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.     

Mosaics in the mangroves: allopatric diversification of tree‐climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo‐West Pacific

The Indo‐Australian Archipelago (IAA) is the richest area of biodiversity in the marine realm, yet the processes that generate and maintain this diversity are poorly understood and have hardly been studied in the mangrove biotope. Cerithidea is a genus of marine and brackish‐water snails restricted to mangrove habitats in the Indo‐West Pacific, and its species are believed to have a short pelagic larval life. Using molecular and morphological techniques, we demonstrate the existence of 15 species, reconstruct their phylogeny and plot their geographical ranges. Sister species show a pattern of narrowly allopatric ranges across the IAA, with overlap only between clades that show evidence of ecological differentiation. These allopatric mosaic distributions suggest that speciation may have been driven by isolation during low sea‐level stands, during episodes preceding the Plio‐Pleistocene glaciations. The Makassar Strait forms a biogeographical barrier hindering eastward dispersal, corresponding to part of Wallace's Line in the terrestrial realm. Areas of maximum diversity of mangrove plants and their associated molluscs do not coincide closely. © 2013 The Natural History Museum. Biological Journal of the Linnean Society © 2013 The Linnean Society of London, 2013, 110 , 564–580.     

Transgressing Wallace's Line brings hyperdiverse weevils down to earth

Wallace's Line, located in the heart of the Indo-Australian archipelago, has historically been hypothesized to strongly inhibit dispersal. Taxa crossing this barrier are confronted with different biota of Asian or Australian origin, respectively, but the extent to which these conditions have affected the evolution of the colonizing lineages remains largely unknown. We examined the potential correlations of body size, lifestyle and biogeographical distribution in the weevil genus Trigonopterus. These beetles are highly diverse both on foliage and in litter east of Wallace's Line but occur exclusively in leaf litter in the west. Based on a comprehensive, dated phylogeny of 303 species, we inferred nine crossing events of Wallace's Line, all from east to west. Five previously foliage-dwelling lineages changed their lifestyle to leaf litter habitats after crossing this barrier. Our results indicate that dispersal is not more likely in edaphic lineages, but rather that abiotic and/or biotic factors may be responsible for the exclusive leaf litter habitat of Trigonopterus in Sundaland. This includes differences in climate, and the different predatory faunas of Australia-New Guinea, Wallacea and Sundaland. A mimicry complex in New Guinea with Trigonopterus species as presumable model may be of relevance in this context.     

Crossing the (Wallace) line: local abundance and distribution of mammals across biogeographic barriers

Past and ongoing vertebrate introductions threaten to rearrange ecological communities in the Indo‐Malay Archipelago, one of Earth's most biodiverse regions. But the consequences of these translocations are difficult to predict. We compared local abundance and distributions in four tropical mammal lineages that have crossed from Asia to Wallacea or New Guinea. The local abundance of macaques (Macaca spp.), which naturally crossed Wallace's Line, was higher in Sulawesi (east of the line; mean = 3.7 individuals per camera station, 95% CI = 2.2: 5.1) than in Borneo (west of the line; mean = 1.1, CI = 0.8: 1.4), but the local abundance of Malay civets (Viverra tangalunga), Rusa deer, and Sus pigs was similar in their native ranges and where they had been introduced by humans east of Wallace's Line. Proximity to rivers increased Malay Civet local abundance and decreased the local abundance of pigs in parts of their introduced ranges (Maluku and New Guinea, respectively), while having no effect on local abundance in their native ranges (Borneo) or other areas where they have been introduced (Sulawesi). That local abundance was higher east of Wallace's Line in just one of four mammal lineages is consistent with findings from plant invasions, where most species have similar abundance in their native and introduced ranges. However, species’ ecology may change as they enter new communities, for example, their patterns of abundance at local scales. This could make it difficult to predict community structure in the face of ongoing species introductions.     

GEOGRAPHIC AND TAXONOMIC DISPARITIES IN SPECIES DIVERSITY: DISPERSAL AND DIVERSIFICATION RATES ACROSS WALLACE'S LINE

Broad‐scale patterns of species diversity have received much attention in the literature, yet the mechanisms behind their formation may not explain species richness disparities across small spatial scales. Few taxa display high species diversity on either side of Wallace's Line and our understanding of the processes causing this biogeographical pattern remains limited, particularly in plant lineages. To understand the evolution of this biogeographical pattern, a time‐calibrated molecular phylogeny of Livistoninae palms (Arecaceae) was used to infer the colonization history of the Sahul tectonic plate region and to test for disparities in diversification rates across taxa and across each side of Wallace's Line. Our analyses allowed us to examine how timing, migration history, and shifts in diversification rates have contributed to shape the biogeographical pattern observed in Livistoninae. We inferred that each of the three genera found in Sahul crossed Wallace's Line only once and relatively recently. In addition, at least two of the three dispersing genera underwent an elevation in their diversification rate leading to high species richness on each side of Wallacea. The correspondence of our results with Southeast Asian geologic and climatic history show how palms emerge as excellent models for understanding the historical formation of fine‐scale biogeographic patterns in a phylogenetic framework.     

