West to east dispersal and subsequent rapid diversification of the mega‐diverse genus Begonia (Begoniaceae) in the Malesian archipelago

Aim The complex palaeogeography of the Malesian archipelago, characterized by the evolution of an ever‐changing mosaic of terrestrial and marine areas throughout the Cenozoic, provides the geographic backdrop for the remarkable diversification of Malesian Begonia (> 450 species). This study aimed to investigate the origin of Malesian Begonia, the directionality of dispersal events within the Malesian archipelago and the impact of ancient water gaps on colonization patterns, and to identify drivers of diversification. Location Asia, Southeast Asia, Malesia. Methods Plastid DNA sequence data of representatives of all families of the Cucurbitales and Fagales (matK, rbcL, trnL intron, trnL–F spacer, 4076 aligned positions, 92 taxa) and a sample of all major Asian Begonia sections (ndhA intron, ndhF–rpl32 spacer, rpl32–trnL spacer, 4059 aligned positions, 112 taxa) were analysed under an uncorrelated‐rates relaxed molecular clock model to estimate the age of the Begonia crown group divergence and divergence ages within Asian Begonia. Ancestral areas were reconstructed using a likelihood approach implementing a dispersal–extinction–cladogenesis model, and with a Bayesian approach to dispersal–vicariance analysis. Results The results indicated an initial diversification of Asian Begonia in continental Asia in the Miocene, and subsequent colonization of Malesia by multiple lineages. There was support for at least six independent dispersal events from continental Asia and western Malesia to Wallacea dating from the late Miocene to the Pleistocene. Begonia section Petermannia (> 270 species) originated in Western Malesia, and subsequently dispersed to Wallacea, New Guinea and the Philippines. Lineages within this section diversified rapidly since the Pliocene, coinciding with rapid orogenesis on Sulawesi and New Guinea. Main conclusions The predominant trend of Begonia dispersals between continental Asia and Malesia, and also within Malesia, has been from west to east. The water bodies separating the Sunda Shelf region from Wallacea have been porous barriers to dispersal in Begonia following the emergence of substantial land in eastern Malesia from the late Miocene onwards. We hypothesize two major drivers of the diversification of Malesian Begonia: (1) the formation of topographical heterogeneity and the promotion of microallopatry by orogenesis in the Pliocene and Pleistocene; and (2) cyclic vicariance by frequent habitat fragmentations and amalgamations due to climate and sea‐level fluctuations during the Pleistocene.     

Wallace's Line and plant distributions: two or three phytogeographical areas and where to group Java?

Wallace's Line or its variants divide the Malay Archipelago or Malesia into a western and eastern area, but is this suitable for plant distributions? Indeed, all boundaries satisfactorily divide Malesia into two parts, stopping far more species east or west of a line than disperse over the boundary. However, phenetic analyses (principal components analysis, nonmetric multidimensional scaling analysis and the unweighted pair group method with arithmetic mean) of 7340 species distributions revealed a stronger partitioning of Malesia into three instead of two regions: the western Sunda Shelf minus Java (Malay Peninsula, Sumatra, Borneo), central Wallacea (Philippines, Sulawesi, Lesser Sunda Islands, Moluccas, with Java), and the eastern Sahul Shelf (New Guinea). Java always appears to be part of Wallacea, probably because of its mainly dry monsoon climate. The three phytogeographic areas equal the present climatic division of Malesia. An everwet climate exists on the Sunda and Sahul Shelves, whereas most of Wallacea has a yearly dry monsoon. During glacial maxima, the Sunda and Sahul Shelves became land areas connected with Asia and Australia, respectively, whereas sea barriers remained within Wallacea. Consequently, the flora of the two shelves is more homogeneous than the Wallacean flora. Wallacea is a distinct area because it comprises many endemic, drought tolerant floristic elements. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 531–545.     

A revision of Ptychopyxis (Euphorbiaceae) in southeast Asia

Ptychopyxis is a genus comprised of 11 species. Nine are found in southeast Asia mainland and west Malesia and two species are endemic to New Guinea. One of the endemic New Guinean species is newly described here, viz. P. esdras Stoops sp. nov.     

