Phylogeographic structure in mitochondrial DNA of a South-east Asian freshwater fish, Hemibagrus nemurus (Siluroidei; Bagridae) and Pleistocene sea-level changes on the Sunda shelf

We postulated that the biogeographical history of South-east Asia contributed to extensive admixture during Pleistocene low sea levels of genetic groups of an obligate freshwater fish (the river catfish, Hemibagrus nemurus) isolated during periods of high sea levels. During Pleistocene glacial maxima, the sea level was lower than at present and the islands of the Sunda shelf (Sumatra, Borneo and Java) and the Asian mainland were connected by lowlands traversed by rivers. Restriction fragment length polymorphisms in mitochondrial DNA were documented for 140 putative H. nemurus analysed from 13 sampling sites resulting in the definition of 35 haplotypes. The high level of haplotype differentiation (mean P × 100 = 2.22, SD = 1.33) indicates that the subdivision of the ancestral H. netnurus group was extensive and probably occurred early in the Pleistocene. The occurrence of some genetically divergent groups of the H. netnurus complex occurring in sympatry in widely separated locations supports the proposition that low sea levels aided the dispersion and mingling of genetic groups. Based on both genetic and morphological evidence, the main H. nemurus line gave rise to three regional groups: (1) a morphologically distinct ‘Indochinese’ group composed of two mtDNA clades overlapping in east peninsular Malaysia; (2) a ‘Sundaic’ group composed of various lineages of differing morphology and genetic identity; (3) a genetically distinct ‘Sarawak’ group in west Borneo, similar in morphology to the ‘Sundaic’ and ‘Indochinese’ groups, but including a small, golden colour morph as a distinct dade. The morphologically similar Sundaic forms from west Java, Sumatra and west Borneo show some degree of genetic divergence, but their phylogenetic relationships are poorly resolved. The most genetically and morphologically distinct Sundaic dade, assigned to H. hoevenii, colonized the Kapuas river (west Borneo), east Sumatra and south peninsular Malaysia. Contrary to our original hypothesis and present biogeographical theory, little exchange of genetic groups has apparently occurred between the mainland and the Sunda Islands during recent glaciations.     

Mammalian species richness on islands on the Sunda Shelf,Southeast Asia

Summary A rich mammalian fauna is found on islands that lie on the Sunda Shelf, a continental shelf extending from Vietnam to Borneo and Java that was periodically exposed as dry land during the Pleistocene. The correlation between log of island area and number of species is high (r ²=0.94); the slope of the curve is moderate (z=0.235). Distance from small islands to source areas (=Borneo, Sumatra, and the Malay Peninsula) does not appear to affect species richness, nor does depth of water to source area (a measure of isolation time). A species-area curve for forest reserves of varying sizes on the Malay Peninsula has a low slope (z=0.104); comparison of the mainlaind and island curves indicates that decreasing island area is strongly correlated with increased extinction. Extinction has left reduced but ecologically balanced sets of species on all islands, except that carnivores are under-represented on all but the largest islands. Initial body size and rarity appear to play a significant role in determining the probability of extinction of individual species.     

Mammals of Borneo – small size on a large island

Aim Island mammals have featured prominently in models of the evolution of body size. Most of these models examine size evolution across a wide range of islands in order to test which island characteristics influence evolutionary pathways. Here, we examine the mammalian fauna of a single island, Borneo, where previous work has detected that some mammal species have evolved a relatively small size. We test whether Borneo is characterized by smaller mammals than adjacent areas, and examine possible causes for the different trajectories of size evolution between different Bornean species. Location Sundaland: Borneo, Sumatra, Java and the Malay/Thai Peninsula. Methods We compared the mammalian body size frequency distributions in the four areas to examine whether the large mammal fauna of Borneo is more depauperate than elsewhere. We measured specimens belonging to 54 mammal species that are shared between Borneo and any of the other areas in order to determine whether there is an intraspecific tendency for Bornean mammals to evolve small body size. Using data on diet, body size and geographical ranges we examine factors that are thought to influence body size. Results Borneo has fewer large mammals than the other areas, but this is not statistically significant. Large Bornean mammals are significantly smaller than their conspecifics in the other regions, while there are no differences between the body sizes of mammals on Sumatra, Java and the Malay/Thai Peninsula. The finding that large mammals show the greatest size difference between Borneo and elsewhere contrasts with some models of size evolution on islands of different areas. Diet does not correlate with the degree of size reduction. Sunda region endemics show a weaker tendency to be small on Borneo than do widespread species. Main conclusions We suggest that soil quality may drive size evolution by affecting primary productivity. On Borneo, where soils are generally poor in nutrients, this may both limit biomass and cause mammals to be reduced in body size. We hypothesize that widespread species respond to low resource abundance by reducing in size, while endemic elements of the fauna have had longer to adjust to local conditions by altering their behaviour, physiology and/or ecology, and are thus similar in size across the region.     

