Infrageneric variability of pollen morphology in Palaua (Malveae,Malvaceae) and the taxonomic utility of quantitative pollen characters

Palaua (Malvaceae) comprises 15 species endemic to the coastal deserts of Chile and Peru. Previous molecular phylogenetic analyses have shown that this genus is monophyletic and can be subdivided into three clades. In the present study, pollen morphology of all species of Palaua was examined using light and scanning electron microscopy to determine whether it provides additional data in support of the proposed infrageneric groups. The pollen grains are spheroidal, medium to large in diameter, spinose and pantocolporate – the ectocolpi are very short. The tectum is perforate and characterised by granula. In all species the nexine is of a similar thickness in all areas of the pollen grain, while the sexine thickness varies, being notably thicker in the areas where the broad spine bases are sited (‘spine cushions’). Many of the quantitative characters have conspicuous variability, but the variability shows considerable overlap between species. Nonetheless, P. guentheri, P. inconspicua, P. malvifolia and P. modesta are united as a group by having the smallest pollen grains with the smallest apertures, the shortest spines and the shortest interspinal distance. This grouping reflects only partially the suggested infrageneric clades, although it does tend to unite the species with the smallest flowers. A possible link between reproductive resources and pollen size is also considered, as well as the influence of polyploidy. However, the taxonomic utility of these quantitative characters is weakened by species with more than one cytotype.     

New insights into the phylogenetics and biogeography of Arum (Araceae): unravelling its evolutionary history

The heat‐ and odour‐producing genus Arum (Araceae) has interested scientists for centuries. This long‐term interest has allowed a deep knowledge of some complex processes, such as the physiology and dynamics of its characteristic lure‐and‐trap pollination system, to be built up. However, mainly because of its large distributional range and high degree of morphological variation, species' limits and relationships are still under discussion. Today, the genus comprises 28 species subdivided into two subgenera, two sections and six subsections. In this study, the phylogeny of the genus is inferred on the basis of four plastid regions, and the evolution of several morphological characters is investigated. Our phylogenetic hypothesis is not in agreement with the current infrageneric classification of the genus and challenges the monophyly of several species. This demonstrates the need for a new infrageneric classification based on characters reflecting the evolution of this enigmatic genus. To investigate the biogeography of Arum deeply, further spatiotemporal analyses were performed, addressing the importance of the Mediterranean basin in the diversification of Arum. Our results suggest that its centre of origin was the European–Aegean region, and that major diversification happened during the last 10 Myr. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 14–32.     

Phylogeny of the spider genus Ixchela Huber, 2000 (Araneae: Pholcidae) based on morphological and molecular evidence (CO1 and 16S), with a hypothesized diversification in the Pleistocene

The genus Ixchela Huber is composed of 20 species distributed from north‐eastern Mexico to Central America, including the five new species described here from Mexico: I xchela azteca sp. nov. , I xchela jalisco sp. nov. , I xchela mendozai sp. nov. , I xchela purepecha sp. nov. and I xchela tlayuda sp. nov. We test the monophyly and investigate the phylogenetic relationships among species of the genus Ixchela using morphological and molecular data. Parsimony (PA) analysis of 24 taxa and 40 morphological characters with equal and implied weights supported the monophyly of Ixchela with eight morphological synapomorphies. The PA analyses with equal and implied weights, and separate Bayesian inference (BI) analyses for the CO1 gene (506 characters), concatenated gene fragments CO1 + 16S (885 characters), morphology + CO1 (546 characters) and the combined evidence data set (morphology + CO1 + 16S) (925 characters) support the monophyly of Ixchela. Our preferred topology shows two large clades; clade 1 has a natural distribution in the Mesoamerican biotic component, whereas clade 2 predominates in the Mexican Montane biotic component. The genus Ixchela diverged in the late Miocene, and the divergence between the internal clades in the genus occurred in the late Pliocene; by contrast, most of the speciation events seem to have occurred mainly during the Pleistocene, where climatic changes brought on by repeated glaciations played an important role in the diversification of the genus. © 2015 The Linnean Society of London     

Phylogeny in Echinocereus (Cactaceae) based on combined morphological and molecular evidence: taxonomic implications