The distribution of the Malay civet Viverra tangalunga (Carnivora: Viverridae) across Southeast Asia: natural or human‐mediated dispersal?

The Malay civet Viverra tangalunga Gray, 1832 is a fairly large viverrid that has a wide distribution in both the Sundaic and Wallacea regions of Southeast Asia. We investigated the genetic diversity of V. tangalunga by analysing the mitochondrial DNA of 81 individuals throughout its range in order to elucidate the evolutionary history of this species and to test the hypotheses of natural dispersal and/or potential human introductions to some islands and regions. Our phylogenetic analyses revealed that V. tangalunga has a low matrilinear genetic diversity and is poorly structured geographically. Borneo is likely to have served as the ancestral population source from which animals dispersed during the Pleistocene. Viverra tangalunga could have naturally dispersed to Peninsular Malaysia, Sumatra, and Belitung, and also to several other Sunda Islands (Bangka, Lingga, and Bintang in the Rhio Archipelago), and to Palawan, although there is possible evidence that humans introduced V. tangalunga to the latter islands. Our results strongly suggested that V. tangalunga was transported by humans across Wallace's Line to Sulawesi and the Moluccas, but also to the Philippines and the Natuna Islands. Our study has shown that human‐mediated dispersal can be an important factor in understanding the distribution of some species in this region. © 2014 The Linnean Society of London     

Alfred R. Wallace's enduring influence on biogeographical studies of the Indo-Australian archipelago

To mark A.R. Wallace's 200th birthday, we review the direct and indirect contributions he made to our understanding of the Indo-Australian Archipelago's biogeography. He is widely known for his field research (1854–1862) and his 1863 boundary line separating the Oriental and Australasian faunal realms (between Bali and Lombok, Borneo and Sulawesi, and the Philippines and the Moluccas). Notably, though, he never accepted Huxley's ‘Wallace Line’ proposal (1868), whose northern part runs between the main Philippine islands and the Palawan Group to the west. Furthermore, in 1910, which was 3 years prior to his demise, he transferred Sulawesi's fauna to the Oriental realm. In 1924, Merrill introduced the ‘Wallacea’ transition zone. Although the label is today widely used to denote a sub-region within the Indo-Australian Archipelago between Wallace's 1863 line and Lydekker's 1896 line (first presented by Darlington in 1957), the western boundary was originally based on Huxley's line, and thus included the Philippine islands minus the Palawan group. Most biogeographers appear to be unaware of Merrill and his intention. Finally, recent attempts to define the faunal break have not led to a consensus view, despite the huge increase in primary data plus the application of modern analytical techniques. This reflects the complexities and diversity of the region's faunal distribution patterns, plus the differences in the ways that researchers choose to process their data.     

Phylogenetics of fanged frogs: testing biogeographical hypotheses at the interface of the asian and Australian faunal zones

The interface of the Asian and Australian faunal zones is defined by a network of deep ocean trenches that separate intervening islands of the Philippines and Wallacea (Sulawesi, the Lesser Sundas, and the Moluccas). Studies of this region by Wallace marked the genesis of the field of biogeography, yet few workers have used molecular methods to investigate the biogeography of taxa whose distribution spans this interface. Some taxa, such as the fanged frogs of the ranid genus Limnonectes, have distributions on either side of the zoogeographical lines of Wallace and Huxley, offering an opportunity to ask how frequently these purported barriers were crossed and by what paths. To examine diversification of Limnonectes in Southeast Asia, the Philippines, and Wallacea, we estimated a phylogeny from mitochondrial DNA sequences obtained from a robust geographic sample. Our analyses suggest that these frogs dispersed from Borneo to the Philippines at least twice, from Borneo to Sulawesi once or twice, from Sulawesi to the Philippines once, and from the Philippines to Sulawesi once. Dispersal to the Moluccas occurred from Sulawesi and to the Lesser Sundas from Java/Bali. Species distributions are generally concordant with Pleistocene aggregate island complexes of the Philippines and with areas of endemism on Sulawesi. We conclude that the recognition of zoogeographic lines, though insightful, may oversimplify the biogeography of widespread taxa in this region.     