Extent and timing of floristic exchange between Australian and Asian rain forests

Aim We tested an entrenched concept – that the Australian rain forest flora is essentially a Gondwanan relict. We also assessed the role of regional‐level source–sink dynamics in the assembly of this flora. Location Eastern Australia. Methods To avoid potential biases inherent in selective studies undertaken to date, we used an analytical, whole‐of‐flora approach integrated with the fossil record. We identified disjunctions between woody Australian rain forest plant taxa and relatives on other land masses. To test the strength of the fossil evidence for the regional antiquity of this flora, we evaluated the proportion of these disjunct clades represented in the Australian fossil record, and to minimize the effects of biases in this record, we compared late Quaternary (i.e. late Pleistocene and Holocene, 126–0 ka), Pliocene and late Oligocene–early Miocene Australian pollen records interpreted as tropical rain forest. Using within‐species disjunctions as a proxy, we assessed the role of recent immigration from Asia into Australia. To assess the role of source–sink dynamics, we performed comparative analyses of disjunctions in major rain forest categories representing a north–south/climatic gradient. Results Southern Australian, cool temperate (microthermal) rain forests contain many floristic disjunctions with Gondwanan fragments and most of these clades have Gondwanan fossils. Disjunct clades in Australian mesothermal rain forest mostly occur in Asia/Malesia and a low proportion of these clades show pre‐Neogene records. Many clades in lowland tropical and ‘dry’ rain forest show disjunctions with Asia/Malesia and few have Australian fossil records. Rates of recent immigration from Asia/Malesia are high in these northern forests, and outweigh rates of recent emigration approximately nine‐fold. The late Quaternary fossil record has many more rain forest angiosperms than Oligocene–Miocene and Pliocene floras, consistent with extensive late Cenozoic immigration. Main conclusions The microthermal rain forests are largely Gondwanan relicts, but there is progressively greater, and more recent contribution from Asia/Malesia into more northern, and more lowland tropical rain forests. This variation reflects a strong gradient in geographic and ecological proximity between these forests and source floras in Asia/Malesia, and is consistent with a source–sink size model of immigration driven by late Cenozoic contractions and expansions of Australian rain forest.     

Biogeography of Australasian flightless weevils (Curculionidae,Celeuthetini) suggests permeability of Lydekker's and Wallace's Lines

The Indo‐Australian region was formed by the collision of the Australian and Asian plates, and its fauna largely reflects this dual origin. Lydekker's and Wallace's Lines represent biogeographic transition boundaries between biotas although their permeability through geological times was rarely assessed. Here, we explore the evolutionary history of flightless weevils of the tribe Celeuthetini in this geologically highly complex region. We generated a DNA sequence data set of 2236 bp comprising two nuclear and two mitochondrial markers for 62 species of the Indo‐Australian tribe Celeuthetini. We used Bayesian Inference and Maximum Likelihood to reconstruct the first molecular phylogeny of the group. Based on this phylogenetic tree, we employed the program BioGeoBEARS to infer the biogeographical history of Celeuthetini in the region. The group's radiation begun east of Wallace's Line, probably during the mid‐Eocene. We unveil multiple transgressions of Lydekker's and Wallace's Lines mostly during the Miocene with a significant role of founder‐event speciation. The phylogeny of Celeuthetini is geographically highly structured with the first lineages occurring in New Guinea and the Moluccas, and a deep divergence between two clades largely confined to Sulawesi and their respective sister clades of the Lesser Sunda Islands. Wallace's Line was crossed once from Sulawesi and three times from the Lesser Sunda Islands to Java whilst Lydekker's Line was crossed once from New Guinea to the Moluccas. Although this beetle group shows extensive local diversification with little dispersal, the biogeographical demarcations of the Australasian region appear to have been rather porous barriers to dispersal.     

