS OF PALM FAMILY PHYLOGENETICS palm family based on DNA sequences do not resolve Subfamilies Calamoideae and Nypoideae the subfamily Coryphoideae as monophyletic (Baker The subfamily Calamoideae is resolved as mono- et al., 1999; Asmussen et al., 2000; Asmussen & phyletic in all palm family phylogenies (Uhl et al., Chase, 2001; Hahn, 2002a). However, the plastid 1995; Baker et al., 1999, 2000a; Baker, Hedderson & RFLP phylogeny of Uhl et al. (1995), in which the Dransfield, 2000b, c; Asmussen et al., 2000; Asmussen taxonomic sampling was heavily biased towards & Chase, 2001; Lewis & Doyle, 2001; Hahn, 2002a). coryphoids, resolved subfamily Coryphoideae includ- Baker et al. (2000a, b, c) explored the relationships ing the tribe Caryoteae from the subfamily within this subfamily and proposed a new classifica- Arecoideae as a monophyletic group. The study of tion for the Calamoideae with three tribes and nine Lewis & Doyle (2001), based on DNA sequences of the subtribes based on a combination of molecular and nuclear gene, malate synthase, and that of Hahn morphological data. (2002a), based on a combined, reduced data set, Nypa, the sole representative of the subfamily resolved the Coryphoideae as monophyletic. It should Nypoideae, is always resolved on an isolated branch be noted, however, that the sample size was small in when maximum parsimony is employed as the opti- both cases. Many data sets group the tribe Caryoteae mality criterion (Uhl et al., 1995; Baker et al., 1999; of the subfamily Arecoideae together with members of Asmussen et al., 2000; Asmussen & Chase, 2001; the subfamily Coryphoideae, often with close relation- Lewis & Doyle, 2001; Hahn, 2002a). In a few analyses ships to the subtribe Coryphinae or the tribe Boras- using maximum likelihood, Nypa is nested in various seae (Uhl et al., 1995; Asmussen et al., 2000; positions among members of the subfamily Calam- Asmussen & Chase, 2001; Hahn, 2002a). oideae or the subfamily Coryphoideae (Hahn, 2002a), but these relationships have received scant support in Subfamilies Ceroxyloideae and Phytelephantoideae other systematic studies. It is clear from most phylogenetic analyses that the The Calamoideae and the Nypoideae are the princi- subfamily Ceroxyloideae (sensu Dransfield & Uhl, pal candidates for the position as the sister taxon to 1998) is not monophyletic (Uhl et al., 1995; Baker the remaining members of Arecaceae. In two recent et al., 1999; Asmussen et al., 2000; Asmussen & papers with extensive taxon and nucleotide character Chase, 2001; Lewis & Doyle, 2001; Hahn, 2002a, b). sampling, the subfamily Calamoideae was resolved as One tribe, the Hyophorbeae, is consistently resolved sister to all other members of the palm family in total with members of the subfamily Arecoideae. The exact evidence analyses based on parsimony (Asmussen & relationships and positions of the tribes Ceroxyleae Chase, 2001; Hahn, 2002a). This finding contrasts and Cyclospatheae are not yet clear, however. In con- markedly with the first phylogenetic study of palms trast, the subfamily Phytelephantoideae is always based on restriction fragment length polymorphisms resolved as monophyletic (Uhl et al., 1995; Asmussen (RFLPs) and morphology, in which Nypa resolved as et al., 2000; Asmussen & Chase, 2001; Hahn, 2002a)

PALM FAMILY PHYLOGENY 17 with the exception of Baker et al. (1999), where the subtribes, 14 tribes and six subfamilies in the classi- two included species are unresolved in a polytomy at fication of Uhl & Dransfield (1987) and all 36 sub- the base of the Arecoid line (Moore, 1973). tribes, 14 tribes and six subfamilies of Dransfield & Recent studies provide modest support for a clade of Uhl (1998; see Appendix) were represented. All tribes Phytelephantoideae, Ceroxyloideae, and Arecoideae, a and subtribes of the revised classification of the group that is equivalent to the Arecoid line sensu Calamoideae of Baker et al. (2000a) were also repre- Moore (1973; Asmussen & Chase, 2001; Hahn, 2002a, sented. The matK region was chosen as an additional b). Various studies give indications of potential rela- plastid DNA region because it has provided many par- tionships between the Ceroxyleae, Cyclospatheae, and simony-informative characters in other monocot stud- Phytelephantoideae, or between at least two of the ies. Other plastid DNA regions (rpl16, rpoC, and three groups (Uhl et al., 1995; Asmussen & Chase, ndhF) were tested on a small sample of species as 2001; Hahn, 2002a, b). Most strikingly, Asmussen & potential new plastid DNA markers, but these regions Chase (2001) provided evidence, albeit weakly sup- showed amplification difficulties, whereas matK ported, that Phytelephantoideae, Cyclospatheae and amplified readily in all palm test samples. All matK Ceroxyleae form a monophyletic sister group to the sequences were produced for this study and are pub- subfamily Arecoideae. lished here for the first time. In addition, rbcL, rps16, and trnL–trnF sequences for taxa not previously Subfamily Arecoideae included in our data sets were generated; all other Most studies point towards a broadly monophyletic data were recycled from three previous studies (Baker subfamily Arecoideae, with the majority including et al., 1999; Asmussen et al., 2000; Asmussen & the tribe Hyophorbeae (subfamily Ceroxyloideae) and Chase, 2001; see Appendix). Ten monocot outgroup excluding the tribe Caryoteae (Uhl et al., 1995; Baker species were selected from among the clades most et al., 1999; Asmussen et al., 2000; Asmussen & closely related to the palm family (Chase et al., 2000; Chase, 2001; Hahn, 2002a, b). see Appendix).

POTENTIAL FOR REVISION OF THE CURRENT DNA EXTRACTION, POLYMERASE CHAIN REACTION CLASSIFICATION (PCR) AND NUCLEOTIDE SEQUENCING Although much progress has been made towards a Total genomic DNA was extracted from fresh or silica robust phylogeny of the palm family, a major revision gel-dried plant material using the 2× CTAB method of of the prevailing classification based on published Doyle & Doyle (1987) or the DNeasy Plant Mini Kit phylogenetic hypotheses is premature. At the highest (Qiagen). Some of the 2× CTAB extractions were fol- level in particular, the lack of resolution and bootstrap lowed by purification on caesium chloride/ethidium support at the basal nodes forming the backbone of bromide gradients (1.55 g ml−1) or with the QIAquick the phylogeny seriously hinders the production of a PCR purification kit (Qiagen) with 35% guanidinium robust, lasting, circumscription of subfamilies. For chloride ((NH2)2C:NH.HCl). The DNA concentrations reasons outlined above, three of the current subfami- were measured on a biophotometer (Eppendorf). All lies are in particular need of clarification, namely samples were vouchered with herbarium specimens Coryphoideae, Ceroxyloideae and Phytelephantoi- (see Appendix). deae. The objectives of this study were to explore fur- The matK sequences were amplified from total ther the phylogeny of the palm family by building on genomic DNA using the primer matK-19F with trnK- previous studies (Baker et al., 1999; Asmussen et al., 2R (Table 1; Steele & Vilgalys, 1994). If amplification 2000; Asmussen & Chase, 2001) with substantially was unsuccessful, reactions were repeated using expanded taxon sampling and by adding the plastid matK-19F with matK-1862R or in two pieces with any DNA region matK to the pre-existing selection of plas- combination of the available primers (Table 1). PCR tid DNA regions (trnL–trnF, rps16 intron and rbcL) reactions (100 µl) were prepared on ice by combining used in these studies, and to use our findings to pro- 65 µl ddH2O, 10 µl 10× DNA polymerase buffer, 8 µl −1 −1 pose a formal revision of the subfamily classification of 20 µmol l MgCl2, 4 µl 10 mmol l each dNTP, 1 µl the Arecaceae. 10 mg ml−1 bovine serum albumin, 5 µl of each primer (10 µmol l−1), 1 µl 5 µ µl−1 Supertaq DNA polymerase MATERIAL AND METHODS (HT Biotechnology), and 25 ng of template DNA. The amplifications were conducted on an MJ Research SAMPLING PTC-200 thermocycler programmed as follows: one This study included 178 palm species, representing cycle at 94 °C for 3 min, 28 cycles of 94 °C for 1 min, 162 of the 189 genera recognized in the 1998 treat- 50 °C (or up to 60 °C for problematic DNA samples) for ment of Arecaceae (Dransfield & Uhl, 1998). All 36 1 min, and 72 °C for 2 min, and a final cycle at 72 °C