The collision of the Indian plate with Asia: molecular evidence for its impact on the phylogeny of freshwater crabs (Brachyura: Potamidae)

Aim We used molecular data to answer the following questions: (1) Is morphology‐based (and to some extent, geography‐based) classification of the freshwater crab family Potamidae congruent with a molecular phylogeny? (2) What historical biogeographical event could have shaped this phylogeny? Location Material from the entire geographical range of the family Potamidae was analysed, including specimens from East Asia (China, Taiwan, the Ryukyus), Southeast Asia, South Asia (northern India, the Middle East and Near East), North Africa, and southern Europe. Methods Mitochondrial DNA sequences encoding 503 base pairs (excluding the variable regions) of the large subunit rRNA (16S rRNA) gene were obtained from 72 species belonging to 49 potamid genera, representing 51% of all known genera in this species‐rich family. Sequences were compared by means of phylogenetic analyses (minimum evolution, Bayesian inference, maximum likelihood and maximum parsimony) and Bayesian relaxed molecular clock estimates. Results The family Potamidae was found to be monophyletic with two major lineages, and there was support for the recognition of two mostly allopatric subfamilies, Potaminae and Potamiscinae. This is largely consistent with the current classification proposed. The ‘Potamiscinae’ clade comprised three subclades: (1) a well‐supported ‘eastern Asia’ subclade that included species from the eastern part of the range (China, Taiwan, the Ryukyus, the Philippines, Indochina, Malay Peninsula, northern India and Myanmar/Burma); (2) a weakly supported ‘Sunda Shelf islands’ subclade that included species from the larger Southeast Asian islands on the Sunda Shelf (Borneo, Sumatra and Java); and (3) a ‘Socotra’ subclade that comprised only Socotrapotamon from Socotra Island, off the north‐east coast of Africa. Main conclusions The discrete distribution of the two subfamilies in Europe/Asia is hypothesized to be the result of vicariance due to the collision of the Indian tectonic plate with the Asian continent, and the orogeny that caused the separation of the two freshwater crab lineages around 22.8 Ma. Within the Potamiscinae, the ‘Sunda Shelf islands’ subclade separated from other potamiscines around 21.1 Ma; and the endemic fauna of the East Asian islands (Taiwan, the Ryukyus and mainland Japan) was isolated from the Asian continent c. 8.4 Ma, following the opening of the Okinawa Trough. The ‘Socotra’ subclade diverged from the ‘eastern Asia’ subclade at 19.1 Ma during the Miocene. Its taxonomic position, however, remains unclear as the members of this clade possess the key potamine character of a transverse ridge on thoracic sternite 8, suggesting that this may in fact be a relict potamid group.     

Mentawai’s endemic,relictual fauna: is it evidence for Pleistocene extinctions on Sumatra?