Phylogeny and spatio‐temporal diversification of Prunus subgenus Laurocerasus section Mesopygeum (Rosaceae) in the Malesian region

The goals of this study were to reconstruct the phylogeny of Prunus subgenus Laurocerasus section Mesopygeum and to provide a preliminary assessment of its spatio‐temporal diversification in the Malesian region. We inferred the phylogeny using nuclear ITS and ETS and plastid psbA‐trnH, rps16, rpl16, and trnC‐petN sequences. Our analyses support the monophyly of sect. Mesopygeum. Within sect. Mesopygeum, we identified four main subclades: (i) Prunus lancilimba from continental Asia; (ii) Prunus ruthii from Malay Peninsula; (iii) a subclade comprising species from areas centered on the Sunda shelf and also a few species from continental Asia and Wallacea; and (iv) a subclade composed of species from areas of the Sahul shelf, with a small number of taxa also from areas of the Sunda shelf, continental Asia, and the Philippines. We estimated that sect. Mesopygeum originated in continental Asia at c. 44.71 Mya (95% HPD: 31.66–46.90). Nine dispersals between major geographic areas were inferred. From continental Asia, three and two dispersals were inferred to the Sunda shelf and the Sahul shelf in the mid‐Oligocene, respectively. Two dispersals were inferred from the Sahul shelf region to the Sunda shelf in late Oligecene and early Miocene, respectively. There were also two dispersals inferred from the Sunda shelf region, one to the Philippines and one to Wallacea, in the middle and late Miocene, respectively. The diversification in sect. Mesopygeum was likely driven by active geologic events and orogenies in the Neogene in the Malesian region.     

Thirteen new species of Dendrochilum (Orchidaceae), a new record from Burma, and a checklist of the genus in East Malesia

Pedersen, H. Æ. 1995. Thirteen new species of Dendrochilum (Orchidaceae), a new record from Burma, and a checklist of the genus in East Malesia. — Nord. J. Bot. 15: 381402. Copenhagen. ISSN 0107–055X.
Seven new species of Dendrochilum are described from the Philippines, viz. D. amesianum, D. flexuosum, D. louisianum, D. parvipapillatum, D. plocoglottoides, D. quisumbingianum and D. tetradactyliferum . The widespread species D. longifolium is recorded from Burma for the first time, and an annotated checklist of Dendrochilum in East Malesia, i.e. the Lesser Sunda Islands, Sulawesi, the Moluccas, and New Guinea is given. This list enumerates a total of 21 species, including D. cornutum and D. pallideflavens which are here recorded from East Malesia for the first time. Six species in the checklist are described as new, viz. D. ambangense, D. citrinum, D. erectila-bium, D. eymae, D. galeatum , and D. longipedicellatum .     

Historical biogeography of Rhododendron section Vireya and the Malesian Archipelago

Aim Vireya rhododendrons are distinctive and easily recognizable by their general form; however, they are virtually circumscribed geographically, predominantly distributed throughout the biogeographically intriguing Malesian Archipelago. Hypotheses of the evolutionary relationships of the group have been proposed but the biogeography of vireyas has not been analysed based on molecular phylogeny. Recently, the first detailed molecular phylogenetic investigation of section Vireya was completed based on cp‐ and nrDNA sequence data, therefore making this cladistic biogeographic study of vireya rhododendrons possible. Location Malesia, Australia, Solomon Islands, Taiwan, Himalayas, north Vietnam and south China. Methods Based on distribution maps, areas of endemism were determined for the biogeographic region of Malesia. Area relationships were analysed based on a recent molecular phylogeny of species in section Vireya. The method of paralogy‐free subtree analysis was applied. Results Individual distribution maps were produced for 74 species of Rhododendron section Vireya. Species clades with bootstrap support proved to be biogeographically informative. Major clades correspond to three regions: eastern Malesia, western/middle Malesia and Taiwan/north Vietnam/south China. Within eastern Malesia, Australia, New Guinea, the Bismarck Archipelago and Solomon Islands are related. In western Malesia, northern Philippines, Borneo, southern Moluccas and north and west Sulawesi are related. These areas are more distantly related to Sumatra, the Malay peninsula, Java, Bali, Palawan, Lesser Sunda islands and the southern Philippines. The position of the Himalayas is equivocal and part of a basal polytomy in the summary area cladogram. Main conclusions Two alternative hypotheses are proposed for the evolution of vireya rhododendrons based on the pattern of area relationships. The first hypothesis is that the vireyas are an old group, with ancestors present on Gondwana, rifting north in the Cretaceous. The second alternative hypothesis is that vireyas are a young group that has dispersed eastwards from India to Australia and the Solomon Islands since the current Malesian islands formed.     