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.     

Evolution of Pacific Rim diving beetles sheds light on Amphi‐Pacific biogeography

The origin of biodiversity in the Neotropics predominantly stems either from Gondwana breakup or late dispersal events from the Nearctic region. Here, we investigate the biogeography of a diving beetle clade whose distribution encompasses parts of the Oriental region, the Indo‐Australian archipelago (IAA) and the Neotropics. We reconstructed a dated molecular phylogeny, inferred diversification dynamics and estimated ancestral areas under different biogeographic assumptions. For the Oriental region and the IAA, we reveal repeated and complex colonization patterns out of Australia, across the major biogeographic lines in the region (e.g. Wallace's Line). The timing of colonization events across the IAA broadly coincides with the proposed timing of the formation of major geographic features in the region. Our phylogenetic hypothesis recovers Neotropical species nested in two derived clades. We recover an origin of the group in the early Eocene about 55 million yr ago, long after the break‐up of Gondwana initiated, but before a complete separation of Australia, Antarctica and the Neotropics. When allowing an old Gondwanan ancestor, we reconstruct an intricate pattern of Gondwanan vicariance and trans‐Pacific long‐distance dispersal from Australia toward the Neotropics. When restricting the ancestral range to more plausible geological area combinations in the Eocene, we infer an Australian origin with two trans‐Pacific long‐distance dispersal events toward the Neotropics. Our results support on one hand a potential Gondwanan signature associated with regional extinctions in the Cenozoic and with Antarctica serving as a link between Australia and the Neotropics. On the other hand, they also support a trans‐Pacific dispersal of these beetles toward the Andean coast in the Oligocene.     

A marine fish follows Wallace's Line: the phylogeography of the three-spot seahorse (Hippocampus trimaculatus, Syngnathidae, Teleostei) in Southeast Asia

Aim To test the potential of two contrasting biogeographical hypotheses (‘Indian/Pacific Ocean Basin’ vs. ‘Wallace's Line’) to explain the distribution of genetic diversity among populations of a marine fish in Southeast Asia. Location The marine waters of Asia and Southeast Asia: from India to Japan, and east to the Indonesian islands of Sulawesi and Flores. Methods We sequenced a 696 base pair fragment of cytochrome b DNA of 100 individuals of Hippocampus trimaculatus Leach 1814 (three‐spot seahorse), obtained from across its range. We tested our hypotheses using phylogenetic reconstructions and analyses of molecular variance. Results Significant genetic divergence was observed among the specimens. Two distinct lineages emerged that diverged by an average of 2.9%. The genetic split was geographically associated, but surprisingly it indicated a major east–west division similar to the terrestrial Wallace's Line (Φ_ST = 0.662, P < 0.001) rather than one consistent with an Indian‐Pacific ocean basin separation hypothesis (Φ_ST = 0.023, P = 0.153). Samples from east of Wallace's Line, when analysed separately, however, were consistent with an Indian/Pacific Ocean separation (Φ_ST = 0.461, P = 0.005). The degree of genetic and geographical structure within each lineage also varied. Lineage A, to the west, was evolutionarily shallow (star‐like), and the haplotypes it contained often occurred over a wide area. Lineage B to the east had greater genetic structure, and there was also some evidence of geographical localization of sublineages within B. Main conclusions Our results indicate that the genetic diversity of marine organisms in Southeast Asia may reflect a more complex history than the simple division between two major ocean basins that has been proposed by previous authors. In particular, the east–west genetic division observed here is novel among marine organisms examined to date. The high haplotype, but low nucleotide diversity to the west of Wallace's Line is consistent with post‐glacial colonization of the Sunda Shelf. Additional data are needed to test the generality of these patterns.     

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 .     

