Phylogeography and molecular phylogeny of Macaronesian island Tarphius (Coleoptera: Zopheridae): why are there so few species in the Azores?

Aim We used a phylogenetic framework to examine island colonization and predictions pertaining to differentiation within Macaronesian Tarphius (Insecta, Coleoptera, Zopheridae), and explain the paucity of endemics in the Azores compared with other Macaronesian archipelagos. Specifically, we test whether low diversity in the Azores could be due to recent colonization (phylogenetic lineage youth), cryptic speciation (distinct phylogenetic entities within species) or the young geological age of the archipelago. Location Macaronesian archipelagos (Azores, Madeira and the Canary Islands), northern Portugal and Morocco. Methods Phylogenetic analyses of mitochondrial and nuclear genes of Tarphius beetles of the Azores, other Macaronesian islands and neighbouring continental areas were used to investigate the origin of island biodiversity and to compare patterns of colonization and differentiation. A comparative nucleotide substitution rate test was used to select the appropriate substitution rate to infer clade divergence times. Results Madeiran and Canarian Tarphius species were found to be more closely related to each other, while Azorean taxa grouped separately. Azorean taxa showed concordance between species and phylogenetic clades, except for species that occur on multiple islands, which segregated by island of origin. Divergence time estimates revealed that Azorean Tarphius are an old group and that the most recent intra‐island speciation event on Santa Maria, the oldest island, occurred between 3.7 and 6.1 Ma. Main conclusions Our phylogenetic approach provides new evidence to understand the impoverishment of Azorean endemics: (1) Tarphius have had a long evolutionary history within the Azores, which does not support the hypothesis of fewer radiation events due to recent colonization; (2) the current taxonomy of Azorean Tarphius does not reflect common ancestry and cryptic speciation is responsible for the underestimation of endemics; (3) intra‐island differentiation in the Azores was found only in the oldest island, supporting the idea that young geological age of the archipelago limits the number of endemics; and (4) the lack of evidence for recent intra‐island diversification in Santa Maria could also explain the paucity of Azorean endemics. Phylogenetic reconstructions of other species‐rich taxa that occur on multiple Macaronesian archipelagos will reveal whether our conclusions are taxon specific, or of a more general nature.     

Biogeography and genetic consequences of anagenetic speciation of Rhaphithamnus venustus (Verbenaceae) in the Juan Fernández archipelago,Chile: insights from AFLP and SSR markers

The genus R haphithamnus (Verbenaceae) consists of two species, one in South America and another endemic to the Juan Fernández archipelago, Chile. The genus represents an example of anagenetic speciation in which the island populations have diverged from their colonizing ancestors to the point where they are recognized as a distinct species. The island species R haphithamnus venustus differs from the continental R . spinosus primarily by floral traits associated with adaptation to hummingbird pollination. Two molecular markers, amplified fragment length polymorphisms (AFLPs) and microsatellites, were used to estimate divergence between the continental and insular species, and to compare diversity in the two species. The comparable or greater diversity in the insular species observed in some diversity indices of AFLPs would support the hypothesis that during the course of anagenetic speciation it has recovered from any reduction of genetic diversity associated with colonization of the archipelago. This pattern of comparable or higher diversity in insular species is seen with other instances of anagenetic speciation in the Juan Fernández archipelago. By contrast, the lower genetic diversity in the insular R . venustus found in microsatellites is likely to be the result of a founder effect from the original colonization of the archipelago; prior molecular studies suggest recent colonization of the Juan Fernández archipelago by R haphithamnus . The seeming non‐concordance between the present results and the widely accepted biogeography of R haphithamnus inferred from other data is discussed and an explanation is presented.     