Echinocereus is a morphologically diverse genus that includes 64 species grouped into eight taxonomic sections based on morphological traits. In previous molecular phylogenetic analyses, the relationships amongst Echinocereus species were not entirely revealed and useful characters to recognize clades were not provided. The inclusion of several sources of evidence in a phylogenetic analysis is likely to produce more supported hypotheses. Therefore, we performed a combined phylogenetic analysis with a set of 44 morphological characters and six chloroplast DNA sequences. Topologies from parsimony and Bayesian analyses were mostly congruent. However, the relationships of E. poselgeri were not consistent between analyses. A second Bayesian analysis using a long-branch extraction test resulted in a topology with the morphological position of E. poselgeri congruent with that in parsimony analysis. Parsimony and Bayesian analyses corroborated the monophyly of Echinocereus, which included eight monophyletic groups. The combined phylogeny integrated into different clades those taxa that were not determined in previous analyses and changed the relationships of some recognized clades. The clades did not recover the recent infrageneric classification. In the present study, a new sectional classification for Echinocereus is proposed based on the eight recovered clades, which is supported by a combination of morphological and molecular characters. An identification key for sections in the genus is included.     

A tale of North and South America: time and mode of dispersal of the amphitropical genus Munroa (Poaceae,Chloridoideae)

Plant disjunctions have provided some of the most intriguing distribution patterns historically addressed by biogeographers. We evaluated the three hypotheses that have been postulated to explain these patterns [vicariance, stepping‐stone dispersal and long‐distance dispersal (LDD)] using Munroa, an American genus of grasses with six species and a disjunct distribution between the desert regions of North and South America. The ages of clades, cytology, ancestral characters and areas of distribution were investigated in order to establish relationships among species, to determine the time of divergence of the genus and its main lineages, and to understand further the biogeographical and evolutionary history of this genus. Bayesian inference recovered the North American M. pulchella as sister species to the rest. Molecular dating and ancestral area analyses suggest that Munroa originated in North America in the late Miocene–Pliocene (7.2 Mya; 8.2–6.5 Mya). Based on these results, we postulate that two dispersal events modelled the current distribution patterns of Munroa: the first from North to South America (7.2 Mya; 8.2–6.5 Mya) and the second (1.8 Mya; 2–0.8 Mya) from South to North America. Arid conditions of the late Miocene–Pliocene in the Neogene and Quaternary climatic oscillations in North America and South America were probably advantageous for the establishment of populations of Munroa. We did not find any relationship between ploidy and dispersal events, and our ancestral character analyses suggest that shifts associated with dispersal and seedling establishment, such as habit, reproductive system, disarticulation of rachilla, and shape and texture of the glume, have been important in these species reaching new areas. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 110–125.     

Phylogeny of Impatiens (Balsaminaceae): integrating molecular and morphological evidence into a new classification

Impatiens L. is one of the largest angiosperm genera, containing over 1000 species, and is notorious for its taxonomic difficulty. Here, we present, to our knowledge, the most comprehensive phylogenetic analysis of the genus to date based on a total evidence approach. Forty‐six morphological characters, mainly obtained from our own investigations, are combined with sequence data from three genetic regions, including nuclear ribosomal ITS and plastid atpB‐rbcL and trnL‐F. We include 150 Impatiens species representing all clades recovered by previous phylogenetic analyses as well as three outgroups. Maximum‐parsimony and Bayesian inference methods were used to infer phylogenetic relationships. Our analyses concur with previous studies, but in most cases provide stronger support. Impatiens splits into two major clades. For the first time, we report that species with three‐colpate pollen and four carpels form a monophyletic group (clade I). Within clade II, seven well‐supported subclades are recognized. Within this phylogenetic framework, character evolution is reconstructed, and diagnostic morphological characters for different clades and subclades are identified and discussed. Based on both morphological and molecular evidence, a new classification outline is presented, in which Impatiens is divided into two subgenera, subgen. Clavicarpa and subgen. Impatiens; the latter is further subdivided into seven sections.     

