Old Aegean palaeoevents driving the diversification of an endemic isopod species (Oniscidea,Trachelipodidae)

The Greek endemic isopod species Trachelipus aegaeus is distributed in Aegean islands and the adjacent coastal parts of the Greek mainland. Major palaeogeographic events of the Aegean archipelago, such as the formation of the mid‐Aegean trench and the Messinian Salinity Crisis, have been often employed as major causal factors of evolutionary events and phylogeographic patterns exhibited by several taxa. Herein, we infer phylogenetic relationships among T. aegaeus populations using partial cytochrome oxidase subunit I (COI) and 16S rRNA sequences. Due to the poor preservation of the specimens, we propose a modified DNA extraction protocol, which returned highly positive results in terms of the quality of the total extracted DNA. We implement a calibrated molecular clock and path sampling analysis, using alternative palaeogeographic events and rates of substitution, to evaluate the biogeographic history of the species and to estimate the chronology of diversification events among its populations. Our results are clearly in favour of the scenario of the MAT triggering vicariance among most T. aegaus populations. Moreover, the large intraspecific genetic divergence (0–19% for COI and 0–20.3% for the 16S rRNA) and the overall phylogeographic patterns depicted herein seem not to have been obscured by more recent palaeogeological events. A role of dispersal, probably human‐aided, is assumed for certain ‘deviant’ cases.     

Cryptic diversity within the endemic prehensile‐tailed gecko Urocotyledon inexpectata across the Seychelles Islands: patterns of phylogeographical structure and isolation at the multilocus level

The deciphering of the process of genetic differentiation of species with insular distributions is relevant for biogeographical and conservation reasons. Despite their importance as old gondwanic islands and part of the western Indian Ocean biodiversity hotspot, little is known about the genetic structure of taxa from the Seychelles Islands. We have examined the patterns of structure and isolation within Urocotyledon inexpectata (Reptilia: Geckkonidae), an endemic species from this archipelago. Genetic diversity was screened from populations across the archipelago for both mitochondrial and nuclear genes. Gene genealogies and model‐based inference were used to explore patterns and timings of isolation between the main lineages. High levels of genetic diversity were found for the mitochondrial and some of the nuclear markers. This species harbours at least two highly differentiated lineages, exclusively distributed across the northern and southern groups of the islands. The main split between these was dated back to the Miocene–late Pliocene, but isolation events throughout the Pliocene and Pleistocene were also inferred. Migration between groups of islands was apparently nonexistent, except for one case. The low dispersal capabilities of this species, together with the intrinsic fragmented nature of its geographical distribution, seem to have resulted in highly structured populations, despite the cyclic periods of contact between the different island groups. These populations may currently represent more than one species, making U. inexpectata another example of a morphologically cryptic lineage with deep genetic divergence within gekkonids. The observed patterns suggest a hypothetical biogeographic scenario (of a main north–south phylogeographic break) for the Seychelles that can be further tested with the exploration of the phylogeographic structure of other Seychellois taxa. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 177–191.     

The xeric side of the Brazilian Atlantic Forest: The forces shaping phylogeographic structure of cacti

In order to investigate biogeographic influences on xeric biota in the Brazilian Atlantic Forest (BAF), a biodiversity hotspot, we used a monophyletic group including three cactus taxa as a model to perform a phylogeographic study: Cereus fernambucensis subsp. fernambucensis, C. fernambucensis subsp. sericifer, and C. insularis. These cacti are allopatric and grow in xeric habitats along BAF, including isolated granite and gneiss rock outcrops (Inselbergs), sand dune vegetation (Restinga forest), and the rocky shore of an oceanic archipelago (islands of Fernando de Noronha). The nucleotide information from nuclear gene phytochrome C and plastid intergenic spacer trnS‐trnG was used to perform different approaches and statistical analyses, comprising population structure, demographic changes, phylogenetic relationships, and biogeographic reconstruction in both spatial and temporal scales. We recovered four allopatric population groups with highly supported branches in the phylogenetic tree with divergence initiated in the middle Pleistocene: southern distribution of C. fernambucensis subsp. fernambucensis, northern distribution of C. fernambucensis subsp. fernambucensis together with C. insularis, southern distribution of C. fernambucensis subsp. sericifer, and northern distribution of C. fernambucensis subsp. sericifer. Further, the results suggest that genetic diversity of population groups was strongly shaped by an initial colonization event from south to north followed by fragmentation. The phylogenetic pattern found for C. insularis is plausible with peripatric speciation in the archipelago of Fernando de Noronha. To explain the phylogeographic patterns, the putative effects of both climatic and sea level changes as well as neotectonic activity during the Pleistocene are discussed.     