Biogeography of polypores in Malesia,Southeast Asia

Biogeography of polypores distributed in the Malesian region is reviewed. Some of the species reported from this region are pantropical or paleotropical and widely distributed along the equator. Several species are restricted to lowland areas of Malesia and adjacent regions, and can be classified as Asian or Asian-Oceanian tropical species. Some species have only been recorded from lowland areas of Malesia at present, but detailed distributions are unclear for many of them. Some lowland Malesian species are also distributed in temperate areas of East Asia; among them, few species are circumglobal in the Northern Hemisphere, whereas many others are tropical species showing continuous distributions up to the warm temperate areas of East Asia. Species recorded from highland areas of Malesia include tropical species common to lowland Malesia, temperate species common to temperate Asia, and endemic species in highland areas of Malesia and adjacent regions. Regarding the factors that delimitate distribution of the Malesian polypores, I discuss polypore host preference, response to physical environmental factors, and dispersal ability. Systematics and ecological characteristics are briefly discussed for selected Malesian polypores: Corneroporus subcitrinus, Paratrichaptum accuratum, Postia stellifera and Roseofavolus eos.     

Regional patterns of biodiversity in New Guinea plants

Regional patterns of biodiversity in seven recently-studied, speciose groups of New Guinea plants (comprising 200 species, or 1–2% of the flora) are analysed with maps showing numbers of species in 1^o grid cells. Patterns are correlated with the tectonic history of New Guinea. The New Guinea orogen involved rocks of the northern margin of the Australian craton as well as the terranes accreted to the margin, and the current axial range is geologically and biologically composite. The southern Nothofagus has a main massing on the Australian craton portion of the New Guinea mountains. In contrast, four typical genera of Malesian rainforest (Parsonsia, Archidendron, Aglaia, Amyema) have centres of biodiversity on the accreted terranes north of the craton. There are 32 distinct tectono-stratigraphic terranes (some composite) which have been accreted to the craton at different times through the Tertiary and these may have travelled hundreds or even thousands of kilometres before docking. Finally, the 'decaisninoid group' of Loranthaceae and the fern Grammitis have centres of diversity on both the craton and the accreted terranes.     

The phylogenetic relationships of the bilby, Macrotis lagotis (Peramelimorphia: Thylacomyidae), to the bandicoots- DNA sequence evidence

Partial sequencing of the 12S ribosomal RNA gene was used to test two competing hypotheses concerning the phylogenetic relationship of the bilby (Macrotis lagotis) to the Australian and New Guinean species of bandicoot. The first hypothesis proposes that the Australian and New Guinean bandicoots are in a monophyletic clade to the exclusion of the bilbies, whereas the second hypothesis proposes that the bilby is monophyletic with the Australian bandicoots to the exclusion of the New Guinean bandicoots. Phylogenies determined by both maximum-likelihood and neighbour-joining approaches supported the first hypothesis in which the bilby is excluded from the clade represented by the Australian and New Guinean bandicoots. Monophyly of the Australian and New Guinean bandicoots is consistent with the biogeographical scenario in which Australia and Papua New Guinea have undergone repeated connection and disconnection over the last 20 million years.     