Phenetic studies in Nuphar Sm. (Nymphaeaceae):variation in sect. Nuphar

Previous taxonomic assessments of the hydrophytic genus Nuphar Sm. (Nymphaeaceae) have inadequately considered the range of morphological variation existing in the genus. A comprehensive statistical analysis of morphological features among the recognized taxa of Nuphar is presented here, first using 10 fruit characters to delimit the separation of two infrageneric groups of taxa, and secondly using 31 characters to quantify the morphological variation found within sect. Nuphar. Results from combined phenetic analyses provided empirical support for recognition of two infrageneric groups of species in the genus (largely by fruit, stigmatic disk margin, neck, and stigmatic disk size) and for separation of taxa within sect. Nuphar (by leaf size, and fruit disk and neck size). Statistical analyses demonstrate that extensive variation in ranges of character states exists among sect. Nuphar taxa. However, a suite of qualitative and quantitative characters can be used to recognize five species and three subspecies within this predominantly Old World assemblage.     

Chloroplast DNA restriction site study of Verbesina (Asteraceae: Heliantheae)

Chloroplast DNA variation was examined for 79 species of Verbesina and 24 outgroups. Two independent analyses of the data were performed. An intergeneric study to elucidate the phylogenetic relationships of Verbesina used 22 genera, most of which are regarded in the literature as closely related to Verbesina. Coreopsis and Hymenoxys served as outgroups for this analysis. The 16 6-bp (base pair) restriction endonucleases used in the intergeneric study revealed 263 phylogenetically informative sites. Wagner analyses of these characters resulted in four equally parsimonious trees with a length of 857 steps and a Consistency Index of 0.492. Results from this study indicate that Verbesina is monophyletic, a member of the tribe Heliantheae, and that its sister taxa are the mostly Mexican genera Podachaenium, Squamopappus, and Tetrachyron. The infrageneric study of Verbesina included species belonging to all of its infrageneric taxa, except the monotypic sect. Stenocarpha. The 17 6-bp restriction endonucleases used in the infrageneric study revealed 137 sites 77 of which were phylogenetically informative. Wagner analyses of these characters generated 180 equally parsimonious trees with a length of 158 steps and a Consistency Index of 0.786. The genera Podachaenium, Squamopappus, and Tetrachyron served as outgroups. Two major clades, which correspond to traditional divisions of the genus based on leaf arrangement, are supported by the study. No support was found for the monophyly of seven of the 11 sections examined; further sampling of sections Lipactinia, Ochractinia, Verbesina, and Ximenesia is needed. Results support a major reassessment of the traditional infrageneric classification of the genus and provide the basis for the reevaluation of the sectional taxonomy of Verbesina. These studies support a North American origin for the genus with several independent introductions into South America producing significant diversity, especially in the Andean region.     

An updated genus‐wide phylogenetic analysis of Arisaema (Araceae) with reference to sections

Arisaema has a long and complicated taxonomic history regarding its infrageneric classification. In the latest system, 14 sections were recognized, based on an unpublished, tentative phylogenetic analysis; in addition the type species and nomenclatural priority for each section were confirmed. Here, we present an updated, genus‐wide phylogenetic analysis, based on four plastid non‐coding regions (3′trnL–trnF, rpl20–5′rps12, psbB–psbH and rpoC2–rps2) for > 150 accessions. The maximum parsimony and maximum likelihood phylogenetic analyses identified eight major clades and one branch with unique sequence variation, although the relationships were unclear due to a polytomy and weak support. In the phylogenetic trees, most of the sections proposed in the latest system were distinct and corresponded to the major clades, but some sections are not monophyletic. On the basis of the phylogenetic relationship: (1) A. schimperianum is treated as a member of section Arisaema, rather than section Tenuipistillata or section Sinarisaema; and (2) section Fimbriata, which was synonymised into section Attenuata in the latest system but is morphologically distinct from the other species, is redefined as a monotypic section. In conclusion, we recognize 15 sections of Arisaema and species‐level classifications are discussed in a phylogenetic context.     