Bridging the gap: western rock skinks (Trachylepis sulcata) have a short history in South Africa

Phylogeographic patterns in wide-ranging species in southern Africa remain largely unexplored, especially in areas north of South Africa. Here, we investigate population structuring, demographic history, and the colonization pattern of the western rock skink (Trachylepis sulcata), a rock-dwelling species with a range extending from southwestern South Africa into Angola. Using 1056 bp from the mitochondrial marker ND2 and > 2.5 kb from three nuclear genes (EXPH5, KIF24, RAG-1), we constructed allele networks, generated extended Bayesian skyline plots and performed population clustering analyses. Analyses of historical demographic patterns show an overall southward range expansion from Northern Namibia into Southern Namibia and South Africa, although we find contrasting genetic breaks across these geographic regions using nuclear and mitochondrial data. We suggest that mtDNA has introgressed across a nuclear break corresponding to the Knersvlakte region of South Africa, a previously proposed biogeographic barrier for rupicolous species. This pattern of mitochondrial variation contrasts sharply to that of other South African taxa previously investigated, which all show significant mtDNA differentiation across the Knersvlakte region. Additionally, while other taxa show divergences dating to the Pliocene, T. sulcata appears to be a recent arrival in southern Africa, having crossed this barrier and colonized South Africa in the mid-Pleistocene. The complex phylogeographic history of T. sulcata corroborates the intricate patterns of genetic variation found in South African taxa and provides novel insight into historical processes affecting species distributed across Namibia.     

Recent colonization and expansion through the Lesser Sundas by seven amphibian and reptile species

The Lesser Sundas Archipelago is comprised of two parallel chains of islands that extend between the Asian continental shelf (Sundaland) and Australo‐Papuan continental shelf (Sahul). These islands have served as stepping stones for taxa dispersing between the Asian and Australo‐Papuan biogeographical realms. While the oceanic barriers have prevented many species from colonizing the archipelago, a number of terrestrial vertebrate species have colonized the islands either by rafting/swimming or by human introduction. Here, we examine phylogeographic structure within the Lesser Sundas for three snake, two lizard and two frog species that each has a Sunda Shelf origin. These species are suspected to have recently colonized the archipelago, though all have inhabited the Lesser Sundas for over 100 years. We sequenced mtDNA from 231 samples to test whether there is sufficiently deep genetic structure within any of these taxa to reject human‐mediated introduction. Additionally, we tested for genetic signatures of population expansion consistent with recent introduction and estimated the ages of Lesser Sundas clades, if any exist. Our results show little to no genetic structure between populations on different islands in five species and moderate structure in two species. Nucleotide diversity is low for all species, and the ages of the most recent common ancestor for species with monophyletic Lesser Sundas lineages date to the Holocene or late Pleistocene. These results support the hypothesis that these species entered the archipelago relatively recently and either naturally colonized or were introduced by humans to most of the larger islands in the archipelago within a short time span.     