Immigration history of amphidromous species on a Greater Antillean island

Aim To use molecular data to test for dispersal structuring in the immigration history of an amphidromous community on an island. Location The Caribbean island of Puerto Rico. Methods Mitochondrial DNA sequences were obtained from 11 amphidromous species, including shrimps, fish and a gastropod, sampled from throughout the island. The timing of population expansion (T_E) in each species was calculated using nucleotide variation and molecular clock dating methods. The order of species accumulation was then reconstructed (oldest to most recent estimate for T_E), and groups of species with non‐overlapping estimates for T_E were identified. The temporal span and average immigration rate for each group were calculated and compared with expectations of two previously published models of island immigration [the ‘dispersal‐structured model of island recolonization’ ( Whittaker & Jones, Oikos, 1994 , 69 , 524–529), which predicts short phases of rapid immigration followed by extended phases with relatively slow immigration rates; and the ‘colonization window hypothesis’ ( Carine, Taxon, 2005 , 54 , 895–903), which suggests that opportunities for island colonization are temporally constrained to discrete waves of colonization]. Results The molecular data indicated the immigration history of Puerto Rican amphidromous fauna from the late Pleistocene through the Holocene and identified two groups of species with non‐overlapping estimates for T_E and one group that overlapped with the other two groups. The temporal span, average immigration rate and lack of discreteness between all three groups indicated a continuum of immigration rather than distinct phases of species arrivals. Main conclusions This study did not support the expectations of the immigration models and suggested that amphidromous species from Puerto Rico comprise a single class of marine‐based dispersers. The immigration sequence we report probably reflects a recolonization chronology in this community, in keeping with the notion of species turnover through time. Four areas of future research into the immigration history of amphidromous species on islands are identified, and indicated the possibility that equilibrium processes govern long‐term community change in amphidromous biota on islands.     

Genomic and phenomic analysis of island ant community assembly

Island biodiversity has long fascinated biologists as it typically presents tractable systems for unpicking the eco‐evolutionary processes driving community assembly. In general, two recurring themes are of central theoretical interest. First, immigration, diversification, and extinction typically depend on island geographical properties (e.g., area, isolation, and age). Second, predictable ecological and evolutionary trajectories readily occur after colonization, such as the evolution of adaptive trait syndromes, trends toward specialization, adaptive radiation, and eventual ecological decline. Hypotheses such as the taxon cycle draw on several of these themes to posit particular constraints on colonization and subsequent eco‐evolutionary dynamics. However, it has been challenging to examine these integrated dynamics with traditional methods. Here, we combine phylogenomics, population genomics and phenomics, to unravel community assembly dynamics among Pheidole (Hymenoptera, Formicidae) ants in the isolated Fijian archipelago. We uphold basic island biogeographic predictions that isolated islands accumulate diversity primarily through in situ evolution rather than dispersal, and population genomic support for taxon cycle predictions that endemic species have decreased dispersal ability and demography relative to regionally widespread taxa. However, rather than trending toward island syndromes, ecomorphological diversification in Fiji was intense, filling much of the genus‐level global morphospace. Furthermore, while most endemic species exhibit demographic decline and reduced dispersal, we show that the archipelago is not an evolutionary dead‐end. Rather, several endemic species show signatures of population and range expansion, including a successful colonization to the Cook islands. These results shed light on the processes shaping island biotas and refine our understanding of island biogeographic theory.     

Bayesian Poisson tree processes and multispecies coalescent models shed new light on the diversification of Nawab butterflies in the Solomon Islands (Nymphalidae,Charaxinae, Polyura)

Butterflies of the genus Polyura form a widespread tropical group distributed from Pakistan to Fiji. The rare endemic Polyura epigenes Godman & Salvin, 1888 from the Solomon Islands archipelago represents a case of marked island polymorphism. We sequenced museum specimens of this species across its geographic range to study the phylogeography and genetic differentiation of populations in the archipelago. We used the Bayesian Poisson tree processes and multispecies coalescent models, to study species boundaries. We also estimated divergence times to investigate the biogeographic history of populations. Our molecular species delimitation and nuclear DNA network analyses unambiguously indicate that Malaita populations form an independent metapopulation lineage, as defined in the generalized lineage concept. This lineage, previously ranked as a subspecies, is raised to species rank under the name Polyura bicolor Turlin & Sato, 1995  stat. nov. Divergence time estimates suggest that this lineage split from its sister taxon in the late Pleistocene. At this time, the bathymetric isolation of Malaita from the rest of the archipelago probably prevented gene flow during periods of lower sea level, thereby fostering allopatric speciation. The combination of molecular species delimitation methods, morphological comparisons, and divergence time estimation is useful to study lineage diversification across intricate geographic regions.     