Phylogenetic signal in the evolution of body colour and spicule skeleton in calcareous sponges

Some of the morphological characters used in Porifera taxonomy have often been shown to be inconsistent. In the present study, we tested the phylogenetic coherence of currently used taxonomic characters of the calcarean genus Clathrina. For this, 20 species of Clathrina and three other calcinean genera (Ascandra, Guancha, and Leucetta) were sequenced for the ITS and D2 region of the 28S ribosomal DNA. Maximum‐likelihood and maximum‐parsimony algorithms were used to reconstruct phylogenetic trees. Deep divergences were observed in our tree and Clathrina was shown to be paraphyletic. The major split in our topology showed a clear‐cut distinction between sponges with and without tetractine spicules. Moreover, a group of yellow‐coloured Clathrina was clearly separated from the remaining white‐coloured species. Our results show that the presence of diactines, water‐collecting tubes, the degree of cormus anastomosis, and actine shapes do not correlate with the major clades of the calcinean phylogeny. On the other hand, the presence of tripods, the absence of tetractines, and the presence of spines in the apical actine of tetractines seem to be good synapomorphies for clades in our tree. Our results demonstrate that skeleton characters can be reliably used in higher level taxonomy in Clathrinida. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 1026–1034.     

Phylogenetics and biogeography of the broad-nosed bats, genus Platyrrhinus (Chiroptera: Phyllostomidae)

The Neotropical broad-nosed bats, genus Platyrrhinus, represent a well-defined monophyletic group of 14 recognized species. A recent study of morphological characters confirmed Platyrrhinus monophyly and species diagnosis, but offered little support to their intra-specific relationships. We conducted phylogenetic analyses of the genus, using dense taxonomic sampling in combination with four gene sequences representing both mitochondrial and nuclear DNA transmission systems. Our aim was to elucidate the phylogenetic structure among species, using the resulting 3341 bp of DNA. Maximum parsimony, maximum likelihood, and Bayesian inference analyses produced similar topologies that confirm the monophyly of the genus Platyrrhinus and strongly support many previously unrecognized groups. Paraphyly of Platyrrhinus helleri and the unclear position of P. brachycephalus in the clades were also apparent in the data. Our biogeographical analysis suggests a Brazilian Shield origin for Platyrrhinus, followed by subsequent radiations of lineages in the Amazon Basin and Andes. Secondary dispersal from Amazonian and Andean centers is responsible for the Platyrrhinus inhabiting the Guianan Shield and the Pacific lowlands and Central America, respectively.     

Molecular phylogeny of the entomopathogenic fungi of the genus Cordyceps (Ascomycota: Clavicipitaceae) and its evolutionary implications

Abstract Cordyceps is an endoparasite ascomycetous genus containing approximately 450 species with a diversity of insect hosts, traditionally included in the family Clavicipitaceae of Ascomycota. Establishing the relationships among species with a varied range of morphologies and hosts is of importance to our understanding of the phylogeny and co‐evolution of parasites and hosts in entomopathogenic ascomycetes. To this end, we used a combination of molecular index and morphological characters from 40 representative species to carry out comprehensive molecular phylogenetic analyses. Based on the phylogenetic tree, we used the program DISCRETE for inferring the rates of evolution and finding ancestral states of morphological character. The phylogenetic analyses revealed two important points. (i) Types of perithecia attached to stroma reflected an evolutionary trend in Cordyceps. The vertically immersed perithecia form was the ancestral state, superficial and obliquely immersed perithecia were derived characters, obliquely immersed was irreversible. Species with obliquely immersed perithecia were in a closely related group and were the derived group. (ii) A strong correlation between fungal relatedness and the microhabitat supported the hypothesis that the host jumps through commingling in soil microhabitats. Based on the results of these analyses, host switching explains the diversity of entomopathogenic fungi of the genus Cordyceps.     

Phylogenetics and biogeography of Nephrolepis – a tale of old settlers and young tramps

The phylogenetic relationships and biogeographical history of the tropical genus Nephrolepis, a popular ornamental fern, are investigated using plastid DNA data from three regions: rbcL, rps4 plus the rps4‐trnS intergenic spacer (IGS) and the trnG intron plus the trnG‐trnR IGS. Our taxon sampling includes all but one of the 19 species of the genus. We confirm the monophyly of Nephrolepis, resolve infrageneric relationships and propose monophyly of the widespread species Nephrolepis biserrata and Nephrolepis abrupta, awaiting a denser sampling of Nephrolepis cordifolia and Nephrolepis undulata. The controversial inclusion of Nephrolepis in Lomariopsidaceae is not clearly supported. With a view to identifying synapomorphies for the clades retrieved, we reconstruct the evolution in Nephrolepis of sorus position and indusium shape. Finally, based on an estimation of divergence times and reconstruction of ancestral distributions for the genus, we propose an origin of the crown group in the Eocene in the forests of the Laurasian tropical belt, from where two main lineages would have dispersed and become isolated, one in the Neotropics and the other in Asia–Australasia. From this clear biogeographical pattern, some species, probably of Asian–Australasian origin, show recent range expansions, now spanning the Palaeotropics or the pantropical zone. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 113–127.     