Phytogeography of Lusitanian Macaronesia: biogeographic affinities in species richness and assemblage composition

Analysis of biogeographic affinities is a key tool to establish and improve the resolution of hierarchical biogeographic systems. We describe patterns of species richness of the marine macroalgal flora across Lusitanian Macaronesia (Azores, Madeira, the Salvage Islands and the Canary Islands), and test (i) whether such differences are related to differences in proximity to the nearest continental shore and size among islands. We also explore biogeographic affinities in the composition of macroalgal assemblages (= presence/absence of each taxon in multivariate datasets) to determine (ii) whether each archipelago is a biogeographic unit within this ecoregion and (iii) whether patterns in assemblage composition are related to proximity (i.e. distances) among islands. Presence/absence matrices were created to test and visualize multivariate affinities among archipelagos. A total of 872 taxa were compiled. Species richness peaked at the Canary Islands and decreased towards the Azores; the pattern matched a progressive increase in distance from the nearest continental shores, matching the classical island biogeography theory. Intra-archipelago differences in species richness were largely related to variations in island size. Biogeographic similarities among archipelagos were hierarchically structured. Madeira and the Salvage Islands constituted one biogeographic unit. Floras from the Azores, Madeira and the Salvage Islands were barely separable from each other, but were different from those at the Canary Islands. Such biogeographic similarities among islands were negatively correlated with the geographical separation (i.e. distances) among them. Proximity to nearby continental shores, in conjunction with large- and meso-scale oceanographic patterns, seems to interact to create patterns in richness and composition of algal assemblages across Lusitanian Macaronesia.     

Phylogeographic analysis detects congruent biogeographic patterns between a woodland agamid and Australian wet tropics taxa despite disparate evolutionary trajectories

Aim To test the congruence of phylogeographic patterns and processes between a woodland agamid lizard (Diporiphora australis) and well‐studied Australian wet tropics fauna. Specifically, to determine whether the biogeographic history of D. australis is more consistent with a history of vicariance, which is common in wet tropics fauna, or with a history of dispersal with expansion, which would be expected for species occupying woodland habitats that expanded with the increasingly drier conditions in eastern Australia during the Miocene–Pleistocene. Location North‐eastern Australia. Methods Field‐collected and museum tissue samples from across the entire distribution of D. australis were used to compile a comprehensive phylo‐geographic dataset based on c. 1400 bp of mitochondrial DNA (mtDNA), incorporating the ND2 protein‐coding gene. We used phylogenetic methods to assess biogeographic patterns within D. australis and relaxed molecular clock analyses were conducted to estimate divergence times. Hierarchical Shimodaira–Hasegawa tests were used to test alternative topologies representing vicariant, dispersal and mixed dispersal/vicariant biogeographic hypotheses. Phylogenetic analyses were combined with phylogeographic analyses to gain an insight into the evolutionary processes operating within D. australis. Results Phylogenetic analyses identified six major mtDNA clades within D. australis, with phylogeographic patterns closely matching those seen in many wet tropics taxa. Congruent phylogeographic breaks were observed across the Black Mountain Corridor, Burdekin and St Lawrence Gaps. Divergence amongst clades was found to decrease in a north–south direction, with a trend of increasing population expansion in the south. Main conclusions While phylogeographic patterns in D australis reflect those seen in many rain forest fauna of the wet tropics, the evolutionary processes underlying these patterns appear to be very different. Our results support a history of sequential colonization of D. australis from north to south across major biogeographic barriers from the late Miocene–Pleistocene. These patterns are most likely in response to expanding woodland habitats. Our results strengthen the data available for this iconic region in Australia by exploring the understudied woodland habitats. In addition, our study shows the importance of thorough investigations of not only the biogeographic patterns displayed by species but also the evolutionary processes underlying such patterns.     

Riding the Kuroshio Current: Stepping stone dispersal of the Okinawa tree lizard across the East Asian Island Arc

Aim

Located hundreds of kilometres offshore of continental mainland Asia, the extremely high level of land vertebrate endemism in the East Asian Island Arc provides an excellent opportunity to test hypotheses regarding biogeographic processes and speciation. In this study, we aim to test alternative explanations for lineage diversification (vicariance versus dispersal models), and further develop a temporal framework for diversification in our focal taxon, which is consistent with the known age of these islands. We achieve these tests by investigating the historical biogeography of the Okinawa tree lizard (Japalura polygonata), one of the few widely‐distributed reptiles across this archipelago.

Location

The East Asian Island Arc: (1) Central Ryukyu (Amami and Okinawa groups); (2) Southern Ryukyu (Miyako and Yaeyama groups); (3) Taiwan and adjacent islands.

