Phylogeny of the Gardenieae (Rubiaceae)

The phylogeny of Gardenieae (Rubiaceae) is evaluated using 70 morphological and anatomical characters and 81 terminal taxa. After successive reweighting of the characters and modified bootstrap tests the following is concluded. The position of the Diplosporinae genera Argocoffeopsis, Calycosiphonia, Cremaspora and Tricalysia with the genera belonging to the tribes Aulacocalyceae, Coffeeae, Octotropideae and Pavetteae, suggests that this subtribe does not belong in Gardenieae. Posoqueria is nested among outgroup genera and consequently is not a member of the tribe. Robbrecht & Puff's informal 'tetrad group' is monophyletic. Likewise, their 'Alibertia group' is supported as monophyletic, although slightly rearranged. On the other hand, their 'Aidia group' must he greatly expanded with several genera in order to be monophyletic, but the monophyly of this group is weakly supported. Several disputed genera should be included in Gardenieae ( Anomanthodia, Amaioua, Brachytome , and supposedly Bertiera ). Most inferred groups, however, exhibit low bootstrap values and should he viewed with caution.     

Morphology and ultrastructure of orbicules in the subfamily Ixoroideae (Rubiaceae)

Orbicules were studied in 43 species belonging to 32 genera of the five tribes of the Ixoroideae (Rubiaceae). The orbicules were investigated with scanning electron microscopy (SEM), light microscopy (LM), and transmission electron microscopy (TEM). Orbicules are present in all species investigated of the tribes Pavetteae, Octotropideae and Coffeeae. In the tribes Gardenieae and Aulacocalyceae orbicules were found in some species, while they were absent in others. 15 species out of 11 genera lack orbicules. Three orbicule types (III, V, and VI) could be distinguished, mainly on the basis of general morphological and ultrastructural variations. Orbicules that belong to type III (0.50-1.29μm) have perforations in their wall, a regular or irregular shape and two or three electron transparent cores. This orbicule type, exclusively found in all Pavetteae species investigated, can be divided into two subtypes. Orbicules of subtype IIIa are present in all genera related to Ixora, and orbicules of subtype IIIb in those genera related to Tarenna. Orbicules of type V (0.97-1.86μm) are present in Himalrandia tetrasperma (Aulacocalyceae), and in all Octotropideae genera investigated, except Feretia. Complexes of more than three individual orbicules characterize this type. They are irregularly shaped and have many perforations as well as sporopollenin granules on the orbicule wall. In all species investigated of tribe Coffeeae, type VI orbicules (0.56-1.60μm) are present. These orbicules are characterized by their embedded position towards the tapetal membrane, their aggregated form and by the presence of perforations as well as sporopollenin granules on their orbicule wall. In the tribe Gardenieae different types of orbicule were found (V, VI and orbicules that cannot be classified in our typology). Our results suggest that orbicule characters in the Ixoroideae may be systematically useful on tribal level.     

Pollen morphology supports the transfer of Wendlandia (Rubiaceae) out of Rondeletieae

Pollen morphology of 27 species, eight subspecies and one variety of Wendlandia was studied using scanning electron microscopy (SEM). Wendlandia pollen are monads, radiosymmetric, small in size, tricolporate (rarely tetracolporate or bicolporate) and spheroidal (rarely subprolate or suboblate) in equatorial view. The compound aperture consists of ectocolpus, mesoporus and endocolpus. In addition, reticulate sexine and granular nexine were observed. The pollen wall ultrastructure of two Wendlandia spp. was examined by transmission electron microscopy (TEM). The exine consists of the tectum, columellae, foot layer and endexine. The endexine is thickened into a costa around the aperture. The intine forms a protruding oncus at the aperture. The palynological characters show a remarkable uniformity among the Wendlandia spp. Differences with Rondeletia, the main genus of tribe Rondeletieae, exist in the exine pattern, the endoaperture and the pollen wall structure. Our observations indicated that the endoaperture type and the structure of the pollen wall of Wendlandia were similar to those of the Gardenieae–Pavetteae–Coffeeae–Octotropideae clade, which provided palynological evidence for a closer relationship of Wendlandia to subfamily Ixoroideae and the transfer of Wendlandia out of Rondeletieae. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 128–141.     