Table 1. Primer sequences designed for this project and sequencer (Perkin-Elmer) or a PE Applied Biosystems not previously published 3100 capillary automated DNA sequencer (Perkin- Elmer). Each base position in the forward and reverse Primer name Primer sequence sequences was checked and assembled using the program SEQUENCHER 3.0 (Gene Codes Corp.). MatK-300F 5′-AGT TCA GTA CTT GTR AAA CG-3′ MatK-445R 5′-GGG AAG ATA CTA ATC GCA GC-3′ MatK-809F 5′-CGA TTA ACA TCT TCT GGA GC-3′ SEQUENCE ALIGNMENT ′ ′ MatK-971R 5 -ATG CAT GAA GGG ATC CTT GA-3 Initial automated alignments of consensus sequences ′ ′ MatK-1315F 5 -TCG TGT GCT AGA ACT TTG GC-3 were performed with the MegAlign program (Laser- MatK-1334R 5′-GCC AAA GTT CTA GCA CAC GA-3′ gene software package, DNASTAR Inc.) and followed MatK-1862R 5′-CAT TGC ACA CGA CTT TAC C-3′ by refinement by hand. The alignment of rbcL RbcL-1407R 5′-CCA GCT TAT CTA CTG GTT CG-3′ sequences was straightforward due to the absence of length variation. The alignment of matK was also relatively straightforward except for a number of indels for 5 min. The resulting PCR products were checked at the 3′ end. For the length-variable rps16 intron and on a 0.8% agarose gel with ethidium bromide and puri- trnL–trnF sequences, the alignments included numer- fied using the QIAquick PCR purification kit (Qiagen) ous indels, but they were not recoded as additional with 35% guanidinium chloride ((NH2)2C:NH.HCl). characters. The aligned matK, rbcL, rps16 intron and The amplification primers and protocols for the rbcL trnL–trnF sequence matrices were combined and region were those described in Asmussen & Chase analysed together. For separate analyses of rbcL, (2001); a new primer, rbcL-1407R was designed and rps16 intron and trnL–trnF, see Asmussen & Chase used for DNAs that would not amplify with rbcL- (2001), Asmussen et al. (2000) and Baker et al. (1999). reverse (Fay et al., 1998). The rps16 intron region was amplified using the primers of Oxelman, Lidén & Berglund (1997) and the protocols of Asmussen CLADISTIC ANALYSES et al. (2000). The trnL–trnF region was amplified using The four data sets were readily combined because they the primers of Taberlet et al. (1991) and the protocols all originated from plastid DNA and therefore have for amplification followed Asmussen et al. (2000) identical evolutionary history, which makes congru- and Baker et al. (1999). ence tests superfluous. However, the tree statistics The concentrations of purified PCR products were indicate that the individual data sets are compatible, measured on a biophotometer and the products were because the number of nodes, the number of supported sequenced using the ABI PRISM BigDye terminator nodes and the number of highly supported nodes cycle sequencing ready reaction kit (Perkin-Elmer, AB increase in the result of the analysis of the combined Applied Biosystems). For matK, the PCR amplification data set (Table 2). The data sets were analysed by primers, matK-19F and trnK-2R, performed poorly as Fitch parsimony (Fitch, 1971; unordered, equally sequencing primers and therefore six new primers weighted characters) using PAUP* version 4.0 Beta 10 were designed as sequencing primers (Table 1). The (Swofford, 2002). The analyses yielded many trees, sequencing primers for rbcL, rps16 and trnL–trnF principally because of zero-length branches resulting were those described in Asmussen & Chase (2001), from an inadequate number of informative characters. Asmussen et al. (2000) and Baker et al. (1999) in Thus, heuristic searches could not be run to com- addition to the new rbcL-1407R (Table 1). Cycle- pletion. Therefore, the following search strategy sequencing reactions (10 µl) were prepared by combin- was used. One thousand random replicate searches ing 1 µl terminator mix, 3 µl 5× cycle-sequencing were conducted using the tree–bisection–reconnection −1 −1 buffer (200 mmol l trizma base, 5 mmol l MgCl2, (TBR) branch-swapping algorithm with steepest pH 9.0, from the BigDye terminator kit), 1 µl primer descent and MULPARS in effect, but holding five trees (1 µmol l−1), 25 ng DNA from the cleaned PCR product per step to minimize the time spent swapping on sub- and ddH2O up to 10 µl. Cycle sequencing was con- optimal trees. A round of TBR swapping was per- ducted on an MJ Research PTC-200 thermocycler pro- formed on the trees collected during the 1000 random grammed as follows: 25 cycles of 96 °C for 30 s, 50 °C replicates, collecting 30 000 optimal trees, and these for 15 s and 60 °C for 4 min trees were swapped to completion. Support for clades Cycle-sequencing products were cleaned using was calculated by conducting 1000 bootstrap repli- Dye-Ex Spin columns (Qiagen) or Sephadex G-50 cates, each with five random replicates, subtree prun- (Roche) following the protocol of the manufacturer. ing–regrafting (SPR) swapping, and saving no more The cleaned cycle-sequencing products were analysed than five trees each replicate. Only groups that on a PE Applied Biosystems 377 automated DNA appeared in > 50% of the trees were retained. Jack-

Table 2. Tree statistics for each of the individual data sets (rbcL, trnL–trnF, rps16 intron, matK) and for the combined rbcL, trnL–trnF, rps16 intron and matK data set

rbcL TrnL–trnF rps16 intron matK Combined

Length of alignment 1 306 1 842 1 569 2 385 7 102 Number of parsimony-informative characters 192 219 248 553 1 212 Tree lengths 762 655 754 1 809 4 176 Number of trees > 30 000 > 30 000 > 30 000 > 30 000 > 30 000 Consistency index 0.33 0.53 0.53 0.47 0.44 Retention index 0.65 0.70 0.71 0.74 0.69 Number of nodes in strict consensus tree 65 35 54 90 125 Number of nodes with > 50% bootstrap support 16 30 45 75 99 Number of nodes with > 90% bootstrap support 4 4 9 28 40

knife percentages and Bremer support values were pairs), and the first 57 and the last 71 base pairs of the calculated for the subfamily clades and the major rbcL gene were excluded from the analysis because nodes connecting the subfamilies. A 10 000 replicate most sequences lacked these positions (primer anneal- jackknife analysis was conducted with collapse ing regions). The data matrix thus consisted of 1306 branches if the minimum length was zero, jackknife positions, of which 192 (14.7%) were potentially with 36.79% deletion, emulate ‘Jac’ resampling in parsimony informative (Table 2). No gaps were effect, and a full heuristic search of five replicates, introduced. saving a maximum of five trees each replicate and The length of the rps16 intron sequences in palms nearest-neighbour interchange swapping. Bremer ranged from 686 (Kerriodoxa elegans) to 954 (Maxbur- support was calculated using the ‘load constraint’ retia rupicola) bases. This is the entire intron except option, and for each node conducting 1000 random for the first 31 and the last 5 base pairs. The align- replicate searches using the TBR branch-swapping ment consisted of 1569 positions (Table 2). There were algorithm with steepest descent and MULPARS in 248 (15.81%) potentially parsimony-informative char- effect and holding five trees per step. All parsimony acters. Sixty gaps varying from 1 to 341 bases in analyses were performed under DELTRAN due to the length were introduced. malfunction of ACCTRAN in PAUP* 4b version 10. The length of the trnL–trnF sequences in palms ranged from 776 (Hedyscepe canterburyana) to RESULTS 884 base pairs (Wettinia hirsuta). The alignment consisted of 1842 positions and no characters were SEQUENCE VARIATION excluded on the grounds of problematic alignment The length of sequences from the matK region (ampli- areas (Table 2). There were 219 (11.89%) potentially fication product of matK-19F and trnK-2R) in palms parsimony-informative characters. Fifty-five gaps ranged from 1800 (Mauritia flexuosa) to 1847 base varying from 1 to 166 base pairs in length were intro- pairs (Kerriodoxa elegans). Most of matK and part of duced in the alignment. the spacer between matK and the 3′ end of the split The combined matrix of matK, rbcL, rps16 intron gene trnK were included in the alignment. The begin- and trnL–trnF consisted of 7102 characters, all ning of matK could not be identified and the last c. included in the analyses (Table 2). There were 1212 100 base pairs of the spacer before the 3′ end of trnK (17.07%) potentially parsimony-informative charac- were excluded from the alignment and analyses ters. No characters were excluded on the grounds of because many sequences lacked these positions due to problematic alignment areas. The 1212 potentially differences in the reverse primer used to obtain the parsimony-informative characters included 844 char- PCR product. The data matrix thus consisted of 2385 acters without any gap positions and 368 characters positions, of which 553 (23.19%) were potentially par- with at least one gap position among the 188 included simony informative (Table 2). Approximately 50 gap taxa. areas varying from 1 to 204 bases in length were intro- The 30 000 equally most-parsimonious trees col- duced. The larger gaps were distributed in the inter- lected in the Fitch parsimony analysis were 4176 steps genic spacer between matK and the 3′ end of trnK. long and had a consistency index of 0.44 (exclud- Only the coding region of the rbcL amplification ing autapomorphies) and a retention index of 0.69 product was included in the alignment (1434 base (Table 2). The tree lengths of the cladograms resulting

© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 15–38 20 C. B. ASMUSSEN ET AL. from the combined analyses were longer than the World genera of the subtribe Thrinacinae (tribe Cory- combined lengths of the four individual data sets pheae; 100% bootstrap support; Fig. 1, clade b). The (762 + 655 + 754 + 1809 = 3980), indicating that the second major clade, which was weakly supported (61% combined analysis recovered homoplasy not present in bootstrap support) as sister to the third (described each of the individual analyses. below), was resolved with 99% bootstrap support and consisted of exclusively Old World taxa: a highly corroborated (100% bootstrap support) monophyletic PHYLOGENETIC ANALYSES group of three members of the subtribe Coryphinae The strict consensus tree of the combined data set was (Nannorrhops, Kerriodoxa and Chuniophoenix; Fig. 1, well resolved and included many well-supported clade c), a highly supported (100% bootstrap support) clades (Fig. 1). The palm family was resolved as mono- tribe Caryoteae (subfamily Arecoideae, Fig. 1, clade d), phyletic with a bootstrap support of 100%. The most the genus Corypha (subtribe Coryphinae; 100% boot- resolved individual tree had seven polytomies of three strap support; Fig. 1, clade e) and a highly supported taxa each (Fig. 2). Five of these seven polytomies of (100% bootstrap support) tribe Borasseae (Fig. 1, the most resolved individual tree were present in all clade f). The clade comprising Caryoteae, Corypha and individual trees (Fig. 2, arrows 1–5). Polytomy 6, the Borasseae was monophyletic with bootstrap sup- including four coryphoid taxa, and polytomy 7, com- port of 91%, whereas the support for Corypha as sister prising three large clades in the subfamily Arecoideae, to the Borasseae was low (66% bootstrap support). The were present only in a subgroup of the 30 000 most- third major clade received 86% bootstrap support and parsimonious trees (Fig. 2, arrows 6 and 7). Five of the consisted of the monogeneric tribe Phoeniceae (100% seven polytomies were positioned in the subfamily bootstrap support; Fig. 1, clade g) and a highly sup- Arecoideae, where particularly backbone branch ported (99% bootstrap support) clade composed of a lengths were short compared with backbone branch paraphyletic subtribe Livistoninae (tribe Corypheae) lengths within the other four subfamilies (Fig. 2). within which a monophyletic, well-supported (86% In the strict consensus tree, the monophyletic (100% bootstrap support) clade of all Old World genera of bootstrap) subfamily Calamoideae (Fig. 1, clade 1) was Thrinacinae (tribe Corypheae) was embedded (Fig. 1, resolved as sister to the rest of the palms. Within clade h). Calamoideae, Eugeissona (tribe Eugeissoneae) was There was 85% bootstrap support for the clade cor- sister to the rest of the Calamoideae (59% bootstrap responding to Moore’s (1973) Arecoid Line comprising support). Additionally, Calamoideae were divided into the subfamilies Ceroxyloideae, Phytelephantoideae two large monophyletic groups corresponding to the and Arecoideae, excluding Caryoteae (Fig. 1, clades 4 two tribes Lepidocaryeae (91% bootstrap support) and and 5). The subfamily Phytelephantoideae was mono- Calameae (70% bootstrap support). The Lepidocar- phyletic (99% bootstrap support) and together with yeae clade consisted of the African and American taxa two monophyletic tribes, Cyclospatheae (100% boot- Mauritia (subtribe Mauritiinae), Raphia (subtribe strap support) and Ceroxyleae (99% bootstrap sup- Raphiinae) and Oncocalamus, Laccosperma and Ere- port) of the subfamily Ceroxyloideae is denoted as mospatha (all three from the subtribe Ancistrophylli- clade 4 on Figure 1 (63% bootstrap support). nae). The Calameae clade consisted of the largely The remaining large clade (Fig. 1, clade 5) was South-east Asian taxa Korthalsia (Korthalsiinae), weakly supported (70% bootstrap support) and con- Salacca (Salaccinae), Calamus (Calaminae), Pigafetta sisted of all genera from the subfamily Arecoideae, (Pigafettinae), Plectocomia (Plectocomiinae) and except for Caryoteae, with the addition of the tribe Metroxylon (Metroxylinae). Hyophorbeae (subfamily Ceroxyloideae). The tribe Iri- Nypa fruticans, from the monospecific subfamily arteeae was monophyletic (98% bootstrap support) Nypoideae, was sister to the remaining palms (namely and sister to a clade (76% bootstrap support) of the Calamoideae not included) with a bootstrap support of remaining members of clade 5. Within this latter 97% (Fig. 1, clade 2). clade, the base of which is highly unresolved, Som- The subfamily Coryphoideae, including Caryoteae mieria and Pelagodoxa (subtribe Iguanurinae) formed from the subfamily Arecoideae, formed a monophyletic a monophyletic group (94% bootstrap support). The lineage with bootstrap support of 97% (Fig. 1, clade 3). monophyly of the tribe Hyophorbeae (subfamily All taxa in this clade have induplicate leaves, except Ceroxyloideae) was highly supported by bootstrap for the anomalous coryphoid genus Guihaia. Three (100%). The tribe Geonomeae was resolved as non- major clades received high bootstrap support. The first monophyletic, Welfia and Pholidostachys forming a of these, with bootstrap support of 98%, consisted clade with Manicaria (subtribe Manicariinae; 89% entirely of the New World taxa: the genus Sabal (100% bootstrap support), whereas a clade of Asterogyne, bootstrap support; Fig. 1, clade a), sole member of the Geonoma, Calyptronoma and Calyptrogyne (80% boot- subtribe Sabalinae (tribe Corypheae), and the New strap support) resolved elsewhere. The latter group of