Methods

A total of 246 tissues were sampled from 10 localities in the Ryukyu archipelago and 17 localities in Taiwan, covering the entire distributional range of this species, including all subspecies. DNA sequences of the mitochondrial cytochrome b, 16S ribosomal RNA, nuclear BACH‐1 and RAG‐1 genes (total: 4,684 bp) were obtained from these samples. We used maximum likelihood and Bayesian methods to infer phylogeny and divergence time, and used a model‐fitting method of biogeographical inference to estimate ancestral range evolution.

Results

Multiple lines of evidence combine to identify a general pattern of dispersal‐mediated diversification northward through the archipelago, following initial dispersal from Taiwan. These included (1) a phylogenetic estimate, revealing a sequential, south‐to‐north branching pattern; (2) ancestral range estimation, inferring multiple overseas dispersals and subsequent colonization of new landmasses; and (3) a reduction in genetic variation observed in successively‐diverging lineages, decreasing from Taiwan northward, towards more remote islands. These results provide strong statistical support for an interpretation of successive bouts of dispersal via the powerful, well‐documented, south‐to‐north Kuroshio Current. Estimation of divergence times suggests that most clades in southern Ryukyu and Taiwan diverged early, giving rise to lineages that have remained isolated, and that more recently‐diverged lineages then colonized northward to subsequently occupy the landmasses of the Central Ryukyu archipelago.

Main conclusions

Our general inference of biogeographic history in Japalura polygonata suggested that this species originated on Taiwan and the Yaeyama group, and arrived at its current distribution in Miyako, Okinawa, Toku and Amami islands by a series of stepping‐stone dispersals, which we report for the first time for a terrestrial vertebrate endemic to this region.     

SEQUENTIAL RADIATIONS AND PATTERNS OF SPECIATION IN THE HAWAIIAN CRICKET GENUS LAUPALA INFERRED FROM DNA SEQUENCES

The tremendous diversity of endemic Hawaiian crickets is thought to have originated primarily through intraisland radiations, in contrast to an interisland mode of diversification in the native Hawaiian Drosophila. The Hawaiian cricket genus Laupala (family Gryllidae) is one of several native genera of flightless crickets found in rain-forest habitat across the Hawaiian archipelago. I examined the phylogenetic relationships among mitochondrial DNA (mtDNA) sequences sampled from 17 species of Laupala, including the 12S ribosomal RNA (rRNA), transfer RNA (RNA)^val and 16S rRNA regions. The distribution of mtDNA variants suggests that species within Laupala are endemic to single islands. The phylogenetic estimate produced from both maximum likelihood and maximum parsimony supports the hypothesis that speciation in Laupala occurred mainly within islands. The inferred biogeographical history suggests that diversification in Laupala began on Kauai, the oldest rain-forested Hawaiian island. Subsequently, colonization to younger islands in the archipelago resulted in a radiation of considerable phylogenetic diversity. Phylogenetic patterns in mtDNA are not congruent with prior systematic or taxonomic hypotheses. Hypotheses that may explain the conflict between the phylogenetic patterns of mtDNA variation and the species taxonomy are discussed.     

Evolutionary basis of mitonuclear discordance between sister species of mole salamanders (Ambystoma sp.)

Distinct genetic markers should show similar patterns of differentiation between species reflecting their common evolutionary histories, yet there are increasing examples of differences in the biogeographic distribution of species‐specific nuclear (nuDNA) and mitochondrial DNA (mtDNA) variants within and between species. Identifying the evolutionary processes that underlie these anomalous patterns of genetic differentiation is an important goal. Here, we analyse the putative mitonuclear discordance observed between sister species of mole salamanders (Ambystoma barbouri and A. texanum) in which A. barbouri‐specific mtDNA is found in animals located within the range of A. texanum. We test three hypotheses for this discordance (undetected range expansion, mtDNA introgression, and hybridization) using nuDNA and mtDNA data analysed with methods that varied in the parameters estimated and the timescales measured. Results from a Bayesian clustering technique (structure ), bidirectional estimates of gene flow (migrate ‐n and IMa2) and phylogeny‐based methods (*beast , buck y) all support the conclusion that the discordance is due to geographically restricted mtDNA introgression from A. barbouri into A. texanum. Limited data on species‐specific tooth morphology match this conclusion. Significant differences in environmental conditions exist between sites where A. texanum with and without A. barbouri‐like mtDNA occur, suggesting a possible role for selection in the process of introgression. Overall, our study provides a general example of the value of using complimentary analyses to make inferences of the directionality, timescale, and source of mtDNA introgression in animals.     