Molecular phylogeny and taxonomic revision of the Philippine endemic Villaria Rolfe (Rubiaceae)

The little known Rubiaceae genus Villaria is endemic mostly to the coastal forests of the Philippines. Traditionally, it has been placed in the tribe Gardenieae. Later it was transferred to Octotropideae sensu Robbrecht and Puff. Villaria was placed among the ??primitive?? genera of the tribe, which are essentially characterized by large fruits, horizontal ovules and numerous seeds. Parsimony and Bayesian analyses of the combined plastid (rps16 and trnT-F) dataset strongly support the inclusion of Villaria in Octotropideae as well as monophyly of the genus. However, our molecular results do not conform to the current informal groups of the tribe delimited by fruit size, ovule position, number of seeds and exotesta thickenings. Instead, a close relationship between Villaria and two ??central genera?? (Hypobathrum and Pouchetia) is revealed for the first time. This clade is sister to a group comprising ??primitive?? (Fernelia), ??advanced?? (Kraussia and Polysphaeria) and ??central?? (Feretia) representatives. In addition, our combined tree strongly supports a sister taxa relationship between Canephora and Paragenipa. Villaria is characterized by unilocular ovaries, parietal placentation and strictly horizontal ovules. These features are unique within the Octotropideae. We recognize a total of five Villaria species, one new species (V. leytensis) is described here, and two species (V. philippinensis and V. rolfei) are transferred into synonymy with V. odorata. Each species is fully described, and a key to the species, a distribution map and illustrations are provided.     

Molecular systematics and morphological character evolution of the Condamineeae (Rubiaceae)

? Premise of the study: The Condamineeae have in previous molecular studies been shown to be part of an early-divergent clade within the subfamily Ixoroideae, together with the tribes Calycophylleae, and Hippotideae, and genera of the former Cinchoneae and Rondeletieae. Generic relationships within this clade have, however, remained largely unresolved. ? Methods: In this study, the systematics of the Condamineeae was further examined by phylogenetic reconstruction of six cpDNA regions and one nrDNA region using parsimony and Bayesian Markov chain Monte Carlo inference. Morphological character evolution within the tribe was assessed by ancestral state reconstruction using likelihood optimization of characters onto Bayesian trees. ? Key results: Calycophylleae appears polyphyletic. "Hippotideae" is monophyletic but nested within the Condamineeae. The phylogenetic hypotheses presented support a resurrection of the genera Holtonia, Schizocalyx, and Semaphyllanthe. Furthermore, Bathysa is found to be polyphyletic, Tresanthera is found nested within Rustia, and the taxonomically disputed genus Dialypetalanthus is here shown to be sister to a Bothriospora-Wittmackanthus clade. Morphological ancestral state reconstructions indicate that protogyny have evolved at least two times within the tribe and that indehiscent fruits, loculicidal fruit dehiscence, and intrapetiolar stipules have evolved independently several times. The occurrence of calycophylls (leaf-like calyx lobes), poricidal anthers, and winged seeds also appear homoplastic within the tribe. ? Conclusions: A diagnosis and delimitation of the tribe Condamineeae is presented, with taxonomic proposals to synonymize Tresanthera and to transfer several species of Bathysa as well as Phitopis to a resurrected Schizocalyx.     

Pollen morphology and phylogeny of the Alyxieae (Apocynaceae)

The pollen of the tribe Alyxieae, including Alyxia , Chilocarpus , Condylocarpon , Lepinia , Lepiniopsis , Plectaneia and Pteralyxia , has been studied with light, scanning electron and transmission electron microscopy. It is characterised by relatively large, irregularly disposed and - unequal-sized pores. 2-porate grains are most common in the Alyxieae, and occur in all genera except for Condylocarpon . 1-, 3-, 4- and 5-porate pollen grains are less common. The orientation of the polar axis and the equatorial plane could not be determined in any of the genera. Within the Alyxieae, Chilocarpus and Plectaneia are the nearest to other Apocynaceae, showing an essentially similar exine ultrastructure. Alyxia , Lepinia , Lepiniopsis and Pteralyxia make up a derived group, characterised by large pollen grain size and a remarkable nexine structure. Pollen of Lepinia and Lepiniopsis might be derived on the basis of their fossulate-verrucate ornamentation. Condylocarpon pollen shows a deviating set of pollen features (tetrads, inaperturate, reduced exine) that suggest it to be neotenic. Pollen morphology supports preliminary results of cladistic analyses of chloroplast data, in which the tribe is a monophyletic group nested in a 3-colporate environment, Chilocarpus and Condylocarpon are sisters at the base of the tree, Plectaneia is sister to a clade made up by Alyxia , Pteralyxia , Lepinia and Lepiniopsis , of which Lepinia and Lepiniopsis are the most derived.     