Genetic variability,local selection and demographic history: genomic evidence of evolving towards allopatric speciation in Asian seabass

Genomewide analysis of genetic divergence is critically important in understanding the genetic processes of allopatric speciation. We sequenced RAD tags of 131 Asian seabass individuals of six populations from South‐East Asia and Australia/Papua New Guinea. Using 32 433 SNPs, we examined the genetic diversity and patterns of population differentiation across all the populations. We found significant evidence of genetic heterogeneity between South‐East Asian and Australian/Papua New Guinean populations. The Australian/Papua New Guinean populations showed a rather lower level of genetic diversity. F_ST and principal components analysis revealed striking divergence between South‐East Asian and Australian/Papua New Guinean populations. Interestingly, no evidence of contemporary gene flow was observed. The demographic history was further tested based on the folded joint site frequency spectrum. The scenario of ancient migration with historical population size changes was suggested to be the best fit model to explain the genetic divergence of Asian seabass between South‐East Asia and Australia/Papua New Guinea. This scenario also revealed that Australian/Papua New Guinean populations were founded by ancestors from South‐East Asia during mid‐Pleistocene and were completely isolated from the ancestral population after the last glacial retreat. We also detected footprints of local selection, which might be related to differential ecological adaptation. The ancient gene flow was examined and deemed likely insufficient to counteract the genetic differentiation caused by genetic drift. The observed genomic pattern of divergence conflicted with the ‘genomic islands’ scenario. Altogether, Asian seabass have likely been evolving towards allopatric speciation since the split from the ancestral population during mid‐Pleistocene.     

Tree diversity and phytogeographical patterns of tropical high mountain rain forests in Central Sulawesi,Indonesia

Tropical high mountain forests in Lore Lindu National Park, Sulawesi, Indonesia, were described by their floristic composition and the importance of tree families (Family importance values, FIV), based on tree inventories conducted on 4 plots (each 0.24 ha) in old-growth forest stands at c. 1800 and 2400 m a.s.l. (mid- and upper montane elevations). To identify general patterns and regional peculiarities of the forests in the SE Asian and SW Pacific context, the biogeography of the tree species was analysed using distribution records. Out of the total of 87 tree species, only 18 species were found at both elevational zones. The discovery of new species and new distribution records (28% of the data set) highlights the deficiencies in the taxonomic and distribution data for Sulawesi. Sulawesi endemism rate was 20%. In the mid-montane Fagaceae–Myrtaceae forests, Lithocarpus spp. (Fagaceae) were overall important (4 spp. occupying around half of the total basal area) and the Myrtaceae the most species rich (8 spp.), thus showing typical features of Malesian montane forests. The upper montane conifer-Myrtaceae forest contained several high mountain tree taxa and showed affinity to the forests of New Guinea. The mountain flora comprised both eastern and western Malesian elements, with the nearest neighbouring islands Borneo and Maluku both sharing species with Sulawesi, reflecting the complex palaeogeography of the island. A separate analysis showed the mid-montane forest to possess greatest biogeographical affinity to Borneo/western Malesia, and the upper montane forest had a number of typical elements of Papuasia/eastern Malesia and the Phillipines, which may be a result of historical patterns in land connection and the emergence of mountain ranges.     

Snakes across the Strait: trans-Torresian phylogeographic relationships in three genera of Australasian snakes (Serpentes: Elapidae: Acanthophis, Oxyuranus, and Pseudechis)

We analyze the phylogeny of three genera of Australasian elapid snakes (Acanthophis-death adders; Oxyuranus-taipans; Pseudechis-blacksnakes), using parsimony, maximum likelihood, and Bayesian analysis of sequences of the mitochondrial cytochrome b and ND4 genes. In Acanthophis and Pseudechis, we find evidence of multiple trans-Torresian sister-group relationships. Analyses of the timing of cladogenic events suggest crossings of the Torres Strait on several occasions between the late Miocene and the Pleistocene. These results support a hypothesis of repeated land connections between Australia and New Guinea in the late Cenozoic. Additionally, our results reveal undocumented genetic diversity in Acanthophis and Pseudechis, supporting the existence of more species than previously believed, and provide a phylogenetic framework for a reinterpretation of the systematics of these genera. In contrast, our Oxyuranus scutellatus samples from Queensland and two localities in New Guinea share a single haplotype, suggesting very recent (late Pleistocene) genetic exchange between New Guinean and Australian populations.     