Aim The four Mentawai islands, south‐west of Sumatra, have long been isolated from the remainder of Sundaland, resulting in a high level of endemism. We examined the distribution of 151 species of the Mentawai Islands in Sundaland and assessed various processes that may have resulted in the various biogeographical patterns. Location Southeast Asia, particularly the Mentawai Islands and nearby large landmasses (Sumatra, Java, Borneo and Peninsular Malaysia). Methods We compared the faunal composition of the Mentawai Islands for selected taxa (43 mammals, 92 reptiles and 16 amphibians) with that of the four nearby large landmasses of Sundaland using morphological comparisons and the most recent molecular phylogenetic analyses available in the literature. These comparisons yielded sister taxa, which were used to simulate species absence data for the four Sundaland landmasses under several scenarios to investigate how patterns of species absence could have arisen. Results In contrast to our expectations, several Mentawai species did not have their closest relatives on neighbouring Sumatra, but rather on the more distant Borneo, Java or Peninsular Malaysia. For mammals, the similarity between species from Mentawai and Borneo was greater than that observed between species from Mentawai and Sumatra. We conclude that the relationships represent traces of species historically distributed throughout Sundaland that became extinct in Sumatra during the Pleistocene. For reptiles and amphibians the observed pattern of species absences generally resembled the simulated pattern expected under the scenario of absence rates increasing with landmass isolation, whereas for mammals we observed more species than expected missing from Java and Sumatra, and fewer than expected from Borneo. Main conclusions The potential extinctions on Sumatra probably had two causes: changes of climate and vegetation during the Pleistocene and environmental impacts from the Toba supervolcanic eruption.     

Vicariance or dispersal? Historical biogeography of three Sunda shelf murine rodents (Maxomys surifer,Leopoldamys sabanus and Maxomys whiteheadi)

The Sunda region of south-east Asia comprises the Malay Peninsula and the islands of Java, Sumatra and Borneo, all of which lie on a shallow continental shelf projecting from Indochina. Pleistocene glacial cycles caused sea levels to drop repeatedly, exposing vast areas of the Sunda shelf and creating land bridges among the islands and mainland. These land bridges, the latest of which connected all three of the major Sunda islands to the Malay Peninsula as recently as 9500 years ago, may have enabled mammalian migrations across the Sunda shelf. Pleistocene land bridges on the Sunda shelf have been invoked to explain the current distributions of mammalian taxa occupying ranges corresponding with the Pleistocene limits of land and the appearance of new mammal species in the Pleistocene fossil record. The ability of mammals to move throughout the exposed shelf during periods of low sea level would, however, have been influenced by topographic and ecological features, which have been variously described as savanna-like or as moist tropical rain forest. Using a phylogeographical approach, we test the hypothesis that Pleistocene land bridges enabled widespread movements in three rain-forest-restricted murine rodents of the Sunda shelf: Maxomys surifer , Leopoldamys sabanus and Maxomys whiteheadi . Our results do not support the hypothesis of broad Pleistocene migrations in these taxa, but instead suggest a deep history of vicariant evolution that may correspond with the Pliocene fragmentation of the Sunda block.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 91–109.     

How island size and isolation affect bee and wasp ensembles on small tropical islands: a case study from Kepulauan Seribu,Indonesia

Aim To study the effects of isolation and size of small tropical islands on species assemblages of bees (superfamily Apoidea) and wasps (superfamily Vespoidea). Location Twenty islands in the Kepulauan Seribu Archipelago off the coast of west Java, Indonesia. The size of surveyed islands ranges between 0.75 and 41.32 ha; their distance from the coast of Java varies between 3 and 62 km. Methods Field work was conducted from February to May 2005. Bees and wasps were caught with a sweep net during sampling units of 15 min, continuing until four consecutive samples revealed no new species. Total species richness was quantified by the estimators Chao 2, first‐order jackknife and Michaelis–Menten. The software binmatnest was used to test for nestedness of species assemblages. Similarities of species composition between islands were quantified by Sørensen’s similarity index. Results Eighty‐two species were recorded on the 20 surveyed islands. Species richness declined with increasing isolation of islands from the source area, Java. Although the size of the largest island exceeded that of the smallest island by a factor of almost 60, island size only very weakly affected species richness of bees; no effect of island size was found for wasps. Mean body size of species decreased with increasing island isolation. Nestedness of island faunas was only weakly developed. Species composition of both superfamilies was affected by island isolation, but not by island size. Main conclusions While the species–isolation relationship on the very small islands of Kepulauan Seribu followed the prediction of MacArthur and Wilson’s equilibrium theory, the absence of a species–area relationship indicated a weak ‘small‐island effect’, at least in wasps. The combination of an only weakly developed pattern of nested species subsets, the shift in species compositions and the decline of mean body size with increasing island isolation from the source area indicates that biotic interactions and different species traits contribute to the shaping of communities of bees and wasps within the archipelago. The potential of biotic interactions for generating distribution patterns of species within the archipelago is also emphasized by the observed restriction of some species with apparently high dispersal abilities to outer islands.     