Geonomeae was sister to a monophyletic subtribe DNA areas were used (Shaw et al., 2005). The rbcL Euterpeinae (56% bootstrap support). The tribe gene produced the fewest parsimony-informative Cocoeae was not supported as monophyletic: Beccari- characters (192), but these variable characters ophoenix (subtribe Beccariophoenicinae) was sister to resulted in 65 resolved nodes in the palm family, Sclerosperma (Sclerospermatinae) with less than 50% whereas the trnL–trnF region and the rps16 intron bootstrap support. The remaining members of Cocoeae produced only 35 and 54 resolved nodes, respectively, formed an unsupported monophyletic group with despite supplying more informative characters (219 Reinhardtia as the sister group, again without sup- and 248; Table 2). The number of clades with more port. The subtribe Elaeidinae (Elaeis) was included in than 90% bootstrap support was relatively low in each a monophyletic group with the subtribe Bactridinae of the individual data sets (four, four and nine), except (Desmoncus, Bactris, Aiphanes and Acrocomia; 73% for matK (28), but the combined data set produced the bootstrap support); and the subtribes Butiinae (Alla- largest number of highly supported (> 90%) clades (40; goptera, Syagrus, Cocos, Voanioala, Jubaeopsis) and Table 2). Attaleinae (Attalea) formed a well-supported monophyletic group (91% bootstrap). Most of the species representing Indo-Pacific pseudomonomerous genera THE NEW SUBFAMILY CLASSIFICATION from the tribe Areceae resolved in an unsupported and The dense taxon sampling and the large number of highly unresolved clade, with some notable exceptions nucleotide characters included in this study and the (Pelagodoxa, Sommieria, Iguanura). However, numer- high levels of resolution and support in the resulting ous smaller groups were resolved within this clade. trees are unprecedented in higher-level palm phyloge- Two subtribe Arecinae species pairs, Areca triandra netic research. Our results are sufficiently robust to and Nenga pumila, and Hydriastele microspadix and justify a formal reclassification of palm subfamilies H. chaunostachys, the latter representing the recently and are equally convincing at lower taxonomic levels synonymized genus Gronophyllum (Baker & Loo, in some areas (Fig. 3). Herein, we describe the ratio- 2004) constituted monophyletic lineages with 84 and nale for recognizing five subfamilies in a forthcoming 78% bootstrap support, respectively, but were not formal reclassification of the palms (Dransfield et al., resolved as sister groups. Further clades resolved 2005). within the Indo-Pacific pseudomonomerous Areceae Subfamily placements for most genera of palms clade and supported by bootstrap include: Rho- remain unchanged in the majority of cases with palostylis baueri and Hedyscepe canterburyana (sub- respect to the previous classification (Uhl & Drans- tribe Archontophoenicinae; 53% bootstrap support), field, 1987; Dransfield & Uhl, 1998; see Appendix). Masoala (subtribe Masoalinae; 82% bootstrap sup- Nevertheless, the new subfamily classification port), Marojejya (subtribe Masoalinae; 93% bootstrap requires three major rearrangements (Figs 1, 2): (1) support), Basselinia and Alloschmidia (subtribe Igua- the tribe Caryoteae from the subfamily Arecoideae nurinae; 87% bootstrap support), Acanthophoenix, sensu Dransfield & Uhl (1998) is moved to a revised Tectiphiala and Oncosperma (subtribe Oncosper- subfamily Coryphoideae; (2) subfamily Phytelephan- matinae; 50% bootstrap support), Heterospathe and toideae changes rank to tribe Phytelephanteae and is Alsmithia (subtribe Iguanurinae; 80% bootstrap included within the new circumscription of the sub- support), Laccospadix and Linospadix (subtribe Lino- family Ceroxyloideae; and (3) the tribe Hyophorbeae spadicinae; 99% bootstrap support), Cyphokentia, from the subfamily Ceroxyloideae (sensu Dransfield & Moratia, Lavoixia, Brongniartikentia and Clino- Uhl, 1998) is moved to the subfamily Arecoideae. sperma (subtribe Iguanurinae; 63% bootstrap support), and Ptychosperma, Ponapea, Balaka, Veitchia, Carpentaria, Wodyetia, Brassiophoenix and Ptycho- SUBFAMILIES CALAMOIDEAE AND NYPOIDEAE coccus (subtribe Ptychospermatinae; 62% bootstrap This study strongly supports the monophyly of the support). subfamily Calamoideae and firmly positions it as sister to the rest of the palms (Fig. 1, clade 1). Moreover, DISCUSSION our results corroborate those of Asmussen & Chase (2001) and the total evidence analyses of Hahn THE SEQUENCES (2002a). The tribal and subtribal classification of The matK sequences produced more than twice the Calamoideae (Baker et al., 2000a) is also corroborated number of parsimony-informative characters (553) for in this study. The position of the subfamily Nypoideae the same taxon sample when compared with the other as sister to all palms excluding Calamoideae is regions: rbcL (192), trnL–trnF (219) and rps16 intron strongly supported and its status as a monogeneric (248; Table 2). This is in agreement with the results subfamily is confirmed, in accordance with all previ- from other studies where two or more of these plastid ous studies (Fig. 1, clade 2; Uhl et al., 1995; Baker

70 continued

5 SUBFAMILY ARECOIDEAE

85 91 Ceroxylon quindiuense 99 Juania australis 58 Oraniopsis appendiculata 67 Ravenea louvelii 82 Aphandra natalia 63 99 Phytelephas aequatorialis Formerly Ammandra decasperma Phytelephantoideae 4 Phytelephas macrocarpa SUBFAMILY CEROXYLOIDEAE 100 Pseudophoenix sargentii Pseudophoenix vinifera Maxburrrtia rupicola Rhapis excelsa 75 Rhapidophyllum hystrix 86 52 Guihaia argyrata Trachycarpus fortunei Chamaerops humilis Brahea berlandieri 53 Acoelorrhaphe wrightii Serenoa repens 100 Colpothrinax wrightii 100 Pritchardia arecina 52 Pritchardia pacifica Washingtonia robusta 99 Johannesteijsmannia altifrons h Pholidocarpus macrocarpus Pritchardiopsis jeanneneyi 86 Licuala kunstleri Livistona chinensis Copernicia prunifera 67 Phoenix reclinata 100 Phoenix dactylifera g Phoenix canariensis 58 Bismarckia nobilis 71 Satranala decussilvae 82 Hyphaene thebaica 61 100 Medemia argun 92 Borassodendron machadonis f 55 Borassus flabillifer 66 65 Lodoicea maldivica Latania verschaffeltii 100 Corypha umbraculifera 97 91 e Corypha taliera 98 Arenga hookeriana 96 Arenga undulatifolia Caryoteae, formerly 99 100 Wallichia distica 87 Caryota mitis Arecoideae d Caryota ophiopellis 97 63 Kerriodoxa elegans 100 Nannorrhops ritchiana c 3 SUBFAMILY CORYPHOIDEAE Chuniophoenix nana 70 Hemithrinax compacta Thrinax morrisii Schippia concolor Trithrinax campestris 100 Zombia antillarum Coccothrinax argentata PALMS 100 b Thrinax radiata 98 Chelyocarpus ulei Cryosophila warscewicziana Itaya amicorum 100 Sabal bermudana a Sabal minor Nypa fruticans 2 Metroxylon salomonense SUBFAMILY NYPOIDEAE 94 Plectocomia mulleri 70 53 Pigafetta elata 70 Calamus aruensis Salacca ramosiana 57 Korthalsia cheb 59 62 Eremospatha wendlandiana 100 Laccosperma acutiflorum 100 91 Oncocalamus tuleyi 66 Raphia farinifera 1 SUBFAMILY CALAMOIDEAE Mauritia flexuosa Eugeissona tristis 100 83 Hanguana malayana 86 Tradescantia pallida 100 Anigozanthos manglesii 100 Canna edulis Musa rosea 63 Fargesia sp. 100 Typha angustifolia Vriesia psittacina Dasypogon bromelifolius Kingia australis

Figure 1. The strict consensus tree of 30 000 equally most-parsimonious cladograms resulting from Fitch parsimony analyses of combined matK, rbcL, rps16 intron and trnL–trnF data sets. Bootstrap percentages for the clades are given above the branches. Five clades are labelled 1–5 for the discussion of the new subfamily classification in the text. The clades corresponding to the five subfamilies of the new classification are labelled with the subfamily name. Another eight clades are labelled a–h for the discussion of the new tribal classification within the subfamily Coryphoideae. The changes to the classification of Dransfield & Uhl (1998) are indicated with boxes to the right.