South Pacific biogeography,tectonic calibration,and pre‐drift tectonics: cladogenesis in Abrotanella (Asteraceae)

Abrotanella is the basal genus in the large tribe Senecioneae (Asteraceae) and has a disjunct distribution in Australasia and South America. A recent molecular phylogeny of the genus was used to investigate whether the main biogeographical patterns in the group could be related to the region's tectonic history in a coherent way. The phylogenetic/biogeographical breaks and overlaps in the genus imply a series of vicariance and range expansion events. Each of these can be related to one of the main tectonic events in the region, including assembly of the New Zealand terranes, crustal extension, and magmatism in Gondwana that preceded seafloor spreading, opening of the Tasman and Pacific basins, and transcurrent movement on the New Zealand Alpine fault. The coincident sequence indicates that pre‐drift tectonics and magmatism have been more important for the origin of trans‐Tasman and trans‐Pacific groups than the final rifting of Gondwana that led to their disjunction. For example, during the pre‐drift phase of break‐up, the Whitsunday volcanic province of Australia and the Median Batholith of New Zealand formed a large, active igneous belt. Its distribution is aligned with the break between New Zealand–south‐eastern Australia clades, and New Zealand–New Guinea clades. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

Repeated evolution of carnivory among Indo‐Australian rodents

Convergent evolution, often observed in island archipelagos, provides compelling evidence for the importance of natural selection as a generator of species and ecological diversity. The Indo‐Australian Archipelago (IAA) is the world's largest island system and encompasses distinct biogeographic units, including the Asian (Sunda) and Australian (Sahul) continental shelves, which together bracket the oceanic archipelagos of the Philippines and Wallacea. Each of these biogeographic units houses numerous endemic rodents in the family Muridae. Carnivorous murids, that is those that feed on animals, have evolved independently in Sunda, Sulawesi (part of Wallacea), the Philippines, and Sahul, but the number of origins of carnivory among IAA murids is unknown. We conducted a comprehensive phylogenetic analysis of carnivorous murids of the IAA, combined with estimates of ancestral states for broad diet categories (herbivore, omnivore, and carnivore) and geographic ranges. These analyses demonstrate that carnivory evolved independently four times after overwater colonization, including in situ origins on the Philippines, Sulawesi, and Sahul. In each biogeographic unit the origin of carnivory was followed by evolution of more specialized carnivorous ecomorphs such as vermivores, insectivores, and amphibious rats.     

Connectivity and the development of population genetic structure in Indo‐West Pacific coral reef communities

Aim To identify connectivity patterns among coral reefs of the Indo‐West Pacific. Projecting connectivity forward in time provides a framework for studying long‐term source–sink dynamics in the region, and makes it possible to evaluate the manner in which migration shapes population genetic structure at regional scales. This information is essential for addressing critical gaps in knowledge for conservation planning efforts in one of the most biologically diverse regions on earth. Location Coral reefs of the Indo‐West Pacific, ranging from 15° S to 30° N and 95° E to 140° E. Methods Individual‐based biophysical dispersal models were used in conjunction with matrix projection to identify the expected patterns of exchange between coral reefs over time. Results Present‐day oceanographic conditions lead to the transport of larvae from the South China Sea into the Coral Triangle region via the Sulu Sea, and from northern Papua New Guinea and the Solomon Islands via Halmahera. The directionality of the system leads to the expected accumulation of organisms from outlying areas into the Coral Triangle region over time, particularly in the vicinity of the Maluku Islands and eastern Sulawesi. Coral reefs in Papua New Guinea, the Sulu Archipelago and areas within the Philippines are expected to be areas of high diversity as well. Main conclusions Biophysical dispersal models, used in conjunction with matrix projection, provide an effective means of simulating connectivity structure across the Indo‐West Pacific and thereby evaluating the directionality of genetic diversity. Migration appears to have a significant influence on population genetic structure in the region. Based on present‐day ocean currents, coral reefs in the South China Sea, northern Papua New Guinea and the Solomon Islands are contributing to high levels of diversity in the Coral Triangle.     

‘After Africa’: the evolutionary history and systematics of the genus Charaxes Ochsenheimer (Lepidoptera: Nymphalidae) in the Indo‐Pacific region

The predominantly Afrotropical genus Charaxes is represented by 31 known species outside of Africa (excluding subgenus Polyura Billberg). We explored the biogeographic history of the genus using every known non‐African species, with several African species as outgroup taxa. A phylogenetic hypothesis is proposed, based on molecular characters of the mitochondrial genes cytochrome oxidase subunit I (COI) and NADH dehydrogenase 5 (ND5), and the nuclear wingless gene. Phylogenetic analyses based on maximum parsimony and Bayesian inference of the combined dataset implies that the Indo‐Pacific Charaxes form a monophyletic assemblage, with the exception of Charaxes solon Fabricius. Eight major lineages are recognized in the Indo‐Pacific, here designated the solon (+African), elwesi, harmodius, amycus, mars, eurialus, latona, nitebis, and bernardus clades. Species group relationships are concordant with morphology and, based on the phylogeny, we present the first systematic appraisal and classification of all non‐African species. A biogeographical analysis reveals that, after the genus originated in Africa, the evolutionary history of Charaxes in the Indo‐Pacific, in particular Wallacea, may be correlated with the inferred geological and climatic history of the region. We propose that Wallacea was the area of origin of all Charaxes (excluding C. solon) occurring to the east of Wallace's [1863] Line. The earliest Indo‐Pacific lineages appear to have diverged subsequent to the initial fragmentation of a palaeo‐continent approximately 13 million years ago. Further diversification in Indo‐Pacific Charaxes appears primarily related to climatic changes during the Pliocene and possibly as recently as the Pleistocene. Although both dispersal and vicariance have played important roles in the evolution of the genus within the region, the latter has been particularly responsible for diversification of Charaxes in Wallacea. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 457–481.     