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.     

Taxonomic revision of shrews of the genus Crociduras from the Sunda Shelf and Sulawesi with description of two new species (Mammalia: Soricidae)

The skulls of 387 shrews of the genus Crocidura sampled in peninsular Malaysia, Sumatra, Borneo, Java and Sulawesi were submitted to principal component and stepwise discriminant analyses. These analyses helped to delineate morphological taxa in this species-rich genus of mammals. Most morphologic groups could be attributed to described species, except one taxon from Sumatra and one from Sulawesi, which are described and named as new. Most of the 21 species recognized in this paper are endemic to one major island. Although Sulawesi has never been connected to the mainland, it supports at least six species, followed by Sumatra (5–6 species), Java and the Malay Peninsula (4 spp) and Borneo (3 spp). C. monticola is apparently the only widespread species whose distribution range covers the entire Malay Archipelago except the Philippines and Sulawesi. In contrast, the continental C. fuliginosa enters only marginally into the Sunda Shelf: its southernmost record is on the Malay Peninsula. This interpretation is completely different from the classical view that C. fuliginosa is a cosmopolitan species occupying the whole of Southeast Asia. Identification keys, tables of measurements and discriminant functions provided in this work may aid in identification of the various species and subspecies of Crocidura living in the Malaya Archipelago.     

How important were stepping stones in the colonization of Krakatau?

In theory, one factor determining the rate and nature of the assembly of island biotas is the presence or absence of stepping stone islands, yet no field studies have demonstrated stepping stone function in practice. Krakatau, in Sunda Strait, is about equidistant from Java and Sumatra. Sebesi lies about half way between Krakatau and Sumatra, but no island intervenes between Krakatau and the nearest coast of Java. We assess the evidence that Sebesi has acted as an important stepping stone for Krakatau's recolonization since the devastating 1883 volcanic eruption. About a quarter of Krakatau's resident land birds, two-fifths of its reptiles, bats and land molluscs, and about two-thirds of its termites, pteridophytes, butterflies and spermatophytes are unknown on Sebesi, evidently having colonized without stepping stone involvement. Identifiable Sumatran taxa do not outnumber identifiable Javan ones on Krakatau, nor do historical distribution records indicate movement from Sebesi to Krakatau in animal groups. Krakatau's biota is not a subset of Sebesi's in predominantly anemochorous or thallassochorous plant groups, butterflies, reptiles or bats, and is only marginally so in termites. It is a subset in predominantly zoochorous spermatophyte groups, except Ficus species, and in birds and land molluscs. Comparison with a weaker stepping stone candidate, Panaitan, provides no evidence for a stepping stone role for Sebesi in butterflies or termites. We discuss the dispersal and establishment constraints on colonization by the groups involved, and conclude that, overall, Sebesi had little impact as a stepping stone. Instead, it is more probable that divergence of the environments of the two islands has led to an increasingly independent recolonization of Krakatau.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 275–317.     

Relationships between morphology, genetics and geography in the cave fruit bat Eonycteris spelaea (Dobson, 1871) from Indonesia

Morphological and genetic analyses of Eonycteris spelaea from 15 islands along the Banda Arc, from Sumatra to Timor and including Kalimantan and Sulawesi, revealed considerable divergence between islands and geographical patterning. On the basis of both morphology and genetics, the populations on the large islands of Greater Sunda (Sumatra, Java, Kalimantan and Sulawesi) are generally distinct from one another and from those on the islands in Nusa Tenggara (Lombok to Timor), which form a more cohesive cluster. These differences may be the result of the Nusa Tenggara populations having been colonized more recently than those on the Greater Sunda, and probably from a single source. All biological measures of the relationships between island populations are positively associated with the extent of the sea-crossing between them, indicating the sea is an important barrier to movement. Multivariate analyses show the presence of a marked trend for body size to increase from west to east. However, individuals from Kalimantan are not consistent with this trend, being smaller than predicted, and on the two outer Banda Arc islands of Sumba and Timor animals are a little larger than predicted from the longitudinal trend. These differences could be due to the relative isolation of these populations or differing environmental conditions. There is also a negative relationship between body size and island area, but this is confounded by the longitudinal trend. No significant longitudinal trends in the genetic data were detected and the trend in body size may be an adaptive response to an environmental cline that is known to occur in this region.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79, 511–522.     