81 Ptychosperma macarthurii Ponapea ledermannianus 92 Balaka seemannii 62 Veitchia arecina 99 Carpentaria arcuminata Wodyetia bifurcata 66 Brassiophoenix schumannii Ptychococcus paradoxus 99 Cyphokentia macrostachys Moratia cerifera 63 87 Lavoixia macrocarpa Brongniartikentia lanuginosa Clinosperma bracteale 99 Linospadix monostachya 53 Laccospadix australasica Dransfieldia micrantha 80 Heterospathe elata Heterospathe longipes 52 Carpoxylon macrospermum Satakentia liukiuensis Bentinckia nicobarica Clinostigma savoryanum 94 Acanthophoenix rubra 50 Tectiphiala ferox Oncosperma tigillarium 84 Areca triandra Nenga pumila 87 Alloschmidia glabrata Basselinia velutina 93 Marojejya darianii Marojejya insignis 82 Masoala madagascariensis Masoala kona 53 Rhopalostylis baueri Hedyscepe canterburyana 78 Hydriastele microspadix Hydriastele chaunostachys 61 Cyphosperma balansae Physokentia rosea Actinorhytis calapparia Archontophoenix purpurea Actinokentia divaricata Chambeyronia macrocarpa Kentiopsis oliviformis Campecarpus fulcitus Cyphophoenix nucele Veillonia alba Neoveitchia storckii Dypsis lutescens Lemurophoenix halleuxii Calyptrocalyx albertisiana Phoenicophorium borsigianum Roscheria melanochaetes Cyrtostachys renda Dictyosperma album Lepidorrhachis mooreana Loxococcus rupicola Rhopaloblaste augusta 60 Allagoptera arenaria 85 Syagrus smithii 74 Attalea allenii 80 Cocos nucifera 91 Voanioala gerardii 76 Jubaeopsis caffra Acrocomia aculeata Acrocomia crispa Aiphanes aculeata 73 Bactris gasipaes Desmoncus orthacanthos Elaeis guineensis Reinhardtia simplex 82 Prestoea pubens 82 Neonicholsonia watsonii 56 Euterpe oleracea Hyospathe macrorhachis 91 Calyptrogyne ghiesbreghtiana Calyptronoma occidentalis 80 Geonoma congesta Asterogyne martiana 56 Chamaedorea microspadix Gaussia maya Hyophorbeae, formerly continued 70 100 Synechanthus warscewiczianus Hyophorbe lagenicaulis Ceroxyloideae 5 SUBFAMILY ARECOIDEAE Wendlandiella gracilis 65 Welfia regia 89 Pholidostachys pulchra 51 Manicaria saccifera Leopoldinia pulchra 100 Podococcus barteri Podococcus barteri Sclerosperma mannii Beccariophoenix madagascariensis 94 Pelagodoxa henryana Sommieria leucophylla 92 Orania lauterbachiana Orania ravaka Roystonea oleracea Iguanura wallichiana 77 Iriartella stenocarpa Wettinia hirsuta 98 Dictyocaryum lamarckianum Iriartea deltoidea Socratea exorrhiza

Figure 1. Continued

Carpentaria arcuminata Wodyetia bifurcata 5 Brassiophoenix schumannii Ptychococcus paradoxus Ptychosperma macarthurii 4 Ponapea ledmannianus Balaka seemannii Veitchia arecina Lepidorrhachis mooreana Archontophoenix purpurea Cyphophoenix nucele Veillonia alba Rhopalostylis baueri Hedyscepe canterburyana Cyphosperma balansae 3 Physokentia rosea Alloschmidia glabrata Basselinia velutina Calyptrocalyx albertisiana 2 Neoveitchia storckii Cyphokentia macrostachys Moratia cerifera Lavoixia macrocarpa Brongniartikentia lanuginosa Clinosperma bracteale Linospadix monostachya Laccospadix australasica Dransfieldia micrantha Heterospathe elata Heterospathe longipes Actinorhytis calapparia Chambeyronia macrocarpa Kentiopsis oliviformis Actinokentia divaricata Masoala madagascariensis Masoala kona Roscheria melanochaetes Hydriastele microspadix Hydriastele chaunostachys 7 Carpoxylon macrospermum SUBFAMI Satakentia liukiuensis Bentinckia nicobarica Clinostigma savoryanum Dictyosperma album Lemurophoenix halleuxii Loxococcus rupicola Rhopaloblaste augusta Marojejya darianii Marojejya insignis L

Dypsis lutescens Y

Acanthophoenix rubra ARECOI Tectiphiala ferox Oncosperma tigillarium Areca triandra Nenga pumila Cyrtostachys renda Phoenicophorium borsigianum Campecarpus fulcitus Pelagodoxa henryana Sommieria leucophylla DEA Iguanura wallichiana Prestoea pubens Neonicholsonia watsonii Euterpe oleracea E Hyospathe macrorhachis Calyptrogyne ghiesbreghtia Calyptronoma occidentalis Geonoma congesta Asterogyne martiana Welfia regia Pholidostachys pulchra Manicaria saccifera Orania ravaka Leopoldinia pulchra Orania lauterbachiana Allagoptera arenaria Syagrus smithii Attalea allenii Cocos nucifera Voanioala gerardii Jubaeopsis caffra Gastrococos crispa Bactris gasipaes Aiphanes aculeata Desmoncus orthacanthos Acrocomia aculeata Elaeis guineensis Reinhardtia simplex Chamaedorea microspadix Gaussia maya Synechanthus warscezianus Hyophorbe lagenicaulis Wendlandiella gracilis Podococcus barteri Podococcus barteri Roystonea oleracea Sclerosperma mannii Beccariophoenix madaensis Iriartella stenocarpa Wettinia hirsuta Socratea exorrhiza Dictyocaryum lamarckianum Iriartea deltoidea Ceroxylon quindiuense Juania australis 1 Oraniopsis appendiculata Ravenea louvelii SUBFAMILY Aphandra natalia Phytelephas aequatorialis Ammandra decasperma CEROXYLOIDEAE Phytelephas macrocarpa Pseudophoenix sargentii Pseudophoenix vinifera Maxburrrtia rupicola Rhapis excelsa Rhapidophyllum hystrix Guihaia argyrata Trachycarpus fortunei Chamaerops humilis Brahea berlandieri Acoelorraphe wrightii Serenoa repens Colpothrinax wrightii Pritchardiaarecina Pritchardia pacifica Washingtonia robusta

Johannesteijst altifrons CORY SUBFAMILY Pholidocarpus macrocarpus Pritchardiopsis jeanneneyi Licuala kunstleri Livistona chinensis Copernicia prunifera Phoenix reclinata Phoenix dactylifera Phoenix canariensis Bismarckia nobilis Satranala decussilvae Hyphaene thebaica Medemia argun Borassodendron machadonis Borassus flabillifer Lodoicea maldivica 10 changes Latania verschaffeltii Corypha umbraculifera Corypha taliera PHOIDE Arenga hookeriana Arenga undulatifolia Wallichia distica Caryota mitis Caryota ophiopellis Kerriodoxa elegans Nannorrhops ritchiana Chuniophoenix nana AE 6 Hemithrinax compacta Thrinax morrisii Zombia antillarum Coccothrinax argentata Schippia concolor Cryosophila warscewicziana OUTGROUPS Itaya amicorum Chelyocarpus ulei Thrinax radiata Trithrinax campestris Sabal bermudana Sabal minor Nypa fruticans SUBFAMILY NYPOIDEAE Metroxylon salomonense Plectocomia mulleri Pigafetta elata Calamus aruensis Salacca ramosiana SUBFAMILY Korthalsia cheb Eremospatha wendlandiana CALAMOIDEAE Laccosperma acutiflorum Oncocalamus tuleyi Raphia farinifera Mauritia flexuosa Eugeissona tristis

Figure 2. One cladogram with branch lengths of the 30 000 equally most-parsimonious cladograms resulting from Fitch analyses of the combined matK, rbcL, rps16 intron and trnL–trnF data sets. A representative of the most resolved cladograms was chosen. Outgroups were excluded to make the cladogram fit one page. The cladogram is fully resolved except for seven polytomies, which are labelled 1–7. The polytomies labelled 1–5 were present in all 30 000 most- parsimonious cladograms. The clades corresponding to the five subfamilies of the new classification are indicated to the right.