LIFE-HISTORY VARIATION AND COMPARATIVE PHYLOGEOGRAPHY OF DARTERS (PISCES: PERCIDAE) FROM THE NORTH AMERICAN CENTRAL HIGHLANDS

Phylogeography seeks to evaluate the relationship between genetic variation and geographic distribution of a species to examine the influence of historical events on divergence. Congruent phylogeographic patterns in codistributed species indicate historical association of the taxa being compared, and the uniform action of biogeographic events in shaping genetic variation. We sought to evaluate the congruence of patterns of genetic variation of five closely related fish species across a well-defined biogeographic boundary. We gathered allozymic and mitochondrial DNA (mtDNA) sequence data for five species of darters (Percidae: Etheostomatini) from populations distributed among biogeographic regions in the Ozark and Ouachita highlands of the south-central United States. Comparisons among species revealed noncongruence in the magnitude of genetic divergence in both allozymes and mtDNA sequences. We hypothesized that noncongruence resulted, in part, from differences in life histories of the species in our comparison. To address this hypothesis, we evaluated the association between gene flow (measured by N_em) and variation in body size and fecundity variables because they have been shown to influence gene flow in fishes. Correlation analysis revealed an association between gene flow and fecundity (r = 0.88), but not with body size (r = 0.36) or reproductive investment per individual (r = –0.23). The result was similar when independent contrasts of the original variables were used in correlation analyses. Phylogeographic analysis of mtDNA sequence data indicated the importance of history, evident in gene trees of Percina nasuta and Percina phoxocephala. Divergence rates between these two taxa may differ because of historically persistent differences in population sizes, reflected in present-day abundance and fecundity differences. Conversely, Percina caprodes showed little evidence of divergence in mtDNA sequences and yielded the highest mean N_em values from allozyme data. Comparisons among closely related, codistributed taxa may help discriminate among the possibilities for noncongruence in biogeographic studies by allowing evaluation of the influence of life history on patterns of gene flow.     

Genetic and morphometric diversity in Wallacea: geographical patterning in the horse shoe bat, Rhinolophus affinis

Genetic and morphometric variation was examined in eleven island populations of the horse‐shoe bat, Rhinolophus affinis, at the easterly end of this widespread species’ range and encompassing the Australian–Oriental biogeographic interface. Allozyme variation revealed mean heterozygosity levels within islands of 0.047, which is near the mammalian average. However, heterozygosity tended to decline from west to east as populations approached the periphery of the species’ distribution, and was lowest in those islands that were separated by the greatest sea‐crossing from source populations. There is extensive between‐island genetic differentiation (mean F_ST = 0.40) and relationships between islands are associated with their arrangement in geographical space; genetic distance is correlated with geographical distance and the genetic arrangement of islands is associated with longitude. The arrangement of islands as indicated by variation in body and skull metrics is also associated with their geographical positions, and the metric and genetic measures are themselves associated. While other taxa in the region have shown genetic‐geographical concordances, R. affinis is the only one that displays concordant patterns in metrical features. These patterns in biological diversity are interpreted as arising from the sequential island population structure and clines in key biogeographic gradients.     