Abundance-age-area relationships in an insular plant community

Two processes are thought to generate positive relationships between species richness and island area. The areaper se hypothesis states that larger islands maintain larger populations, which are less susceptible to extinction. The habitat hypothesis states that larger islands contain more habitats, and therefore a greater number of habitat specialists. However, the importance of each mechanism is debated. I tested the areaper se and habitat hypotheses by comparing relationships between plant abundance, age and island area in five shrub species on islands off the coast of British Columbia, Canada. Results showed that two shrub species increased in both abundance and age with island area. The remaining three species showed no differences in abundance and age with island area. Conifer abundances increased with island area, which generated differences in habitat availability. Smaller islands were dominated by open habitat, while larger islands contained both open and forested habitats. Changes in habitat availability with island area could explain patterns in plant abundance and age. The two species that increased in abundance with island area were commonly found in conifer forest on the mainland, and their distributions were consistent with the distribution forest habitat. Positive relationships between plant age and island area in these two species may result from lower survivorship in the open habitat, which dominated small islands. The three species that showed no relationship between abundance and island area are commonly found in open habitat on the mainland, and their island distributions paralleled the availability of open habitat on islands. Similar plant ages on different sized islands may result from their occurrence in open habitat on both large and small islands. Overall results support the habitat hypothesis and indicate that species distributions result from the interaction between habitat affinities and changes in habitat availability with island area.     

Heterochrony: The evolution of ontogeny. By Michael L. McKinney and Kenneth J. McNamara. New York: Plenum Press. 1991. XIX + 437 pp. ISBN 0-306-43638-8. $ 49.50 (cloth)

Macaca fascicularis is broadly distributed in Southeast Asia across 30° of latitude and 35° of longitude (Indochinese Peninsula, Isthmus of Kra, Malay Peninsula, Greater and Lesser Sunda Islands, Philippine Islands, and numerous small, neighboring islands). The range is divisible into 1) a core area comprised of mainland Southeast Asia, Borneo, Sumatra, and Java (large land masses interconnected during the last glacial maximum, 18,000 B. P.); 2) shallow-water fringing islands, which are smaller islands connected to the core area during the last glacial maximum; and 3) deep-water fringing islands, which are peripheral islands not connected to the core area during the last glacial maximum. Skull length was used to study effects of latitude and insularity on patterns of size variation. The data are from 802 adult M. fascicularis specimens from 140 core-area localities, 63 shallow-water islands, and 29 deep-water islands. Sex-specific polynomial regressions of skull length on latitude were used to describe skull length variation in the core area. These regressions served as standards for evaluating variation among samples from shallow-water and deep-water islands. The core area exhibits Bergmannian latitudinal size clines through most of the species range. Thus, skull length decreases from about 8°S (Java) to the equator (Sumatra and Borneo), then increases as far north as about 13°N (Isthmus of Kra). Farther north, to the northernmost Indochinese localities at about 17°N, skull length in M. fascicularis decreases with increasing latitude, contrary to Bergmann's rule. Latitudinal size variation in shallow-water fringing islands generally parallels that in the core area. However, skull length tends to be smaller than in the core area at similar latitudes. Deep-water fringing islands are markedly more variable, with relatively small specimens in the Lesser Sunda Islands and relatively large specimens in the Nicobar Islands. These analyses illustrate how a primate species may vary in response to latitudinal temperature variation and to isolation. © 1993 Wiley-Liss, Inc.     

Forest fruit production is higher on Sumatra than on Borneo

Background

Various studies have shown that the population densities of a number of forest vertebrates, such as orangutans, are higher on Sumatra than Borneo, and that several species exhibit smaller body sizes on Borneo than Sumatra and mainland Southeast Asia. It has been suggested that differences in forest fruit productivity between the islands can explain these patterns. Here we present a large-scale comparison of forest fruit production between the islands to test this hypothesis.