70 Arecoideae The position of Coryphoideae as sister to all palms 63/1 112 genera except the Calamoideae and Nypa had only previ- 85 ously been recovered by Hahn (2002a) in a highly 87/3 reduced taxon sample. 63 Ceroxyloideae The relationship between the Coryphoid genera is 100 62/1 8 genera well resolved, and there are high bootstrap values for 100/16 many of the subclades (Fig. 1, clade 3). Two of the 97 97 Coryphoideae three tribes in Dransfield & Uhl’s (1998) classification 99/6 93/5 45 genera of the subfamily Coryphoideae are resolved as monophyletic: the tribes Borasseae (Fig. 1, clade f) 100 Nypoideae and Phoeniceae (Fig. 1, clade g). The third tribe, 100/46 1 genus Corypheae, is not monophyletic, and just one of the four constituent subtribes, Sabalinae, is monophyletic 100 Calamoideae (Fig. 1, clade a). The significance of these relationships 100/28 21 genera for classification depends on which nodes are recognized and the ranks that they are allocated. In the Figure 3. A summary tree showing the relationship of the interests of nomenclatural stability, we propose that five subfamilies in the new classification. The number of as many as possible of the current tribes and subtribes genera in each subfamily is given below the branches for are maintained, but major rearrangements of the tribe each subfamily. Bootstrap percentages that support the Corypheae and three of its four subtribes are needed subfamilies are given above the branches and jackknife to satisfy the criterion of monophyly (Fig. 1, clade 3). percentages and Bremer support values are given below Such a reorganization (Dransfield et al., 2005) results the branches. in eight tribes (Fig. 1, clades a–h) and a number of subtribes, all of which find support among other stud- et al., 1999; Asmussen et al., 2000; Asmussen & ies (Uhl et al., 1995; Asmussen et al., 2000; Asmussen Chase, 2001; Lewis & Doyle, 2001; Hahn, 2002a), not- & Chase, 2001; Hahn, 2002a). withstanding a few unusual maximum likelihood topologies presented by Hahn (2002a). SUBFAMILIES CEROXYLOIDEAE AND ARECOIDEAE SUBFAMILY CORYPHOIDEAE The bootstrap support for the subfamilies Ceroxy- The matK sequences were particularly useful for the loideae and Arecoideae is low (63 and 70%, respec- resolution of the subfamily Coryphoideae. However, tively). However, the monophyly of both subfamilies is the addition of more genera probably also contributed strongly supported by data from low copy nuclear to the improved resolution of the relationships com- DNA genes (W. J. Baker, unpubl. data). Furthermore, pared with previous studies (Asmussen & Chase, the Arecoideae is well defined by the floral triad, not- 2001). The new subfamily Coryphoideae (Fig. 1, clade withstanding the floral cluster of the Hyophorbeae 3) is modified only by the inclusion of the tribe and the presence of triads in Caryoteae. Although the Caryoteae, a relationship that can also be found Ceroxyloideae is morphologically heterogeneous, it is among the most-parsimonious solutions emerging defined by all taxa having solitary flowers. The sub- from many other phylogenetic analyses of molecular family Phytelephantoideae (sensu Dransfield & Uhl, data in the palm family (Uhl et al., 1995; Baker et al., 1998) is highly supported as monophyletic (99% boot- 1999; Asmussen et al., 2000; Asmussen & Chase, strap support), which is in agreement with other stud- 2001; Hahn, 2002a). However, until now, a robust ies (Barfod, 1991; Uhl et al., 1995; Asmussen et al., monophyletic group consisting of the subfamily 2000; Asmussen & Chase, 2001; Hahn, 2002a, b). Coryphoideae and the tribe Caryoteae had only However, given that Phytelephantoideae is nested been recovered by Uhl et al. (1995) and Hahn (2002a). between two tribes of Ceroxyloideae, Ceroxyleae and

Cyclospatheae, as sister to the former, the subfamily era formerly referred to the subtribes Livistoninae can no longer be recognized at the same rank and is and Thrinacinae. To address these problems and to placed as a tribe within the new concept of the sub- consolidate further our findings, we plan to add low family Ceroxyloideae (Fig. 1, clade 4). In the studies of copy nuclear DNA sequences and additional plastid Hahn (2002a, b), the Phytelephantoideae and the tribe DNA sequences to this data set and expand the taxon Ceroxyleae were similarly resolved, but the Cyclos- sample to include all genera of palms. Despite these patheae had a different position. The remaining shortcomings, however, we are confident that the well- studies on palm family phylogenies placed the supported relationships presented here will be robust Phytelephantoideae unresolved as a member of a poly- to the addition of new data and that our revised sub- tomy (Uhl et al., 1995; Baker et al., 1999; Asmussen family circumscriptions represent significant steps et al., 2000; Lewis & Doyle, 2002). The inclusion of the towards a natural and stable classification of palms tribe Hyophorbeae, formerly of the subfamily Ceroxy- that will stand the test of time. loideae (sensu Dransfield & Uhl, 1998), in the subfamily Arecoideae, as well as the exclusion of the tribe ACKNOWLEDGEMENTS Caryoteae, is in accordance with all previous molecular phylogenies of the palm family (Fig. 1, clade 5). The This project was supported by grants from the Danish limits of the subfamily Arecoideae require no further Research Council to Conny Asmussen and Anders alterations. Barfod. We thank Charlotte Hansen, University of Copenhagen and Hans Hjort, University of Aarhus for performing the automated sequencing. We are CONCLUSION AND FUTURE PLANS extremely grateful to all those individuals and insti- The addition of matK sequences and more taxa to the tutions who have supported our work by providing previous palm data sets of Asmussen & Chase (2001) material for DNA extraction, especially Fairchild provided the resolution and support required to refine Tropical Botanic Garden, the Montgomery Botanical the subfamily classification of the palm family (Fig. 3). Centre, the Royal Botanic Gardens, Kew, Dr Finn Five subfamilies, all monophyletic, rather than six, Borchsenius and Phillipp Trénel, University of are now recognized (Dransfield et al., 2005): (1) the Aarhus, Dr Carl Lewis, Fairchild Tropical Botanic subfamily Calamoideae, as circumscribed in Drans- Garden, and Dr Natalie Uhl, Bailey Hortorium. We field & Uhl (1998); (2) the subfamily Nypoideae, with also wish to thank an anonymous reviewer for helpful just one species, Nypa fruticans; (3) the subfamily comments. Coryphoideae, comprising those genera included by Dransfield & Uhl (1998), with the addition of the tribe REFERENCES Caryoteae; (4) the subfamily Ceroxyloideae, including the tribes Cyclospatheae and Ceroxyleae, and the Asmussen CB, Baker WJ, Dransfield J. 2000. Phylogeny of three phytelephantoid genera; (5) the subfamily the palm family (Arecaceae) based on rps16 intron and trnL– Arecoideae, following the concept of Dransfield & Uhl trnF plastid DNA sequences. In: Wilson KL, Morrison DA, (1998), but with the addition of the tribe Hyophorbeae eds. Systematics and evolution of monocots. Collingwood, and the exclusion of the tribe Caryoteae. This new Victoria: CSIRO Publishing, 525–537. subfamily classification will form the backbone of a Asmussen CB, Chase MW. 2001. Coding and noncoding plas- new edition of Genera Palmarum (Uhl & Dransfield, tid DNA in palm family systematics. American Journal of 1987; J. Dransfield, N. W. Uhl, C. B. Asmussen, W. J. Botany 88: 1103–1117. Baker, M. M. Harley & C. E. Lewis, unpubl. data). Baker WJ, Asmussen CB, Barrow S, Dransfield J, Within the new subfamilies, high resolution and Hedderson TA. 1999. A phylogenetic study of the palm bootstrap support are recovered in the Calamoideae, family (Palmae) based on chloroplast DNA sequences from the trnL–trnF region. Plant Systematics and Evolution 219: Nypoideae, Coryphoideae and, to some extent, the 111–126. Ceroxyloideae. The subfamily Arecoideae is, however, Baker WJ, Dransfield J, Hedderson TA. 2000a. Phylogeny, poorly resolved, and the internal nodes generally character evolution, and a new classification of the Calamoid receive low bootstrap support. The low resolution and palms. Systematic Botany 25: 297–322. bootstrap support in Arecoideae are principally a Baker WJ, Hedderson TA, Dransfield J. 2000b. Molecular result of a relatively low number of parsimony- phylogenetics of subfamily Calamoideae (Palmae) based on informative characters in this portion of the tree. The nrDNA ITS and cpDNA rps16 intron sequence data. Molec- most significant phylogenetic ambiguities remain in ular Phylogenetics and Evolution 14: 195–217. three areas: (1) poorly supported nodes for and some Baker WJ, Hedderson TA, Dransfield J. 2000c. Molecular within the Ceroxyloideae; (2) poor resolution and sup- phylogenetics of Calamus (Palmae) based on 5S nrDNA port for and within the subfamily Arecoideae; (3) poor spacer sequence data. Molecular Phylogenetics and Evolu- support and resolution in the clades of coryphoid gen- tion 14: 218–231.

Baker WJ, Loo AHB. 2004. A synopsis of the genus Hydri- gene malate synthase in the palm family (Arecaceae). Molec- astele (Arecaceae). Kew Bulletin 59: 61–68. ular Phylogenetics and Evolution 19: 409–420. Barfod AS. 1991. A monographic study of the subfamily Lewis CE, Doyle JJ. 2002. A phylogenetic analysis of tribe Phytelephantoideae (Arecaceae). Opera Botanica 105: 1–73. Areceae (Arecaceae) using two low-copy nuclear genes. Plant Chase MW, Soltis DE, Soltis PS, Rudall PJ, Fay MF, Systematics and Evolution 236: 1–17. Hahn WJ, Sullivan S, Joseph J, Molvray M, Kores PJ, Moore HE. 1973. The major groups of palms and their distri- Givnish TJ, Sytsma KJ, Pires JC. 2000. Higher-level sys- bution. Gentes Herbariorum 11: 27–141. tematics of the monocotyledons: an assessment of current Oxelman B, Lidén M, Berglund D. 1997. Chloroplast rps16 knowledge and a new classification. In: Wilson KL, Morrison intron phylogeny of the tribe Sileneae (Caryophyllaceae). DA, eds. Systematics and evolution of monocots. Colling- Plant Systematics and Evolution 206: 393–410. wood, Victoria: CSIRO Publishing, 3–16. Shaw J, Lickey EB, Beck JT, Farmer SB, Liu W, Miller J, Doyle JJ, Doyle JL. 1987. A rapid DNA isolation procedure Siripun KC, Winder CT, Schilling EE, Small RL. 2005. from small quantities of fresh leaf tissue. Phytochemical Bul- The tortoise and the hare II: relative utility of 21 noncoding letin 19: 11–15. chloroplast DNA sequences for phylogenetic analysis. Amer- Dransfield J, Uhl NW. 1998. Palmae. In: Kubitzki K, ed. The ican Journal of Botany 91: 142–166. families and genera of vascular plants, IV: Flowering plants, Steele KP, Vilgalys R. 1994. Phylogenetic analyses of monocotyledons. Berlin: Springer, 306–389. Polemoniaceae using nucleotide sequences of the plastid Dransfield J, Uhl NW, Asmussen CB, Baker WJ, gene matK. Systematic Botany 19: 126–142. Harley MM, Lewis CE. 2005. A new phylogenetic classifi- Swofford DL. 2002. PAUP*. Phylogenetic analysis using par- cation of the palm family, Arecaceae. Kew Bulletin 60: 559– simony (*and other methods), Version 4.0. Beta 10. Sunder- 569. land, Massachusetts: Sinauer. Fay MF, Bayer C, Alverson WS, de Bruijn AY, Chase MW. Taberlet P, Gielly L, Pautou G, Bouvet J. 1991. Universal 1998. Plastid rbcL sequence data indicate a close affinity primers for amplification of three non-coding regions of chlo- between Diegodendron and Bixa. Taxon 47: 43–50. roplast DNA. Plant Molecular Biology 17: 1105–1109. Fitch WM. 1971. Toward defining the course of evolution: min- Uhl NW, Dransfield J. 1987. Genera Palmarum: a classifica- imal change for a specific tree topology. Systematic Zoology tion of palms based on the work of H. E. Moore Jr. Lawrence, 20: 406–416. Kansas: International Palm Society and L. H. Bailey Hahn WJ. 2002a. A molecular phylogenetic study of the Hortorium. Palmae (Arecaceae) based on atpB, rbcL, and 18S nrDNA Uhl NW, Dransfield J, Davis JI, Luckow MA, Hansen KS, sequences. Systematic Biology 51: 92–112. Doyle JJ. 1995. Phylogenetic relationships among palms: Hahn WJ. 2002b. A phylogenetic analysis of the Arecoid Line cladistic analyses of morphological and chloroplast DNA of palms based on plastid DNA sequence data. Molecular restriction site variation. In: Rudall PJ, Cribb PJ, Cutler DF, Phylogenetics and Evolution 23: 189–204. Humphries CJ, eds. Monocotyledons: systematics and evolu- Lewis CE, Doyle JJ. 2001. Phylogenetic utility of the nuclear tion. Richmond: Royal Botanic Gardens, 623–661.