Caribbean golden orbweaving spiders maintain gene flow with North America

The Caribbean archipelago offers one of the best natural arenas for testing biogeographic hypotheses. The intermediate dispersal model of biogeography (IDM) predicts variation in species richness among lineages on islands to relate to their dispersal potential. To test this model, one would need background knowledge of dispersal potential of lineages and their biogeographic patterns, which has been problematic as evidenced by our prior work on the Caribbean tetragnathid spiders. In order to investigate the biogeographic imprint of an excellent disperser, we study Trichonephila in the Americas. Trichonephila is a nephilid genus that contains globally distributed species known to overcome long, overwater distances. The results of our phylogenetic and population genetic analyses on T. clavipes suggest that populations over the Caribbean and North America maintain a lively gene flow. However, the single species status of T. clavipes over the entire New World is challenged by our species delimitation analyses. Combined with prior evidence from spider genera of different dispersal ability, these patterns coming from an excellent disperser (Trichonephila) that is species-poor and of a relatively homogenous genetic structure, support the IDM predictions.     

Causal or spurious relationship? Climate and the distribution of Phelsuma geckos on Grand Comoro Island

The volcanic island of Grand Comoro, Malagasy biogeographic region, is inhabited by three species of Phelsuma day geckos; two island‐endemic taxa (Phelsuma comorensis and Phelsuma v‐nigra comoraegrandensis) and the introduced Phelsuma dubia. Phelsuma comorensis is restricted to elevations of greater than 150 m above sea level on the northern of the island's two volcanoes and is the only Phelsuma above 300 m. The other species are widespread at low elevations but also reach levels above 900 m at the southern volcano. To investigate these divergent distribution patterns, we used environmental niche models based on climate and habitat data and tested whether predicted climate change may influence species distributions. Analyses of niche overlap did not show significant differences between present‐day and predicted future potential distributions of any Phelsuma species studied, which could be seen as an indicator of resilience towards climate change. Climate models reflected the restricted distribution of P. comorensis with precipitation of the wettest month detected as most important variable, whereas habitat models predicted an island‐wide distribution. While climate appears to determine the distribution of P. comorensis, we propose isolation by migration barriers as an alternative and discuss the detection of causal versus spurious relationships in ecological niche models.     

Spiders on a Hot Volcanic Roof: Colonisation Pathways and Phylogeography of the Canary Islands Endemic Trap-Door Spider Titanidiops canariensis (Araneae,Idiopidae)

Studies conducted on volcanic islands have greatly contributed to our current understanding of how organisms diversify. The Canary Islands archipelago, located northwest of the coast of northern Africa, harbours a large number of endemic taxa. Because of their low vagility, mygalomorph spiders are usually absent from oceanic islands. The spider Titanidiops canariensis, which inhabits the easternmost islands of the archipelago, constitutes an exception to this rule. Here, we use a multi-locus approach that combines three mitochondrial and four nuclear genes to investigate the origins and phylogeography of this remarkable trap-door spider. We provide a timeframe for the colonisation of the Canary Islands using two alternative approaches: concatenation and species tree inference in a Bayesian relaxed clock framework. Additionally, we investigate the existence of cryptic species on the islands by means of a Bayesian multi-locus species delimitation method. Our results indicate that T. canariensis colonised the Canary Islands once, most likely during the Miocene, although discrepancies between the timeframes from different approaches make the exact timing uncertain. A complex evolutionary history for the species in the archipelago is revealed, which involves two independent colonisations of Fuerteventura from the ancestral range of T. canariensis in northern Lanzarote and a possible back colonisation of southern Lanzarote. The data further corroborate a previously proposed volcanic refugium, highlighting the impact of the dynamic volcanic history of the island on the phylogeographic patterns of the endemic taxa. T. canariensis includes at least two different species, one inhabiting the Jandia peninsula and central Fuerteventura and one spanning from central Fuerteventura to Lanzarote. Our data suggest that the extant northern African Titanidiops lineages may have expanded to the region after the islands were colonised and, hence, are not the source of colonisation. In addition, T. maroccanus may harbour several cryptic species.     