Methodology/Principal Findings

Data on fruit production were collated from Sumatran and Bornean sites. At six sites we assessed fruit production in three forest types: riverine, peat swamp and dryland forests. We compared fruit production using time-series models during different periods of overall fruit production and in different tree size classes. We examined overall island differences and differences specifically for fruiting period and tree size class. The results of these analyses indicate that overall the Sumatran forests are more productive than those on Borneo. This difference remains when each of the three forest types (dryland, riverine, and peat) are examined separately. The difference also holds over most tree sizes and fruiting periods.

Conclusions/Significance

Our results provide strong support for the hypothesis that forest fruit productivity is higher on Sumatra than Borneo. This difference is most likely the result of the overall younger and more volcanic soils on Sumatra than Borneo. These results contribute to our understanding of the determinants of faunal density and the evolution of body size on both islands.     

Local endemism and within‐island diversification of shrews illustrate the importance of speciation in building Sundaland mammal diversity

Island systems are important models for evolutionary biology because they provide convenient, discrete biogeographic units of study. Continental islands with a history of intermittent dry land connections confound the discrete definitions of islands and have led zoologists to predict (i) little differentiation of terrestrial organisms among continental shelf islands and (ii) extinction, rather than speciation, to be the main cause of differences in community composition among islands. However, few continental island systems have been subjected to well‐sampled phylogeographic studies, leaving these biogeographic assumptions of connectivity largely untested. We analysed nine unlinked loci from shrews of the genus Crocidura from seven mountains and two lowland localities on the Sundaic continental shelf islands of Sumatra and Java. Coalescent species delimitation strongly supported all currently recognized Crocidura species from Sumatra (six species) and Java (five species), as well as one undescribed species endemic to each island. We find that nearly all species of Crocidura in the region are endemic to a single island and several of these have their closest relative(s) on the same island. Intra‐island genetic divergence among allopatric, conspecific populations is often substantial, perhaps indicating species‐level diversity remains underestimated. One recent (Pleistocene) speciation event generated two morphologically distinct, syntopic species on Java, further highlighting the prevalence of within‐island diversification. Our results suggest that both between‐ and within‐island speciation processes generated local endemism in Sundaland, supplementing the traditional view that the region's fauna is relictual and primarily governed by extinction.     

Island history affects faunal composition: the treeshrews (Mammalia: Scandentia: Tupaiidae) from the Mentawai and Batu Islands,Indonesia

The Mentawai and Batu Island groups off the west coast of Sumatra have a complicated geological and biogeographical history. The Batu Islands have shared a connection with the Sumatran ‘mainland’ during periods of lowered sea level, whereas the Mentawai Islands, despite being a similar distance from Sumatra, have remained isolated from Sumatra, and probably from the Batu Islands as well. These contrasting historical relationships to Sumatra have influenced the compositions of the respective mammalian faunas of these island groups. Treeshrews (Scandentia, Tupaiidae) from these islands have, at various times in their history, been recognized as geographically circumscribed populations of a broadly distributed Tupaia glis, subspecies, or distinct species. We used multivariate analyses of measurements from the skull and hands to compare the island populations from Siberut (Mentawai Islands) and Tanahbala (Batu Islands) with the geographically adjacent species from the southern Mentawai Islands (T. chrysogaster) and Sumatra (T. ferruginea). Results from both the skull and manus of the Siberut population show that it is most similar to T. chrysogaster, whereas the Tanahbala population is more similar to T. ferruginea, confirming predictions based on island history. These results are further corroborated by mammae counts. Based on these lines of evidence, we include the Siberut population in T. chrysogaster and the Tanahbala population in T. ferruginea. Our conclusions expand the known distributions of both the Mentawai and Sumatran species. The larger geographical range of the endangered T. chrysogaster has conservation implications for this Mentawai endemic, so populations and habitat should be re‐evaluated on each of the islands it inhabits. However, until such a re‐evaluation is conducted, we recommend that the IUCN Red List status of this species be changed from ‘Endangered’ to ‘Data Deficient’. Published 2013. This article is a U.S. Government work and is in the public domain in the USA, Biological Journal of the Linnean Society, 2014, 111 , 290–304.     