trn btribal classiﬁcation of Dransﬁeld 16 intron rps L rbc (K) AJ404775 AJ240870 AJ241279 AM114551 (K)(K) AM117812 AJ240868 AJ404772 AJ241277 AJ240867 AM114542 AJ241276 AM114543

(K) AJ404776 AJ240871 AJ241376 AM114541

(K) AJ829907 AJ242184 AM113612 AM114544

(BH) AJ404778 AJ240873 AJ241282 AM114552

(K) AJ404777 AJ240872 AJ241281 AM114545

(K) AJ829899 AJ242168 AM113617 AM114550 (K)(K) AM110188 AJ242175 AM113613 AM114546 AJ829897 AJ242171 AM113616 AM114549 (KEP) AJ404774 AJ240869 AJ241278 AM114540

(FTG) AM110190 AM116769 AM113615 AM114548

.W. Uhl s.n. .W. 1964–4540 (K) AM110191 AM116770 AM113618 AM114554 Dransfield JD 7571 1982–5602 (K) AJ404766 AJ240862 AJ241271 AM114553 Baker 563 Zona 651 Ely et al. 17 al. et Ely Baker 513 Baker 508 Dransfield JD 7004 Dransfield JD 7006 Rutherford 156 Baker 501 Sunderland 1759 APPENDIX Blume Mogea 1979–4409 (K) AM110189 AJ242176 AM113614 AM114547

Becc. Griff. L.f.

urmb N Becc.

Becc.

W (Mart.)

(N.J.Jacquin)

ersoon P L.H.Bailey (Warb.) Becc. (Warb.) H.Wendl. (Becc.) J.Dransf. (Becc.) (Gaertn.) Hylander wendlandiana Sunderl. Sabal bermudana Sabal minor Calamus aruensis Nypa fruticans Metroxylon salomonense Mauritia flexuosa Korthalsia cheb Salacca ramosiana Pigafetta elata Plectocomia mulleri Laccosperma acutiflorum Raphia farinifera Eremospatha Oncocalamus tuleyi Eugeissona tristis , tribe , orthalsiinae CORYPHOIDEAE Sabaleae Calamineae NYPOIDEAE subtribe Species information Voucher SUBFAMILY Metroxylinae Mauritiinae Calameae K Salaccinae Pigafettinae Plectocomiinae Raphiinae Lepidocaryeae Ancistrophyllinae CALAMOIDEAE Eugeissoneae (in press) . ALAMOIDEAE oucher and database information (EMBL/GenBank/DDBJ databases) for the taxa used in this study. Herbarium acronyms are given in oucher and database information (EMBL/GenBank/DDBJ databases) for the taxa used in this study. lassification (1998) CORYPHOIDEAE NYPOIDEAE column gives the subfamily classification of Dransfield & Uhl (1998) and the second column gives the new subfamily, tribal and su column gives the subfamily classification of Dransfield & Uhl (1998) and the second column gives new subfamily, et al Subfamily c V C

© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 15–38 PALM FAMILY PHYLOGENY 29 K mat F trn L– trn 16 intron rps L rbc (FTG) AJ829884 AM116779 AM113624 AM114572 (HAJB) AJ829869 AM116772 AM113620 AM114559 (K) AJ404767 AJ240863 AJ241272 AM114565

(FTG) AM110192 AM116771 AM113619 AM114557

(K) AJ404752 AJ404925 AJ404892 AM114570

(FTG) AM110193 AM116773 AM113621 AM114560

(AAU) AJ404746 AJ240845 AJ241254 AM114562

(K)(K) AM110195 AM116776 AJ404754 AM113623 AM116777 AM114567 AJ241260 AM114568 (K) AM110194 AM116775 AM113622 AM114566 (FTG) AJ404748 AJ404923 AJ404890 AM114564 (FTG) AJ404749 AJ404924 AJ404891 AM114555

1092 (K) AJ404747 AJ240846 AJ241255 AM114563 − Roncal 043

.E. Lewis 02–080 .E. .E. Lewis 02–072 .E. 013–6701301 (K)C 1987–2573 (K) AJ404753 AM116778 AJ241259 AM114571 AJ404756 AJ240853 AJ241262 AM114573 Barrow 77 Barrow 76 1984–4470 (K)Dransﬁeld s.n. AJ404755 AJ240852 AJ241261 AM114569 1991 Baker 990 3928 al. Goyder et Barrow 75 Balslev 6407 J. 1938–27003 (K) Sanders 1763 R. 1973–12608 (K) AJ404745 AJ240844C AJ241253 AM114556 AJ404751 AJ240848 AJ241257 AM114558 1933–047031 (K) AJ404750 AM116774 AJ241256 AM114561 Baker 995 L.

L. (Desc.) acq.

Dammer Bartlett Burret (S.K.Lee J

H.Wendl.

Lodd. ex Lodd.

H.E.Moore

Henry ex

ei & J.Dransf. W N. & F.N.Wei) S.K.Lee, & F.N.Wei) F. (Pursh) H.Wendl. & Drude (Pursh) H.Wendl. (Ridley) Furtado Rehder (Hook.) H.Wendl. Chabaud warscewicziana (Burmeist.) Drude & Griseb. L.H.Bailey L.H.Bailey (Jacq.) & H.Wendl.) (Griseb. f. Hook. Schult. & Schult.f. Schult. hrinax radiata hrinax morrisii rachycarpus fortunei rachycarpus rithrinax campestris Guihaia argyrata Maxburretia rupicola Rhapis excelsa Rhapidophyllum hystrix T Phoenix dactylifera Chamaerops humilis Phoenix reclinata Phoenix canariensis Cryosophila Itaya amicorum T T Zombia antillarum Coccothrinax argentata Hemithrinax compacta T Chelyocarpus ulei Chelyocarpus Schippia concolor Schippia , tribe , subtribe Species information Voucher SUBFAMILY Livistoneae Rhapidinae Phoeniceae Cryosophileae lassiﬁcation (1998) Subfamily c

© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 15–38 30 C. B. ASMUSSEN ET AL. K mat F trn L– trn 16 intron rps L rbc (FTG) AM110201 AM116789 AM113633 AM114586 (FTG) AM110198(FTG) AM116783 AM113628 AM110199 AM114580 AM116785 AM113630 AM114582

(FTG) AM110197 AM116782 AM113627 AM114579 (KEP) AJ829894 AM116780 AM113625 AM114577

(FTG) AJ829862 AM116784 AM113629 AM114581 (P) AM110196 AM116781 AM113626 AM114578

(K) AJ829905 AM116786 AM113631 AM114583

(FTG) AJ404789 AJ404929 AJ404896 AM114593 (FTG) AJ404791 AJ404930 AJ404897 AM114591

Continued Bogler 1247 Chee 47332

.E. Lewis 03–013 .E. .E. Lewis 03–011 .E. Lewis 03–012 .E. 085–85.01492 (K)Baker 987 AJ404788 AJ240882 AJ241291 AM114592 Baker 989 R. Racine 41 R. 1952–35501 (K)1958–66102 (K)C AM110200 AM116787 AJ404760 AM113632 AM116788 AM114584 AJ241266 AM114585 C C Baker 1183 1987–2685 (K)1990–1132 (K) AJ404765 AJ240861 AJ404763 AJ241270 AJ240859 AM114588 AJ241268 AM114589 D. Pintaud 315 B. 1982–5601 (K)1985–1515 (K) AJ404757 AJ240854 AJ404758 AJ241263 AJ240855 AM114574 AJ241264 AM114576 Appendix H.

Becc. Burret 1986–3018 (K) AJ404764 AJ240860 AJ241269 AM114587

Seem. Becc. Dowe

(Jacq.) .Anders. L-80.0770 (BH) AJ404792 AJ240884 AJ241293 AM114594 Bartlett (Becc.)

Becc. 1985–1497 (K) AJ404759 AJ240856 AJ241265 AM114575

(Bartram) Becc.

Lour. 464–85.05037 (K) AJ404790 AJ240883 AJ241292 AM114590

(Reichb.f. & (Reichb.f.

endl. .K.Small .Dransf. Whitmore W & H.Wendl. J H.Wendl. (Mill.) H.E.Moore J H.Wendl. H.Wendl. Becc. R.Br. ex Mart. R.Br. altifrons Zoll.) H.E.Moore macrocarpus allichia disticha allichia ashingtonia robusta ohannesteijsmannia W Arenga hookeriana Arenga undulatifolia Serenoa repens W Caryota ophiopellis Copernicia prunifera paciﬁca Pritchardia Kerriodoxa elegans Colpothrinax wrightii arecina Pritchardia Chuniophoenix nana Nannorrhops ritchiana Caryota mitis Brahea berlandieri Acoelorraphe wrightii Pritchardiopsis jeanneneyi Pritchardiopsis Livistona chinensis Licuala kunstleri J Pholidocarpus , tribe , Livistoneae subtribe Species information Voucher SUBFAMILY Chuniophoeniceae Caryoteae Unplaced genera in Livistoninae lassiﬁcation (1998) Subfamily c ARECOIDEAE

© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 15–38 PALM FAMILY PHYLOGENY 31 K mat F trn L– trn 16 intron rps L rbc AM110202 AM116793 AM113637 AM114604 AJ404771 AJ240866 AJ241275 AM114598

(K) AJ829852 AM116790 AM113634 AM114597

(FTG) AF829878 AM116792 AM113636 AM114601

(FTG) AJ404780 AJ404928 AJ404895 AM114606

(AAU) AJ829874 AM116794 AM113638 AM114608 (FTG) AJ829885 AM116791 AM113635 AM114600 (FTG) AJ404762 AJ404926 AJ404893 AM114596

(K)

rénel 4 Roncal 64 T 1160 (K) ilkin, Suddee & Thapyai Thapyai Suddee & ilkin,

1988–227 (K) AJ404782 AJ240876 AJ241285 AM114609 1994–3231 (K)1989–3394 (K)W AJ404769 AJ240864 AJ404768 AJ241273 AJ404927 AM114602 AJ404894 AM114603 P. J. FTG 82–441C (BH)Baker 1002 AJ404779 AJ240874 AJ241283 AM114605 1976–1160 (K) AJ404781 AJ240875 AJ241284 AM114607 Baker 986 Beentje & Dransﬁeld 4810 Baker 984 43 Bayton R.P. L. 1954–35301 (K) AJ404761 AJ240858 AJ241267 AM114595

Lem.