Phylogenetic relationships of Moxostoma and Scartomyzon (Catostomidae) based on mitochondrial cytochrome b sequence data

A recent phylogenetic study based on morphological, biochemical and early life history characters resurrected the genus Scartomyzon (jumprock suckers, c . eight−10 species) from Moxostoma (redhorse suckers, c . 10–11 species) and advanced the understanding of relationships among species in these two genera, and the genealogical affinities of these genera with other evolutionary lineages within the tribe Moxostomatini in the subfamily Catostominae. To further examine phylogenetic relationships among moxostomatin suckers, the complete mitochondrial (mt) cytochrome b gene was sequenced from all species within this tribe and representative outgroup taxa from the Catostomini and other catostomid subfamilies. Phylogenetic analysis of gene sequences yielded two monophyletic clades within Catostominae: Catostomus + Deltistes + Xyrauchen + Erimyzon + Minytrema and Moxostoma + Scartomyzon + Hypentelium + Thoburnia . Within the Moxostomatini, Thoburnia was either unresolved or polyphyletic; Thoburnia atripinnis was sister to a monophyletic Hypentelium . In turn, this clade was sister to a monophyletic clade containing Scartomyzon and Moxostoma . Scartomyzon was never resolved as monophyletic, but was always recovered as a polyphyletic group embedded within Moxostoma , rendering the latter genus paraphyletic if ' Scartomyzon ' continues to be recognized. Relationships among lineages within the Moxostoma and' Scartomyzon 'clade were resolved as a polytomy. To better reflect phylogenetic relationships resolved in this analysis, the following changes to the classification of the tribe Moxostomatini are proposed: subsumption of' Scartomyzon 'into Moxostoma ; restriction of the tribe Moxostomatini to Moxostoma ; resurrect the tribe Erimyzonini, containing Erimyzon and Minytrema , classified as incertae sedis within Catostominae; retain the tribe Thoburniini.     

Aperture variation in the pollen of Nelumbo (Nelumbonaceae)

The evolutionary and developmental origin of tricolpate pollen is of great interest because pollen of this kind defines a major clade of angiosperms (eudicots), a clade that is also well supported by molecular data. We examined evidence that tricolpate and monosulcate pollen types are produced alongside each other in the anthers of Nelumbo flowers, as has previously been reported. Observations of pollen in situ within individual anthers revealed mainly tricolpate pollen produced in tetrahedral tetrads, but also a small percentage of clearly aberrant pollen grains that have a great variety of aperture configurations. Previously published evidence for tetragonal tetrads is not supported, and previously reported monosulcate grains are part of a continuum of variation among the aberrant grains in aperture number, position and form. Other eudicots show similar variability in their pollen apertures. The variation in the pollen of Nelumbo is not exceptional, and may not be more significant than variation seen in the other taxa with regard to the origin of the tricolpate and tricolpate‐derived pollen characteristic of eudicots. Nevertheless further studies of aberrant pollen in Nelumbo and other eudicots, together with comparisons of pollen development in “normal” eudicots and closely related species that show radical, and developmentally fixed, reorganization of apertures and pollen polarity, may be helpful in understanding the processes that controlled the transition from the monosulcate to the tricolpate condition.     

虎尾草亚科系统发育关系的初步研究

运用广义形态学性状对虎尾草亚科（Chloridoideae）进行系统发育分析。内类群包括虎尾草亚科52属的69种植物,代表虎尾草亚科的主要类群;芦竹亚科（Arundinoideae）扁芒草族（Danthonieae）的Centropodia和Danthonia被选作外类群。分支分析表明,虎尾草亚科是一个单系类群。其严格一致树包括A、B、C、D、E5个分支。两个大族画眉草族（Eragrostideae）和虎尾草族（Chlorideae）代表虎尾草亚科内部类群分化的两个方向,分开处理较合理。细穗草族（Leptureae）放到虎尾草族中较合理。冠芒草族（Pappophoreae）是虎尾草亚科的基部类群,与画眉草族近缘。我们的研究支持虎尾草亚科从旧世界向新世界扩散的地理分布假说,并提供了虎尾草亚科属上类群的系统发育关系的框架。     