Cyclamen: time, sea and speciation biogeography using a temporally calibrated phylogeny

Aim The Mediterranean region is a species‐rich area with a complex geographical history. Geographical barriers have been removed and restored due to sea level changes and local climatic change. Such barriers have been proposed as a plausible mechanism driving the high levels of speciation and endemism in the Mediterranean basin. This raises the fundamental question: is allopatric isolation the mechanism by which speciation occurs? This study explores the potential driving influence of palaeo‐geographical events on the speciation of Cyclamen (Myrsinaceae), a group with most species endemic to the Mediterranean region. Cyclamen species have been shown experimentally to have few genetic barriers to hybridization. Location The Mediterranean region, including northern Africa, extending eastwards to the Black Sea coast. Methods A generic level molecular phylogeny of Myrsinaceae and Primulaceae is constructed, using Bayesian approximation, to produce a secondary age estimate for the stem lineage of Cyclamen. This estimate is used to calibrate temporally an infrageneric phylogeny of Cyclamen, built with nrDNA ITS, cpDNA trnL‐F and cpDNA rps16 sequences. A biogeographical analysis of Cyclamen is performed using dispersal–vicariance analysis. Results The emergence of the Cyclamen stem lineage is estimated at 30.1–29.2 Ma, and the crown divergence at 12.9–12.2 Ma. The average age of Cyclamen species is 3.7 Myr. Every pair of sister species have mutually exclusive, allopatric distributions relative to each other. This pattern appears typical of divergence events throughout the evolutionary history of the genus. Main conclusions Geographical barriers, such as the varying levels of the Mediterranean Sea, are the most plausible explanation for speciation events throughout the phylogenetic history of Cyclamen. The genus demonstrates distributional patterns congruent with the temporally reticulate palaeogeography of the Mediterranean region.     

Dynamic colonization exchanges between continents and islands drive diversification in paradise‐flycatchers (Terpsiphone,Monarchidae)

Aim We use parametric biogeographical reconstruction based on an extensive DNA sequence dataset to characterize the spatio‐temporal pattern of colonization of the Old World monarch flycatchers (Monarchidae). We then use this framework to examine the role of dispersal and colonization in their evolutionary diversification and to compare plumages between island and continental Terpsiphone species. Location Africa, Asia and the Indian Ocean. Methods We generate a DNA sequence dataset of 2300 bp comprising one nuclear and three mitochondrial markers for 89% (17/19) of the Old World Monarchidae species and 70% of the Terpsiphone subspecies. By applying maximum likelihood and Bayesian phylogenetic methods and implementing a Bayesian molecular clock to provide a temporal framework, we reveal the evolutionary history of the group. Furthermore, we employ both Lagrange and Bayes‐ Lagrange analyses to assess ancestral areas at each node of the phylogeny. By combining the ancestral area reconstruction with information on plumage traits we are able to compare patterns of plumage evolution on islands and continents. Results We provide the first comprehensive molecular phylogenetic reconstruction for the Old World Monarchidae. Our phylogenetic results reveal a relatively recent diversification associated with several dispersal events within this group. Moreover, ancestral area analyses reveal an Asian origin of the Indian Ocean and African clades. Ancestral state reconstruction analyses of plumage characters provide an interpretation of the plumage differentiation on islands and continents. Ancestral plumage traits are inferred to be close to those of the Asian paradise‐flycatcher (Terpsiphone paradisi), and island species display a high degree of plumage autapomorphy compared with continental species. Main conclusions Terpsiphone paradisi is polyphyletic and comprises populations that have retained the ancestral plumage of the widespread Terpsiphone genus. The genus appears to have colonized south‐west Asia, the Indian Ocean and Africa from eastern Asia. The phylogeny and divergence time estimates indicate multiple simultaneous colonizations of the western Old World by Terpsiphone. These results reinforce a hypothesis of range expansions of a Terpsiphone paradisi‐like ancestor into eastern Asia and the western Old World.     