Historical biogeography of Australian Rhamnaceae, tribe Pomaderreae

Aim To discover the pattern of relationships of areas of endemism for Australian genera in the plant family Rhamnaceae tribe Pomaderreae for comparison with other taxa and interpretation of biogeographical history. Location Australian mainland, Tasmania and New Zealand. Methods A molecular phylogeny and geographic distribution of species within four clades of Pomaderreae are used as a basis for recognition of areas of endemism and analysis of area relationships using paralogy‐free subtrees. The taxon phylogeny is the strict consensus tree from a parsimony analysis of 54 taxa, in four clades, and sequence data for the internal transcribed spacer regions of ribosomal DNA (ITS1‐5.8S‐ITS2) and the plastid DNA region trnL‐F. Results The biogeographical analysis identified five subtrees, which, after parsimony analysis, resulted in a minimal tree with 100% consistency and seven resolved nodes. Three sets of area relationships were identified: the areas of Arnhem and Kimberley in tropical north Australia are related based on the phylogeny of taxa within Cryptandra; the moister South‐west of Western Australia, its sister area the coastal Geraldton Sandplains, the semi‐arid Interzone region and arid Western Desert are related, based on taxa within Cryptandra, Spyridium, Trymalium and Pomaderris; and the eastern regions of Queensland, McPherson‐Macleay, south‐eastern New South Wales (NSW), Victoria, southern Australia, Tasmania and New Zealand are related based on Cryptandra, Pomaderris and Spyridium. Tasmania and NSW are related based entirely on Cryptandra, but the position of New Zealand relative to the other south‐eastern Australian regions is unresolved. Main conclusions The method of paralogy‐free subtrees identified a general pattern of geographic area relationships based on Australian Pomaderreae. The widespread distribution of clades, the high level of endemicity and the age of fossils for the family, suggest that the Pomaderreae are an old group among the Australian flora. Their biogeographical history may date to the early Palaeogene with subsequent changes through to the Pleistocene.     

Systematics and biogeography of Indo-Pacific ground-doves

Ground-doves represent an insular bird radiation distributed across the Indo-Pacific. The radiation comprises sixteen extant species, two species believed to be extinct and six species known to be extinct. In the present study, we present a molecular phylogeny for all sixteen extant species, based on two mitochondrial markers. We demonstrate that the Gallicolumba as currently circumscribed is not monophyletic and recommend reinstalling the name Alopecoenas for a monophyletic radiation comprising ten extant species, distributed in New Guinea, the Lesser Sundas and Oceania. Gallicolumba remains the name for six species confined to New Guinea the Philippines and Sulawesi. Although our phylogenetic analyses fail to support a single origin for the remaining Gallicolumba species, we suspect that the addition of nuclear sequence data may alter this result. Because a number of ground-dove taxa have gone extinct, it is difficult to assess biogeographical patterns. However, the Alopecoenas clade has clearly colonized many remote oceanic islands rather recently, with several significant water crossings. The Gallicolumba radiation(s), on the other hand, is significantly older and it is possible that diversification within that group may in part have been shaped by plate tectonics and corresponding re-arrangements of land masses within the Philippine and Sulawesi region.     