Phylogenetic relationships of butterflies of the tribe Acraeini (Lepidoptera, Nymphalidae, Heliconiinae) and the evolution of host plant use

The tribe Acraeini (Nymphalidae, Heliconiinae) is believed to comprise between one and seven genera, with the greatest diversity in Africa. The genera Abananote, Altinote, and Actinote (s. str.) are distributed in the Neotropics, while the genera Acraea, Bematistes, Miyana, and Pardopsis have a Palaeotropical distribution. The monotypic Pardopsis use herbaceous plants of the family Violaceae, Acraea and Bematistes feed selectively on plants with cyanoglycosides belonging to many plant families, but preferentially to Passifloraceae, and all Neotropical species with a known life cycle feed on Asteraceae only. Here, a molecular phylogeny is proposed for the butterflies of the tribe Acraeini based on sequences of COI, EF-1alpha and wgl. Both Maximum Parsimony and Bayesian analyses showed that the tribe is monophyletic, once the genus Pardopsis is excluded, since it appears to be related to Argynnini. The existing genus Acraea is a paraphyletic group with regard to the South American genera, and the species of Acraea belonging to the group of "Old World Actinote" is the sister group of the Neotropical genera. The monophyly of South American clade is strongly supported, suggesting a single colonization event of South America. The New World Actinote (s. str.) is monophyletic, and sister to Abananote+Altinote (polyphyletic). Based on the present results it was possible to propose a scenario for the evolution in host plant use within Acraeini, mainly concerning the use of Asteraceae by the South American genera.     

Phylogeny and morphological evolution of tribe Menispermeae (Menispermaceae) inferred from chloroplast and nuclear sequences

The Menispermaceae family contains ca. 72 genera with 450 species that are almost entirely tropical. Its phylogeny at the tribal level has never been examined using molecular data. Here we used DNA sequences of the chloroplast matK gene and trnL-F regions, and the nuclear ITS region to study the delimitation and position of the tribe Menispermeae within the family and its subtribal monophyletic groups. Family-wide phylogenetic analyses of the chloroplast data produced two strongly supported clades. The first clade contains two subclades: Coscinieae including Arcangelisia and Anamirta, and Tinosporeae sensu lato including Fibraureae, supported by morphological characters, such as traits of the cotyledon, stylar scar and embryo. The second clade consists of the tribes Menispermeae sensu DC. and Tiliacoreae Miers. All our analyses surprisingly recognized that tribe Menispermeae is not monophyletic unless tribe Tiliacoreae is included, suggesting that characters of cotyledon and stylar scar are very important for the infrafamilial classification, and that endosperm presence vs. absence was over-emphasized in traditionally tribal division of the family. Our topologies indicate a secondary loss of endosperm. The monophyly of two subtribes of the tribe Menispermeae, Stephaniinae and Cissampelinae, is supported by the cpDNA and ITS data, as well as by morphological characters, including aperture types and shapes, and colpal membrane features of pollen grains, and sepal number of male flowers. The Cocculinae was recognized as a paraphyletic group containing the remaining genera of the tribe Menispermeae.     

Pollen and the Evolution of Arctotideae (Compositae)

Recent molecular studies have elucidated the phylogeny of Compositae tribe Arctotideae, and found it to contain two, well supported, monophyletic subtribes, Arctotidineae and Gorteriinae, as well as some polyphyletic and problematic genera. On the basis of this new information, it may now be possible to identify diagnostic characters and synapomorphies to support the groupings defined within Arctotideae. Pollen characters have been shown to be particularly variable in Compositae. This paper aims to investigate the utility of those characters in the context of recent molecular phylogenies, in order to determine synapomorphic and diagnostic characters in Arctotideae. The pollen of each genus is described, illustrated with scanning electron micrographs, and optimised on a phylogeny of the tribe. Many pollen characters were found to be very informative when considered in the context of the current best estimate of phylogenetic relationships. Pollen morphology provides synapomorphies for clades at a number of hierarchical levels within Arctotideae, including the two subtribes, Arctotidinae and Gorteriinae, the grouping of Eremothamnus and Hoplophyllum, and smaller clades. It also supports the exclusion of Platycarpha from the tribe. The plesiomorphic palynological state for the tribe is discussed. Particular attention is paid to the evolution of different patterns of lophae (surface ridges). A single origin for the lophate condition is proposed as the most parsimonious mode of evolution in Arctotideae.     