Systematics,biogeography and evolution of the endemic Hemidactylus geckos (Reptilia,Squamata, Gekkonidae) of the Cape Verde Islands: based on morphology and mitochondrial and nuclear DNA sequences

A total of 1854 bp of mitochondrial DNA (669 bp of cytochrome b (cyt b) and 386 bp of 12S rRNA), and 804 bp of a nuclear gene (RAG2) were investigated in endemic Hemidactylus from eight Cape Verde Islands, and used to explore their phylogeny, biogeography and evolution. Maximum‐likelihood, maximum‐parsimony and Bayesian analyses based on mtDNA revealed four well‐supported clades with uncorrected genetic divergences of 7.8–12.4% in the cyt b plus 12S rRNA genes, which were also supported by nuclear DNA. A population from the southern island of Fogo is the most divergent in both molecules and morphology and is described as Hemidactylus lopezjuradoi sp. n., and the populations on Sal and Boavista are also assigned species status as H. boavistensis. Although divergent in their DNA, the clade on S. Nicolau and that in the north‐western islands are morphologically similar and both are assigned to H. bouvieri for the present. Hemidactylus b. razoensis from Raso is genetically similar to H. b. bouvieri and differs only in its smaller body size. A molecular clock suggests that the ancestor of the endemic Hemidactylus of the Cape Verde Islands colonized the archipelago approximately 10 ± 2.48 Mya, perhaps reaching the north‐eastern islands first. The H. lopezjuradoi lineage separated soon after, and the north‐western islands were colonized progressively but slowly, S. Nicolau probably being reached first, then S. Vicente and islands on the same bank, and finally Sto. Antão, which is likely to have been colonized less than 1 Mya. Hemidactylus boavistensis is abundant on the arid islands where it occurs, but H. bouvieri appears to have been uncommon at least since it was described 130 years ago, and the same may be true of H. lopezjuradoi sp. n. The impact of introduced H. angulatus and H. mabouia on the endemic Hemidactylus of the Cape Verde Islands is not clear, but the discovery of substantial genetic diversity in endemic Cape Verde Hemidactylus means that the conservation requirements of the group should be reassessed.     

Effects of historical forest contraction on the phylogeographic structure of Australo‐Papuan populations of the red‐legged pademelon (Macropodidae: Thylogale stigmatica)

Geographic patterns of genetic variation are strongly influenced by historical changes in species habitats. Whether such patterns are common to co‐distributed taxa may depend on the extent to which species vary in ecology and vagility. We investigated whether broad‐scale phylogeographic patterns common to a number of small‐bodied vertebrate and invertebrate species in eastern Australian forests were reflected in the population genetic structure of an Australo‐Papuan forest marsupial, the red‐legged pademelon (Macropodidae: Thylogale stigmatica). Strong genetic structuring of mtDNA haplotypes indicated the persistence of T. stigmatica populations across eastern Australia and southern New Guinea in Pleistocene refugial areas consistent with those inferred from studies of smaller, poorly dispersing species. However, there was limited divergence of haplotypes across two known historical barriers in the northeastern Wet Tropics (Black Mountain Barrier) and coastal mideastern Queensland (Burdekin Gap) regions. Lack of divergence across these barriers may reflect post‐glacial recolonization of forests from a large, central refugium in the Wet Tropics. Additionally, genetic structure is not consistent with the present delimitation of subspecies T. s. wilcoxi and T. s. stigmatica across the Burdekin Gap. Instead, the genetic division occurs further to the south in mideastern Queensland. Thus, while larger‐bodied marsupials such as T. stigmatica did persist in Pleistocene refugia common to a number of other forest‐restricted species, species‐specific local extinction and recolonization events have resulted in cryptic patterns of genetic variation. Our study demonstrates the importance of understanding individualistic responses to historical climate change in order to adequately conserve genetic diversity and the evolutionary potential of species.     