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.     

Genetic structure of Gymnures (genus Hylomys; Erinaceidae) on continental islands of Southeast Asia: historical effects of fragmentation

Eustatic sea level changes during Pleistocene climatic fluctuations produced several cycles of connection-isolation among continental islands of the Sunda shelf. To explore the potential effects of these fluctuations, we reconstructed a model of the vicariant events that separated these islands, based on bathymetric information. Among many possible scenarios, two opposite phylogenetic patterns of evolution were predicted for terrestrial organisms living in this region: one is based on the classical allopatric speciation mode of evolution, while the other is the outcome of a sequential dispersal colonization of the archipelago. We tested the applicability of these predictions with an analysis of sequence variation of the cytochrome b gene from several taxa of Hylomys. They were sampled throughout SE-Asia and the Sunda islands. High levels of haplotype differentiation characterize the different island taxa. Such levels of differentiation support the existence of several allopatric species, as was suggested by previous allozyme and morphological data. Also in accordance with previous results, the occurrence of two sympatric species from Sumatra is suggested by their strongly divergent haplotypes. One species, Hylomys suillus maxi, is found both on Sumatra and in Peninsular Malaysia, while the other, H. parvus, is endemic to Sumatra. Its closest relative is H. suillus dorsalis from Borneo. Phylogenetic reconstructions also demonstrate the existence of a Sundaic clade composed of all island taxa, as opposed to those from the continent. Although there is no statistical support for either proposed biogeographic model of evolution, we argue that the sequential dispersal scenario is more appropriate to describe the genetic variation found among the Hylomys taxa. However, despite strong differentiation among island haplotypes, the cladistic relationships between some island taxa could not be resolved. We argue that this is evidence of a rapid radiation, suggesting that the separation of the islands may have been perceived as a simultaneous event rather than as a succession of vicariant events. Furthermore, the estimates of divergence times between the haplotypes of these taxa suggest that this radiation may actually have predated the climatic fluctuations of the Pleistocene. Further refinement of the initial palaeogeographic models of evolution are therefore needed to account for these results.     

Clambidae of Sulawesi with reference to the biogeography of the Sunda Islands (Coleoptera, Eucinetoidea)

Abstract. Eight new species of Clambidae from Sulawesi (Celebes), and two new species and a new subspecies from Borneo and Sumatra are described. The emerging distribution patterns reflect the long-standing isolation of Sulawesi from the rest of the Sunda Islands and from the Asian continent. The clambid fauna of Borneo, Sumatra and the Malayan Peninsula indicates close faunal-historic contacts.     

Macrogeographical variability in the great call of Hylobates agilis: assessing the applicability of vocal analysis in studies of fine‐scale taxonomy of gibbons

Vocal characteristics have been used extensively to distinguish different taxonomic units of gibbons (family Hylobatidae). The agile gibbon (Hylobates agilis) has a disjunct distribution range in the Southeast Asian archipelago (remnants of the former Sunda landmass), and populations on different islands are currently recognized as distinct subspecies or even species. We recorded great calls from female agile gibbons from two populations on Sumatra and two populations on Borneo and examined the vocal variability on four levels: within‐individuals, between‐individuals, between‐populations and between‐islands. The primary objective was to evaluate the effect of geographical isolation on variability in song pattern and to test whether proposed island‐specific song characteristics exist, reflecting evolutionary divergence between Sumatran and Bornean agile gibbons. One hundred great calls were recorded from 20 females and analyzed for 18 spectral and temporal acoustic parameters. Principal component analysis followed by a nested ANOVA on components revealed a complex pattern of song variability not likely to reflect taxonomic or evolutionary relationship. We found no evidence that Sumatran and Bornean agile gibbons have evolved different vocal characteristics, refuting a distinction between them based on vocal characteristics. A high level of plasticity was found in great calls from the same individual, and generally the inferred pattern of variability suggested that ecological or social factors may confound any genetically based island dialects. Am. J. Primatol. 72:142–151, 2010. © 2009 Wiley‐Liss, Inc.