L. Mart. 1994–3803 (K) AJ404770 AJ240865 AJ241274 AM114599

uert. ex uert. Drude ex Beentje 1988–2369 (K) AJ404783 AJ240877 AJ241286 AM114610

Roxb.

(Ridl.) Becc.

(F.M.Bailey) J.Dransf., (F.M.Bailey) A.K.Irvine & N.W.Uhl (J.F.Gmel.) Pers. (J.F.Gmel.) machadonis Hook.f. H.Wendl. (Mart.) Becc. (H.Karst) H.Wendl. H.Wendl. Beentje & J.Dransf. Hildebr. & H.Wendl. Hildebr. uania australis Oraniopsis appendiculata Ravenea louvelii Lodoicea maldivica Borassodendron Borassus ﬂabellifer Pseudophoenix sargentii Pseudophoenix vinifera Medemia argun Ceroxylon quindiuense J Latania verschaffeltii Satranala decussilvae Hyphaene thebaica Corypha umbraculifera Corypha taliera nobilis Bismarckia , tribe , SUBFAMILY subtribe Species information Voucher CEROXYLOIDEAE Cyclospatheae Ceroxyleae Lataniinae Borasseae Hyphaeninae lassiﬁcation (1998) CEROXYLOIDEAE Subfamily c CORYPHOIDEAE Corypheae

© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 15–38 32 C. B. ASMUSSEN ET AL. K mat F trn L– trn 16 intron rps L rbc AJ404786 AJ240880 AJ241787 AM114622 AM110203 AM116795 AM113639 AM114615 (K) AM110208 AM116799 AM113643 AM114626

(BH) AJ404793 AJ240885 AJ241294 AM114617

(CP) AM110204 AM116796 AM113640 AM114616 (UNMSM) (BH) AM110207 AJ240886 AJ241295 AM114625

(FTG) AJ404794 AJ404931 AJ404898 AM114619 (FTG) AM110205 AM116797 AM113641 AM114618 (K) AJ404837 AJ404954 AJ404921 AM114612

(FTG) AM110206 AM116798 AM113642 AM114621

(AAU) 94 (K) AJ404835 AJ240908 AJ241317 AM114613

egas 728 − V Continued 640 C. .Millán, J.C.Pintaud, J.C.Pintaud, .Millán, Sunderland 1803 Reitsma 2840 BH 60–811 (BH)1958–80101 (K) AJ404787 AJ240881 AJ404784 AJ241290 AJ240878 AM114623 AJ241287 AM114624 Knudsen & Asmussen Knudsen & Baker 991 1983–674 (K)Zona 754 AJ404785 AJ240879 AJ241288 AM114620 Baker 992 Henderson 042 Asmussen 111 B 1993 1992–2480 (K) AJ404836 AJ240907 AJ241316 AM114614 L-77.08309 (BH)Baker 985 AJ404838 AJ404955 AJ404922 AM114611 Appendix Burret .Mann .Mann

G G (A.J.

Ruiz &

(Mart.)

Burret

(O.F.Cook) polyclada

.Cook F & H.Wendl. & H.Wendl. Burret & Read H.J.Quero warscewiczianus H.Wendl. (L.H.Bailey) H.E.Moore var. Damm. (Burret) A.Henderson H.Wendl. Pav. H.Wendl. lamarckianum Spruce Ruiz & Pav. Henderson & Balslev) Barfod O. endlandiella gracilis ettinia hirsuta odococcus barteri odococcus barteri P Gaussia maya P Chamaedorea microspadix Synechanthus Hyophorbe lagenicaulis W W Socratea exorrhiza Iriartea deltoidea Dictyocaryum Phytelephas aequatorialis Iriartella stenocarpa Phytelephas macrocarpa Aphandra natalia Ammandra decasperma , tribe , odococceae subtribe Species information Voucher SUBFAMILY P Chamaedoreeae ARECOIDEAE Iriarteeae Phytelepheae lassiﬁcation (1998) ARECOIDEAE CEROXYLOIDEAE Subfamily c ARECOIDEAE PHYTELEPHANT- OIDEAE

© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 15–38 PALM FAMILY PHYLOGENY 33 K mat F trn L– trn 16 intron rps L rbc AJ404829 AJ240903 AJ241312 AM114636 (K) AJ404823 AJ404948 AJ404915 AM114629

(NH) AJ829876 AM116801 AM113645 AM114633

(K) AM110210 AM116802 AM113646 AM114634 (K) AM110209 AM116800 AM113644 AM114628

(FTG) AM110214 AM116806 AM113650 AM114642 (FTG) AJ404797 AJ240888 AJ241297 AM114645

(AAU) AJ404831 AJ404953 AJ404920 AM114641

(AAU) AJ404827 AJ240901 AJ241310 AM114638 (FTG) AM110213 AM116805 AM113649 AM114640

(FTG) AM110212 AM116804 AM113648 AM114639

(FTG) AM110215 AM116807 AM113651 AM114643

(AAU)

Sikhakhane 139

Roncal 79 612 B.

.E. Lewis 02–027 .E. .E. Lewis 03–010 .E. J. 599 Borchsenius C Zona 620 C Balslev 6404 Baker 1000 Dransﬁeld JD 6389 BH 79.312 (BH)Asmussen Knudsen & AJ404828 AJ240902 AJ241311 AM114635 T. 1989–3532 (K) AJ404826 AJ404951 AJ404918 AM114632 1963–57401 (K)1988–366 (K) AJ404805 AJ404936 AJ404799 AJ404903 AJ404933 AM114630 AJ404900 AM114631 Dransﬁeld JD 7731 Sunderland TCHS 1794 L-78.0662 (BH) AJ404796 AJ240887 AJ241296 AM114627 um. um. illd. (Jacq.) J (Jacq.)

J. acq. 1987–216 (K) AJ404830 AJ404952 AJ404919 AM114644 W

Becc. J (Kunth)

Kunth

H.Beentje L. 1968–4480 (K) AM110211 AM116803 AM113647 AM114637 H.E.Moore

.Cook .Baker ex Becc. F F Mart. Gaertn. C. (H.E.Moore) Glassm. ex Mart. Lodd. Dransf. (Gomes) Kuntze et H.Perrier (H.Wendl.) Drude ex (H.Wendl.) Dammer madagascariensis O. H.Wendl. Becc. oanioala gerardii ubaeopsis caffra Aiphanes aculeata Bactris gasipaes Desmoncus orthacanthos Elaeis guineensis Manicaria saccifera Acrocomia crispa Syagrus smithii Acrocomia aculeata V Cocos nucifera Allagoptera arenaria Attalea allenii J Reinhardtia simplex Beccariophoenix Sclerosperma mannii Roystonea oleracea Orania lauterbachiana Orania ravaka , tribe , subtribe Species information Voucher SUBFAMILY Elaeidinae Manicarieae Bactridinae Reinhardtieae Cocoseae Attaleinae Sclerospermeae Roystoneae Oranieae lassiﬁcation (1998) Subfamily c

© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 15–38 34 C. B. ASMUSSEN ET AL. K mat F trn L– trn 16 intron rps L rbc AM110218 AM116811 AM113655 AM114652 AM110217 AM116810 AM113654 AM114651 AJ829917 AM116809 AM113653 AM114650 AM110216 AM116808 AM113652 AM114648 (FTG) AJ829847 AM116814 AM113658 AM114659

(AAU) AM110219 AJ240906 AJ241315 AM114655

(VEN) AJ404798 AJ404932 AJ404899 AM114656 (AAU) AJ404804 AJ240891 AJ241300 AM114646 (TL)(K) NY)(K, AJ404806 NY) NOU, (K, AJ404937 AM110221 AJ404809 AM110222 AJ404904 AM116815 AJ240892 AM116816 AM114660 AM113659 AM113660 AJ241788 AM114661 AM114662 AM114663

(AAU) (AAU) (AAU) (AAU) 2935 (K) AJ829892 AM116812 AM113656 AM114657

− Continued 627 613 607 619 .E. Lewis 97–011 .E. Romero 3060 1992–3477 (K)C Pintaud 492 AM110220 AM116813Pintaud 351 AM113657Pintaud 361 AM114658 Pintaud 358 Knudsen & Asmussen Knudsen & FTG 71375 D (FTG)L-81.0284 (BH) AJ404832 348 Borchsenius AJ240904 AJ241313 AJ404833 AM114653 AJ240905 AJ241314 AM114654 Knudsen & Asmussen Knudsen & Knudsen & Asmussen Knudsen & L-81.0303 (BH) AJ404803 AJ240890 AJ241299 AM114649 Knudsen & Asmussen Knudsen & Balslev 6421 Appendix Becc. 1988

Mart.