Checkerboard Patterns,Interspecific Competition,and Extinction: Lessons from Distribution Patterns of Tarsiers (<Emphasis Type="Italic">Tarsius</Emphasis>) and Slow Lorises (<Emphasis Type="Italic">Nycticebus</Emphasis>) in Insular Southeast Asia

Tarsiers (Tarsius) and slow lorises (Nycticebus) are the only extant nocturnal primates occurring in Southeast Asia. Harcourt (1999) hypothesized that in insular Southeast Asia, slow lorises and tarsiers showed a checkerboard distribution on 12 small (<12,000 km²) islands, i.e., only one or the other occurs, and attributed this to extreme levels of competition between these 2 largely faunivorous primates. Further, he predicted slow lorises were able to persist on smaller islands than tarsiers. We re-evaluated these findings using an expanded dataset including 49 islands where tarsiers or slow lorises occur. Tarsiers and slow lorises live on islands of similar size (median size of ca. 300–900 km²), and both taxa inhabit an equal proportion of small, medium, and large islands. On small islands within their area of sympatry tarsiers occur on 1 island, slow lorises on 8, both genera on 3, and we can assume they have become extinct from 11 small islands since the Last Glacial Maximum. Sizes of islands where tarsiers or slow lorises have become extinct do not differ from islands where they are still extant. We show that slow lorises occur on more islands in insular Southeast Asia than perhaps previously assumed, but these islands are not smaller on average than islands where tarsiers occur. A checkerboard distribution between these taxa is not evident. More studies are needed at the macroecological level to assess the importance of biogeographic history in explaining their present-day distribution patterns.     

Invasion of the Hawaiian Islands by a parasite infecting imperiled stream fishes

Points of origin and pathways of spread are often poorly understood for introduced parasites that drive disease emergence in imperiled native species. Co‐introduction of parasites with non‐native hosts is of particular concern in remote areas like the Hawaiian Islands, where the introduced nematode Camallanus cotti has become the most prevalent parasite of at‐risk native stream fishes. In this study, we evaluated the prevailing hypothesis that C. cotti entered the Hawaiian Islands with poeciliid fishes from the Americas, and spread by translocation of poeciliid hosts across the archipelago for mosquito control. We also considered the alternative hypothesis of multiple independent co‐introductions with host fishes originating from Asia. We inferred conduits of introduction and spread of C. cotti across the archipelago from geographic patterns of mtDNA sequence variation and allelic variation across 11 newly developed microsatellite markers. The distribution of haplotypes suggests that C. cotti spread across the archipelago following an initial introduction on O'ahu. Approximate Bayesian Computation modeling and allelic variation also indicate that O'ahu is the most likely location of introduction, from which C. cotti dispersed to Maui followed by spread to the other islands in the archipelago. Evidence of significant genetic structure across islands indicates that contemporary dispersal is limited. Our findings parallel historical records of non‐native poeciliid introductions and suggest that remediating invasion hotspots could reduce the risk of infection in native stream fishes, which illustrates how inferences on parasite co‐introductions can improve conservation efforts by guiding responses to emerging infectious disease in species of concern.     

Colonization history of Galapagos giant tortoises: Insights from mitogenomes support the progression rule

Galapagos giant tortoises (Chelonoidis spp.) are a group of large, long-lived reptiles that includes 14 species, 11 of which are extant and threatened by human activities and introductions of non-native species. Here, we evaluated the phylogenetic relationships of all extant and two extinct species (Chelonoidis abingdonii from the island of Pinta and Chelonoidis niger from the island of Floreana) using Bayesian and maximum likelihood analysis of complete or nearly complete mitochondrial genomes. We also provide an updated phylogeographic scenario of their colonization of the Galapagos Islands using chrono-phylogenetic and biogeographic approaches. The resulting phylogenetic trees show three major groups of species: one from the southern, central, and western Galapagos Islands; the second from the northwestern islands; and the third group from the northern, central, and eastern Galapagos Islands. The time-calibrated phylogenetic and ancestral area reconstructions generally align with the geologic ages of the islands. The divergence of the Galapagos giant tortoises from their South American ancestor likely occurred in the upper Miocene. Their diversification on the Galapagos adheres to the island progression rule, starting in the Pleistocene with the dispersal of the ancestral form from the two oldest islands (San Cristóbal and Española) to Santa Cruz, Santiago, and Pinta, followed by multiple colonizations from different sources within the archipelago. Our work provides an example of how to reconstruct the history of endangered taxa in spite of extinctions and human-mediated dispersal events and provides a framework for evaluating the contribution of colonization and in situ speciation to the diversity of other Galapagos lineages.