SYSTEMATICS OF GRATELOUPIA FILICINA (HALYMENIACEAE,RHODOPHYTA), BASED ON rbcL SEQUENCE ANALYSES AND MORPHOLOGICAL EVIDENCE,INCLUDING THE REINSTATEMENT OF G. MINIMA AND THE DESCRIPTION OF G. CAPENSIS SP. NOV.1

Grateloupia filicina (C. Agardh) Lamouroux, originally described from the Mediterranean Sea, has long been considered a textbook example of a marine red alga with a cosmopolitan distribution. An rbcL‐based molecular phylogeny, encompassing samples covering the entire geographic distribution of the species, revealed a plethora of “cryptic” species, whereby the presence of genuine G. filicina is limited to the Mediterranean basin. The phylogeny revealed a strong biogeographic imprint, with specimens from temperate regions resolved in clades composed of species inhabiting the same geographic region. Presence of widely divergent morphologies in the temperate clades indicated that several lineages have converged independently to a G. filicina‐type morphology. Tropical representatives are resolved in a single clade with very uniform G. filicina‐type morphology and pairwise sequence divergences that are lower than the average divergence observed in temperate lineages. This, combined with a lack of clear geographic structure among the tropical lineages, may indicate a more recent divergence with long‐range dispersal capacities. Violations to the biogeographic signal in temperate lineages seemed to be due to either inadequate taxonomy or recent introductions. Grateloupia minima P. & H. Crouan, a taxon placed in synonymy under G. filicina, is reinstated as a separate species distributed in the northeast Atlantic Ocean. Grateloupia capensis sp. nov. is described to accommodate specimens from South Africa with a G. filicina‐type morphology, and G. filicina var. luxurians is elevated to species status. Morphological and anatomical characters were put forward that support the distinctiveness of these three distinct species.     

The infrageneric taxonomy of Chaerophyllum (Apiaceae) revisited: new evidence from nuclear ribosomal DNA ITS sequences and fruit anatomy

Evolutionary relationships among 65 Chaerophyllum spp. were inferred from nuclear ribosomal DNA internal transcribed spacer (ITS) sequence variation. Thirty‐one species, represented by 158 mericarp samples, were analysed for fruit anatomical character variation, employing phylogenetic and phenetic methods to explore their congruence with infrageneric divisions based on molecular data. Phylogenetic trees inferred from molecular data using maximum likelihood (ML) and Bayesian inference (BI) methods corroborated the division of the genus into four sections: Chaerophyllum, Dasypetalon, Physocaulis and Chrysocarpum. From among the newly sequenced species, the Greek endemic C. heldreichii was grouped with section Chaerophyllum, whereas the highly variable Asian C. reflexum–C. villosum complex formed an early‐branching paraphyletic assemblage in section Chrysocarpum. The recently described C. karsianum has an identical ITS sequence to C. bulbosum, whereas C. aksekiense was clearly separated from the morphologically similar C. macrospermum. Our study confirmed the postulated synonymy of several species on the basis of morphology, but also demonstrated distant relationships between some morphologically similar species. With the exception of the monotypic section Physocaulis, we were unable to find carpological traits matching sectional divisions. We hypothesize that fruit characters evolved rapidly as a result of diversification of members of the genus in different habitats. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 298–313.     

Further fragmentation of the polyphyletic genus Polyalthia (Annonaceae): molecular phylogenetic support for a broader delimitation of Marsypopetalum

The species-rich genus Polyalthia has previously been shown to be highly polyphyletic, with species represented in at least five different clades. The Polyalthia species that are associated with Marsypopetalum and Trivalvaria (as revealed either by previous phylogenetic studies or inferred on the basis of comparative morphology) were included in a molecular phylogenetic study based on three chloroplast DNA regions (matK, rbcL and trnL-F). Maximum parsimony, maximum likelihood and Bayesian analyses consistently revealed that several Polyalthia species form a well-supported clade with Marsypopetalum pallidum, and that this clade is sister to Trivalvaria. Diagnostic morphological characters for the clades are re-evaluated and shown to be congruent with the molecular phylogeny. Five Polyalthia species (P. crassa, P. littoralis, P. lucida, P. modesta and P. tristis) are accordingly transferred to Marsypopetalum.