A new phylogenetic tribal classification of the grape family (Vitaceae)

Vitaceae (the grape family) consist of 16 genera and ca. 950 species primarily distributed in tropical regions. The family is well‐known for the economic importance of grapes, and is also ecologically significant with many species as dominant climbers in tropical and temperate forests. Recent phylogenetic and phylogenomic analyses of sequence data from all three genomes have supported five major clades within Vitaceae: (i) the clade of Ampelopsis, Nekemias, Rhoicissus, and Clematicissus; (ii) the Cissus clade; (iii) the clade of Cayratia, Causonis, Cyphostemma, Pseudocayratia, Tetrastigma, and an undescribed genus “Afrocayratia”; (iv) the clade of Parthenocissus and Yua; and (v) the grape genus Vitis and its close tropical relatives Ampelocissus, Pterisanthes and Nothocissus, with Nothocissus and Pterisanthes nested within Ampelocissus. Based on the phylogenetic and morphological (mostly inflorescence, floral and seed characters) evidence, the new classification places the 950 species and 16 genera into five tribes: (i) tribe Ampelopsideae J.Wen & Z.L.Nie, trib. nov. (47 species in four genera; Ampelopsis, Nekemias, Rhoicissus and Clematicissus); (ii) tribe Cisseae Rchb. (300 species in one genus; Cissus); (iii) tribe Cayratieae J.Wen & L.M.Lu, trib. nov. (370 species in seven genera; Cayratia, Causonis, “Afrocayratia”, Pseudocayratia, Acareosperma, Cyphostemma and Tetrastigma); (iv) tribe Parthenocisseae J.Wen & Z.D.Chen, trib. nov. (ca. 16 spp. in two genera; Parthenocissus and Yua); and (v) tribe Viteae Dumort. (ca. 190 species in two genera; Ampelocissus and Vitis).     

Identifying clades in Asian Annonaceae: monophyletic genera in the polyphyletic Miliuseae

The tribe Miliuseae (Annonaceae) comprises six genera distributed in Asia: Alphonsea, Mezzettia, Miliusa, Orophea, Platymitra, and Phoenicanthus. A phylogenetic study to investigate the putative monophyly of the tribe and the intergeneric relationships is presented here. Nucleotide sequences of the plastid gene rbcL, trnL intron, and trnL-trnF intergenic spacer were analyzed from 114 Annonaceae taxa, including 24 Miliuseae species and two outgroups using maximum parsimony and Bayesian inference. The two data sets (rbcL and the trnL-trnF regions) were analyzed separately and in combination. Miliuseae were found to be polyphyletic due to the position of Mezzettia and are part of a large, predominantly Asian and Central-American clade (miliusoid clade). Although intergeneric relationships were poorly resolved, all genera, except Polyalthia, were monophyletic, supporting previous generic delimitation based on morphology. A group of three Polyalthia species seems the most likely sister group of Miliusa. Several infrageneric groups of Miliusa, Orophea, and Polyalthia are supported by both molecular and morphological data. No morphological synapomorphies have yet been found for the miliusoid clade. Molecular clades within the miliusoid clade, however, can be characterized by size and the shape of the outer petals, number of ovules per carpel, and the size of the fruits.     

Phylogeny of the plant bug subfamily Mirinae (Hemiptera: Heteroptera: Cimicomorpha: Miridae) based on total evidence analysis

The first comprehensive phylogenetic analyses of the most diverse subfamily of plant bugs, Mirinae, is presented in this study, for 110 representative taxa based on total evidence analysis. A total of 85 morphological characters and 3898 bp of mitochondrial (16S, COI) and nuclear (18S, 28S) sequences were analysed for each partitioned and combined dataset based on parsimony, maximum likelihood and Bayesian inference. Major results obtained in this study include monophyly of the tribe Mecistoscelini. The largest tribe, Mirini, was recovered as polyphyletic, and Stenodemini was recovered as paraphyletic. The clade of Stenodemini + Mecistoscelini is the sister group of the remaining Mirinae. The monophyly of two complexes composed of superficially similar genera were tested; the Lygus complex was recovered as nonmonophyletic, and the Adelphocoris–Creontiades–Megacoelum complex was confirmed to be monophyletic. The generic relationships of the main clades within each tribe based on the phylogeny, as well as their supported morphological characters, are discussed.     