Diversification of a ‘great speciator’ in the Wallacea region: differing responses of closely related resident and migratory kingfisher species (Aves: Alcedinidae: Todiramphus)

The Collared Kingfisher species complex is the most widespread of the ‘great speciator’ lineages of the Indo‐Pacific. They have shown a remarkable ability to spread and diversify. As a result of this rapid diversification, Todiramphus species are often found in secondary sympatry. In Southeast Sulawesi, Indonesia, two Todiramphus species are present, the breeding resident Collared Kingfisher Todiramphus chloris and the overwintering migratory Sacred Kingfisher Todiramphus sanctus. We investigated the effect of isolation on these closely related species by comparing their populations on mainland Sulawesi and its larger continental islands, with populations on the small, oceanic Wakatobi Islands. Within our wider analysis we provide further support for the distinctiveness of the Sulawesi Collared Kingfisher population, perhaps isolated by the deep water barrier of Wallace's line. Within Sulawesi we found that populations of Collared Kingfisher on the Wakatobi Islands had diverged from those on mainland Sulawesi, differing both in morphology and in mitochondrial DNA. In contrast, there was no divergence between Sacred Kingfisher populations in either morphology or mitochondrial DNA. We propose that a difference in habitat occupied by Collared Kingfisher populations between the mainland and continental islands vs. oceanic islands has caused this divergence. Mainland Collared Kingfishers are predominately found inland, whereas Wakatobi Collared Kingfishers are also found in coastal habitats. The larger body size of Wakatobi Collared Kingfisher populations may be a result of increased competition with predominantly coastal Sacred Kingfisher populations. The uniform nature of Sacred Kingfisher populations in this region probably reflects their consistent habitat choice (coastal mangrove) and their migratory nature. The demands of their breeding range are likely to have an even stronger selective influence than their Sulawesi wintering range, limiting their scope for divergence. These results provide insight into the adaptability of the widespread Todiramphus lineage and are evidence of the need for further taxonomic revision of Collared Kingfisher populations.     

Phylogeny of Cerberus (Serpentes: Homalopsinae) and phylogeography of Cerberus rynchops: diversification of a coastal marine snake in Southeast Asia

Aim The biogeography of Southeast Asia has been greatly affected by plate tectonic events over the last 10 Myr and changing sea levels during the Quaternary. We investigated how these events may have influenced the evolution of Cerberus Cuvier, a marine coastal snake belonging to the Homalopsinae (Oriental‐Australian Rear‐fanged Water Snakes). This study is an expansion of a previous study on the biogeography and systematics of Cerberus. Location We obtained species from localities across the range of the widely distributed Cerberus: India, Sri Lanka, the Andaman islands, Myanmar, the Philippines, Borneo, Suluwesi, Sumatra, Vietnam, Thailand, Singapore and Australia. Methods We analysed mtDNA sequences (12S, ND3, ATPase, 2338 nucleotide characters) from 21 localities. The sample consisted of 65 Cerberus rynchops (Schneider), three Cerberus australis (Gray) and four Cerberus microlepis Boulenger. One Homalopsis buccata (Linnaeus), one Bitia hydroides Gray, one Enhydris enhydris (Schneider), and two Enhydris plumbea (Boie) were used as outgroups. Results We produced phylogenetic trees based on parsimony, maximum likelihood and Bayesian analysis. We did not find unambiguous support for the monophly of Cerberus. Cerberus austalis, H. buccata and all other Cerberus populations formed a three‐way basal polytomy under parsimony and C. australis formed the sister group to a clade consisting of H. buccata and all other Cerberus in likelihood and Bayesian analysis. The non‐Australian Cerberus were monophyletic and consisted of four primary biogeographical clades: Indian and Mayanmar, Philippines, Greater Sunda Islands and Suluwesi, and the Thai‐Malay peninsula and Gulf of Thailand. The range of genetic divergence between these clades and Australian Cerberus was 0.06–0.12. Genetic divergence among clades to the west of Australia was less pronounced (Thai‐Malay peninsula and Gulf of Thailand = 0.02–0.05; Sunda Islands and Suluwesi = 0.02–0.05; Philippines = 0.02–0.06; India and Myanmar = 0.04–0.06, Philippines = 0.02–0.5). Main conclusions Gyi [University of Kansas Publications, Museum of Natural History 20 (1970), 47] recognized three species of Cerberus: C. australis (from Australia), C. microlepis (known only from Lake Buhi in the Philippines), and the widely distributed C. rynchops (India to Wallacea). We did not find strong support for the monophyly of the genus. Cerberus australis is highly divergent from all other Cerberus lineages sampled from this region. The geographically widespread C. rynchops is resolved into four biogeographical clades (Indian and Myanmar, Philippines, Greater Sunda Islands and Suluwesi, and the Thai‐Malay Peninsula and Gulf of Thailand). We discuss how the dispersal biology of a salt‐water tolerant, coastal marine taxon and the complex geological history of the region (Tertiary plate tectonic movements and Quaternary sea‐level changes) could produce the observed patterns of diversification.