Independent histories of human Y chromosomes from Melanesia and Australia

To investigate the origins and relationships of Australian and Melanesian populations, 611 males from 18 populations from Australia, Melanesia, and eastern/southeastern Asia were typed for eight single-nucleotide polymorphism (SNP) loci and seven short tandem-repeat loci on the Y chromosome. A unique haplotype, DYS390.1del/RPS4Y711T, was found at a frequency of 53%-69% in Australian populations, whereas the major haplotypes found in Melanesian populations (M4G/M5T/M9G and DYS390.3del/RPS4Y711T) are absent from the Australian populations. The Y-chromosome data thus indicate independent histories for Australians and Melanesians, a finding that is in agreement with evidence from mtDNA but that contradicts some analyses of autosomal loci, which show a close relationship between Australian and Melanesian (specifically, highland Papua New Guinean) populations. Since the Australian and New Guinean landmasses were connected when first colonized by humans > or =50,000 years ago but separated some 8,000 years ago, a possible way to reconcile all the genetic data is to infer that the Y-chromosome and mtDNA results reflect the past 8,000 years of independent history for Australia and New Guinea, whereas the autosomal loci reflect the long preceding period of common origin and shared history. Two Y-chromosome haplotypes (M119C/M9G and M122C/M9G) that originated in eastern/southeastern Asia are present in coastal and island Melanesia but are rare or absent in both Australia and highland Papua New Guinea. This distribution, along with demographic analyses indicating that population expansions for both haplotypes began approximately 4,000-6,000 years ago, suggests that these haplotypes were brought to Melanesia by the Austronesian expansion. Most of the populations in this study were previously typed for mtDNA SNPs; population differentiation is greater for the Y chromosome than for mtDNA and is significantly correlated with geographic distance, a finding in agreement with results of similar analyses of European populations.     

Species diversity and endemism of five major Malesian islands: diversity–area relationships

Aim A comparison of biodiversity patterns within Malesia in relation to surface area. Location Analysis of the patterns in species richness and endemism of vascular plants in the five major Malesian islands, i.e. Java, Sulawesi, Sumatra, Borneo and New Guinea. Methods Available data on species richness and species ranges in correlation with the surface area of the respective islands were examined in this work. Estimations of total species numbers for these islands are presented based on extrapolation of all available published Flora Malesiana information and recent checklists, all in all comprising 12,000 different species. The regression analysis of overall species richness and endemism were studied for all species together as well as for different plant families to compare the fit with the Arrhenius species–area model. Results The five islands form a series of independent areas of increasing size suited for an analysis of the species–area relationships at the regional scale. All species taken together and those of families with even distribution throughout Malesia show significant species–area relationships. Non‐significant relationships were detected in families with western or eastern‐centred Malesian distribution patterns. Relationships between number of endemic species and surface area are significant for all species and for the majority of the families with significant species–area relationships. Main conclusions Species–area relationships of families appear to be dependent on species number. Families with high numbers of species usually have a significant species–area relationship whereas small families have not. For the families that display an eastern or western Malesian centred pattern, a historical biogeographical explanation should be invoked. Island surface area appears to be a predictor for island percent endemism in Malesian vascular plants. None of the islands appears to be a hotspot of endemism nor of species diversity, as no significant departure from the Arrhenius model was noted for any of them.     

A monograph of the betel nut palms (Areca: Arecaceae) of East Malesia

The palm genus Areca is widespread in tropical Asia and includes the economically important betel nut palm, A. catechu. The genus has three centres of high species diversity: the Sunda Region, the Philippines and East Malesia (to the east of Wallace's line). The taxonomy of the genus in East Malesia has been neglected. Prior to this study, 19 species were accepted for this area, all but one endemic, but their limits and differences were not understood. Here, we provide a taxonomic monograph of East Malesian Areca spp., based on an extensive study of the genus in herbaria and in the field. We recognize six species of Areca in East Malesia, including the widespread cultivated A. catechu. Five wild species are accepted, namely A. macrocalyx, A. mandacanii, A. novohibernica, A. oxycarpa and A. vestiaria. We place 12 of the previously accepted species into synonymy, and provide additional new synonymy in A. catechu. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 147–173.     