(Linden

Mart. L-70.0017 (BH) AJ404802 AJ240889 AJ241298 AM114647 H.E.Moore

H. Wendl. H.

Becc. & Drude H.Wendl. Hodel & Dowe (Brongn.) Dammer ex Becc. Vieill. (Brongn. & Gris) Brongn. (Brongn. & H.Wendl.) H.Wendl. & H.Wendl.) H.E.Moore (Sw.) H.Wendl. (H.Wendl.) ex Drude ex Spruce H.Wendl. H.Wendl. ex Burret H.Wendl. ghiesbreghtiana Dammer Burret elﬁa regia elagodoxa henryana Actinorhytis calapparia purpurea Archontophoenix Actinokentia divaricata Chambeyronia macrocarpa Kentiopsis oliviformis Sommieria leucophylla P Calyptronoma occidentalis Calyptronoma Asterogyne martiana Geonoma congesta Leopoldinia pulchra Calyptrogyne Pholidostachys pulchra Pholidostachys W Neonicholsonia watsonii Neonicholsonia Prestoea pubens Euterpe oleracea Hyospathe macrorhachis , tribe , elagodoxeae subtribe Species information Voucher SUBFAMILY Areceae Archontophoenicinae P Leopoldinieae Geonomateae Euterpeae lassiﬁcation (1998) Subfamily c

© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 15–38 PALM FAMILY PHYLOGENY 35 K mat F trn L– trn 16 intron rps L rbc AJ829888 AM116826 AM113669 AM114675 AJ404821 AM116820 AJ241309 AM114669 (K, NOU, NY, NY, NOU, (K, (K)(P)(TL) AJ829849 AM116817 AM110223 AM110224 AM113661 AM116818 AM116819 AM114666 AM113662 AM113663 AM114667 AM114668 (K)(TL)(TL) AM110227 AM116825(P) AJ829864 AM113668 AM114674 (P) AM116827 AM110228 AM113670 AM116828 NY)(K, AM114676 AJ829879 AM113671 AM114677 AM116829 AJ829854 AJ829861 AM113672 AM116830 AM116831 AM114678 AM113673 AM113674 AM114679 AM114680 (TL)(TL) AM110225(TL) AM116821 AM113664 AJ829896 AM114670 AM116822 AM110226 AM113665 AM116823 AM114671 AM113666 AM114672

(FTG) AJ404818 AJ404944 AJ404911 AM114665

(FTG) AJ829859 AM116824 AM113667 AM114673

P) Baker 994 Pintaud 468 Pintaud 365 Pintaud 483 Pintaud 372 Zona 722 Pintaud 446 Roncal 73 Pintaud 482 Pintaud 469 Pintaud 364 Pintaud 368 Pintaud 349 Pintaud 491 Pintaud 452 Pintaud 470 Becc. Becc.

H.Wendl.

(Blume) Roxb. 1984–2295 (K) AJ404819 AJ404945 AJ404912 AM114664

(Mart.)

H.E.Moore

ernando F H.E.Moore (Becc.) ex (Brongn.) H.Wendl. Becc. H.E.Moore & Drude (Hatusima) H.E.Moore Brongn. H.E.Moore H.E.Moore lanuginosa (Brongn.) Becc. H.Wendl. var. var. H.Wendl. pachystachys (Brongn.) H.Wendl. (Brongn.) H.Wendl. ex Salomon H.E.Moore macrospermum eillonia alba Alloschmidia glabrata Alloschmidia Basselinia velutina Campecarpus fulcitus Cyphophoenix nucele Satakentia liukiuensis storckii Neoveitchia Cyphokentia macrostachya Moratia cerifera Lavoixia macrocarpa Brongniartikentia Clinosperma bracteale Nenga pumila Physokentia rosea V Carpoxylon Cyphosperma balansae Areca triandra , tribe , subtribe Species information Voucher SUBFAMILY Basseliniinae Clinospermatinae Carpoxylinae Arecinae lassiﬁcation (1998) Subfamily c

(BH)(BH) AM110234 AM116836 AJ829914 AM113679 AM116837 AM114691 AM113680 AM114692

(FTG) AM110233 AM116835 AM113678 AM114690

(FTG) AJ404813 AJ404942 AJ404909 AM114696 (K) AM110232 AM116834 AM113677 AM114687 (K)(K) AM110230 AM116832 AM113675 AM114684 AM110231 AM116833 AM113676 AM114686 (K) AM110229 AJ404935 AJ404902 AM114682

(K) AJ829906 AM116842 AM113685 AM114700 (FTG) AJ829858 AM116840 AM113683 AM114697 (K) AJ404825 AJ404950 AJ404917 AM114683

(FTG) AM110235 AM116838 AM113681 AM114693 (FTG) AJ829903 AM116839 AM113682 AM114694

Continued .E. Lewis 98–067 .E. Lewis 98–070 .E. redriksen et al. C-210 al. redriksen et L-72.0031 (BH)Baker 572 AJ404815 AJ240897 AJ241306 AM114699 FTG 85–11113 (FTG) AM110236 AM116841 AM113684 AM114698 Baker 999 Baker 1003 F L-79.0850 (BH) Sanders 1768 R. C AJ404812C AJ240895 AJ241304 AM114689 Baker 1109 Zona 869 Baker 1016 1992–3552 (K)Baker 1038 AJ404824 AJ404949 AJ404916 AM114685 Zona 878 1978–1196 (K)Baker 1008 Baker 998 AJ404800 AJ404934 AJ404901 AM114681 Appendix Becc. L-69.0404 (BH) AJ404814 AJ240896 AJ241305 AM114695

(H.Wendl.) Becc.

Becc.

Beentje (Becc.) Essig (Becc.)

um. .Dransf. & N.Uhl .Dransf. (Scheff.) Becc. (Scheff.) A.K.Irvine schumannii (H.Wendl. & Drude) Becc. (H.Wendl. (Mart.) H. Wendl. (Mart.) H. & Drude H.Wendl. Ridl. (Jack) (Bory) H.Wendl. H.E.Moore (H.Wendl. ex H.J.Veitch) (H.Wendl. f. ex Hook. H.Wendl. Humbert J Becc. albertisianus Beentje & J.Dransf. J J odyetia bifurcata eitchia arecina eitchia onapea ledermanniana ectiphiala ferox Ptychococcus paradoxus Ptychococcus W Brassiophoenix Carpentaria arcuminata V Laccospadix australasica Oncosperma tigillarium Acanthophoenix rubra T Linospadix monostachya Ptychosperma macarthurii Ptychosperma P Masoala madagascariensis Marojejya insignis Masoala kona Balaka seemannii Calyptrocalyx Marojejya darianii Dypsis lutescens Lemurophoenix halleuxii , tribe , subtribe Species information Voucher SUBFAMILY Oncospermatinae Ptychospermatinae Linospadicinae Dypsidinae lassiﬁcation (1998) Subfamily c

© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 15–38 PALM FAMILY PHYLOGENY 37 K mat F trn L– trn 16 intron rps L rbc (FTG) AM110244 AM116852 AM113695 AM114717 (GUAM) AM110243 AM116848 AM113691 AM114710 (BH) AM110241 AM116846 AM113689 AM114708

(FTG) AM110242 AM116847 AM113690 AM114709

(TL) AJ829850 AM116849 AM113692 AM114711 (NY)(TL) AJ404808 AJ404939 AJ404807 AJ404906 AJ404938 AM114701 AJ404905(TL) AM114702 (MAK) AM110239 AM116844 AM110240 AM116845 AM113687 AM114705 AM113688 AM114706

(K) AJ829881 AM116850 AM113693 AM114715

(K) AJ404817 AJ404943 AJ404910 AM114712

1825 (K) AM110238 AJ404947 AJ404914 AM114704 − .E. Lewis 99–044 .E. .E. Lewis 99–034 .E. .E. Lewis 98–061 .E. Lewis s.n. .E. 1990–2497 (K)C AJ829882 AM116851 AM113694 AM114716 L-72.0359 (BH)1985–1488 (K)Baker 1167 AJ404816 AJ240898 AJ404820 AJ241307 AJ404946 AM114713 AJ404913 AM114714 C Pintaud 461 Baker 573 Pintaud 384 Pintaud 407 1986–2346 (K)1985 Pintaud 457 AM110237 AM116843Pintaud 442 AM113686 AM114703 C C Blume 1982–5882 (K) AJ404810 AJ404940 AJ404907 AM114707 (Becc.)

(K.Koch)

baueri .Essig & B.E.Young endl. & Drude endl. B H.Wendl. (Kurz) H.E.Moore (Burret) H.E.Moore (Mart.) Bentham et ex Becc. Hook.f. O.F.Cook (F.Muell.) (H.E.Moore) Norup Burret (Becc.) H.Wendl. & Drude H.Wendl. var. H. & F.Muell.) (C.Moore W borsigianum Stuntz H.Wendl. (H.Wendl.) Becc. (Rehder & E.H.Wilson) H.E.Moore & Fosberg & Drude H.Wendl. Baker & Zona W.J. (Becc.) F. Rhopaloblaste augusta Loxococcus rupicola Hydriastele chaunostachys Iguanura wallichiana mooreana Lepidorrhachis Heterospathe longipes Hydriastele microspadix Rhopalostylis baueri Hedyscepe canterburyana Phoenicophorium melanochaetes Roscheria Clinostigma savoryanum renda Cyrtostachys Dictyosperma album Dransﬁeldia micrantha Heterospathe elata Bentinckia nicobarica Bentinckia , tribe , Areceae erschaffeltiinae subtribe Species information Voucher SUBFAMILY V Rhopalostylidinae Unplaced genera in lassiﬁcation (1998) Subfamily c

© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 15–38 38 C. B. ASMUSSEN ET AL. K mat F trn L– trn 16 intron rps L rbc (CP)(CP) AM110253 AM110254 AM116861 AM116862 AM113704 AM114726 AM113705 AM114727 (CP) AM110252 AM116860 AM113703 AM114725 (CP)(CP)(CP) AM110249 AM110250 AM116857 AM110251 AM116858 AM113700 AM116859 AM114722 AM113701 AM114723 AM113702 AM114724 (CP) AM110248 AM116856 AM113699 AM114721

(K)(K) AM110245 AM110246 AM116853 AM116854 AM113696 AM114718 AM113697 AM114719

Continued Asmussen 102 Asmussen 103 Asmussen 101 Asmussen 105 Asmussen 107 Asmussen 104 Chase 2229 1998–1475 (K)Asmussen 109 AM110247 AM116855 AM113698 AM114720 Chase 2230 Appendix L.

Lindl. -Gawl.

R.Br.

Ker

Baker

sp.

. .Don (Rose) D.R.Hunt R.Br. Merr D radescantia pallida ypha angustifolia argesia T Vriesia psittacina T Canna edulis Musa rosea F Dasypogon bromelifolius Hanguana malayana Anigozanthos manglesii Kingia australis , tribe , oaceae subtribe Species information Voucher SUBFAMILY Commelinaceae Bromeliaceae Typhaceae Cannaceae Musaceae Dasypogonaceae Hanguanaceae P Haemodoraceae Outgroups (family) Dasypogonaceae lassiﬁcation (1998) Subfamily c