Phylogenetic analyses confirm polyphyly of the genus Campanula (Campanulaceae s. str.), leading to a proposal for generic reappraisal

Previous phylogenetic analyses found that the largest genus in the tribe Campanuleae, Campanula L. is polyphyletic. The genus is extremely intermingled, involving more than 50 genera, but no generic reappraisal has been attempted. For undertaking further phylogenetic analyses and subsequent generic reappraisal of the tribe, we sampled 333 samples, representing 27 of 28 genera currently recognized. Among them, 146 samples and two genera, Cylindrocarpa Regel and Sergia Fed., were newly sequenced. Six chloroplast DNA loci (atpB-rbcL, matK, petD-intron, rbcL, rpl16, and trnL-F) and internal transcribed spacer were used to undertake phylogenetic analyses. Our chloroplast DNA phylogeny comprises 24 clades, that is, 18 Cam clades and six genera, Feeria Buser, Homocodon D. Y. Hong, Jasione L., Peracarpa Hook. f. & Thomson, Trachelium L., and Favratia Feer. Campanula species are scattered among the 18 Cam clades and the six genera, and some of them join with well-established genera like Phyteuma L. and Adenophora Fisch. In the phylogeny Musschia Dumort. is at the basal position, but Jasione's position is unclear, whether in the tribe Campanuleae or in Wahlenbergieae; the other 22 lineages are grouped into two major clades: clade A comprising Cam 13–17 plus Feeria and Trachelium, and clade B comprising Cam 02, 03, 04-1, 04-2, and 06-12 plus Homocodon and Peracarpa. We found that the molecular phylogeny is closely correlated with morphology, particularly pollen morphology: clade A with pollen 3-porate and capsule dehiscent mostly by basal pores; and clade B with pollen mostly 4 (5–15)-porate and capsule dehiscent mostly by apical-middle pores. A generic reappraisal of the tribe is suggested based on the integrated phylogenetic analyses.     

Phylogenetic relationships of the downy mildews (Peronosporales) and related groups based on nuclear large subunit ribosomal DNA sequences

In order to investigate phylogenetic relationships of the Peronosporomycetes (Oomycetes), nuclear large subunit ribosomal DNA sequences containing the D1 and D2 region were analyzed of 92 species belonging to the orders Peronosporales, Pythiales, Leptomitales, Rhipidiales, Saprolegniales and Sclerosporales. The data were analyzed applying methods of neighbor-joining as well as maximum parsimony, both statistically supported using the bootstrap method. The results confirm the major division between the Pythiales and Peronosporales on the one hand and the Saprolegniales, Leptomitales, and Rhipidiales on the other. The Sclerosporales were shown to be polyphyletic; while Sclerosporaceae are nested within the Peronosporaceae, the Verrucalvaceae are merged within the Saprolegniales. Within the Peronosporomycetidae, Pythiales as well as Peronosporales as currently defined are polyphyletic. The well supported Albugo clade appears to be the most basal lineage, followed by a Pythium-Lagenidium clade. The third, highly supported clade comprises the Peronosporaceae together with Sclerospora, Phytophthora, and Peronophythora. Peronophythora is placed within Phytophthora, indicating that both genera should be merged. Bremiella seems to be polyphyletic within the genus Plasmopara, suggesting a transfer to Plasmopara. The species of Peronospora do not appear as a monophyletic group. Peronospora species growing on Brassicaceae form a highly supported clade.     

A phylogenetic analysis of tribe Areceae (Arecaceae) using two low-copy nuclear genes

 A phylogenetic study of the largest tribe of palms, the Areceae, was conducted using sequences of two low-copy nuclear genes. Previous morphological and plastid DNA studies have not supported the monophyly of the tribe, but have placed its members in a large clade that includes the monophyletic tribes Geonomeae, Cocoeae, Podococceae, and Hyophorbeae. We analyzed this large clade to test the monophyly of tribe Areceae with nuclear data, to explore relationships among its subtribes, and to identify other monophyletic groups. For 54 palm species, including members of all 17 subtribes of tribe Areceae, we sequenced regions of the malate synthase (MS) and phosphoribulokinase (PRK) genes. Simultaneous analysis of these regions revealed 52 shortest trees, all of which resolved tribe Areceae as polyphyletic. Subtribes Iguanurinae, Dypsidinae, Oncospermatinae, and Arecinae were also resolved as polyphyletic. A clade of Indo-Pacific taxa was resolved with strong support, and would be a suitable target for more focused study. Received February 7, 2001; accepted April 9, 2002 Published online: December 3, 2002