Phylogenetic relationships of living and recently extinct bandicoots based on nuclear and mitochondrial DNA sequences

Bandicoots (Peramelemorphia) are a major order of australidelphian marsupials, which despite a fossil record spanning at least the past 25 million years and a pandemic Australasian range, remain poorly understood in terms of their evolutionary relationships. Many living peramelemorphians are critically endangered, making this group an important focus for biological and conservation research. To establish a phylogenetic framework for the group, we compiled a concatenated alignment of nuclear and mitochondrial DNA sequences, comprising representatives of most living and recently extinct species. Our analysis confirmed the currently recognised deep split between Macrotis (Thylacomyidae), Chaeropus (Chaeropodidae) and all other living bandicoots (Peramelidae). The mainly New Guinean rainforest peramelids were returned as the sister clade of Australian dry-country species. The wholly New Guinean Peroryctinae was sister to Echymiperinae. The poorly known and perhaps recently extinct Seram Bandicoot (Rhynchomeles) is sister to Echymipera. Estimates of divergence times from relaxed-clock Bayesian methods suggest that living bandicoots originated in the late Oligocene or early Miocene, much earlier than currently thought based on fossils. Subsequent radiations within Peramelemorphia probably took place on the Australian mainland during the Miocene, with diversification of rainforest taxa on the newly emergent New Guinean landmasses through the middle-late Miocene and complete establishment of modern lineages by the early Pliocene.     

New taxa and name changes in tropical Asiatic Rubiaceae

Four new genera, viz. Deccania Tirveng., Dioecrescis Tirveng., Fagerlindio Tirveng., Kailarsenia Tirveng., one new subgenus, viz. Aidia subgen. Lankaidia Tirveng., seven new species and thirty four of new combinations for South and South East Asia are published, pending the publication of the generic delimitation and revision of the Rubiaceae–Gardenieae in forthcoming flora treatments. New combinations are also included for a few taxa from Malesia, Indonesia and the Philippines.     

ORIGINAL ARTICLE: Cycads in the insular South‐west Pacific: dispersal or vicariance?

Aim Cycads constitute an ancient plant group that is generally believed to disperse poorly. However, one group of cycads (subsection Rumphiae) is thought to have dispersed relatively recently from a Malesian source area westwards to East Africa and eastwards into the Pacific, using a floatation‐facilitating layer in their seeds. We use morphological and allozyme characters to investigate the relationships among the species within this group and to deduce whether the wide distribution was achieved by recent dispersal (as evidenced by high genetic similarity) or more distant vicariance events (high genetic differentiation). Location We examined specimens collected throughout the range of subsection Rumphiae, from East Africa through Southeast Asia to Tonga in the South‐west Pacific. Methods We investigated relationships within subsection Rumphiae of the genus Cycas by analysing 18 variable (11 informative) morphological characters and 22 allozyme loci for seven of the 10 species currently assigned to this taxon. Results Distinctive morphological characters are few and fail to resolve relationships within the group. Allozyme data show that species within this subsection are closely related and suggest that there are two groups within the subsection, one comprising Cycas thouarsii (East Africa) and C. edentata (the Philippines), and the other the remaining species (from Malesia and the Pacific). The Australian species C. silvestris is sister to subsection Rumphiae in the morphological analysis but is closely allied to C. rumphii (nested within the subsection) in the allozyme analysis, suggesting that Rumphiae may be paraphyletic and that characters thought to be taxonomically important may need to be re‐evaluated. Main conclusions Cycads within subsection Rumphiae are closely related, and the wide distribution of this group was probably achieved through relatively recent oceanic dispersal events. Separate events probably account for the dispersal of these cycads into the Pacific and to Africa. The origin and distribution of C. silvestris (Australia) could be explained by a dispersal event from New Guinea or may have resulted from a former land connection between Australia and New Guinea.