New insights into tateid gastropods and their radiation on Fiji based on anatomical and molecular methods (Caenogastropoda: Truncatelloidea)

The South Pacific archipelago of Fiji is characterized by a predominantly Indo‐Malesian flora and fauna. We provide a first systematic study on Fiji's tateid gastropods – previously classified as Hydrobiidae – describing 18 new species, combining morphological, anatomical, and molecular data. The molecular phylogeny of tateid gastropods based on 16S rRNA and cytochrome c oxidase subunit I (COI) showed that the species from Fiji were closer related to New Zealand than to Australian or New Caledonian taxa, which is rather exceptional. Performing an ancestral range reconstruction we inferred the colonization history across the two main islands. The radiation had its origin in southern Viti Levu, with a subsequent dispersal over the western and central parts of the island. The chronology of the radiation over eastern Viti Levu and Vanua Levu remained unresolved because of incomplete lineage sorting, a phenomenon typical for young radiations. © 2014 The Linnean Society of London     

Colonization and demographic expansion of freshwater fauna across the Hawaiian archipelago

It is widely accepted that insular terrestrial biodiversity progresses with island age because colonization and diversification proceed over time. Here, we assessed whether this principle extends to oceanic island streams. We examined rangewide mtDNA sequence variation in four stream‐dwelling species across the Hawaiian archipelago to characterize the relationship between colonization and demographic expansion, and to determine whether either factor reflects island age. We found that colonization and demographic expansion are not related and that neither corresponds to island age. The snail Neritina granosa exhibited the oldest colonization time (~2.713 mya) and time since demographic expansion (~282 kya), likely reflecting a preference for lotic habitats most prevalent on young islands. Conversely, gobioid fishes (Awaous stamineus, Eleotris sandwicensis and Sicyopterus stimpsoni) colonized the archipelago only ~0.411–0.935 mya, suggesting ecological opportunities for colonization in this group were temporally constrained. These findings indicate that stream communities form across colonization windows, underscoring the importance of ecological opportunities in shaping island freshwater diversity.     

Divergent evolutionary processes associated with colonization of offshore islands

Oceanic islands have been a test ground for evolutionary theory, but here, we focus on the possibilities for evolutionary study created by offshore islands. These can be colonized through various means and by a wide range of species, including those with low dispersal capabilities. We use morphology, modern and ancient sequences of cytochrome b (cytb) and microsatellite genotypes to examine colonization history and evolutionary change associated with occupation of the Orkney archipelago by the common vole (Microtus arvalis), a species found in continental Europe but not in Britain. Among possible colonization scenarios, our results are most consistent with human introduction at least 5100 bp (confirmed by radiocarbon dating). We used approximate Bayesian computation of population history to infer the coast of Belgium as the possible source and estimated the evolutionary timescale using a Bayesian coalescent approach. We showed substantial morphological divergence of the island populations, including a size increase presumably driven by selection and reduced microsatellite variation likely reflecting founder events and genetic drift. More surprisingly, our results suggest that a recent and widespread cytb replacement event in the continental source area purged cytb variation there, whereas the ancestral diversity is largely retained in the colonized islands as a genetic ‘ark’. The replacement event in the continental M. arvalis was probably triggered by anthropogenic causes (land‐use change). Our studies illustrate that small offshore islands can act as field laboratories for studying various evolutionary processes over relatively short timescales, informing about the mainland source area as well as the island.     

Phylogeny and divergence time of island tiger beetles of the genus Cylindera (Coleoptera: Cicindelidae) in East Asia

To examine the diverse colonization histories in eight tiger beetle species of the genus Cylindera (Coleoptera: Cicindelidae) on the East Asian islands, we conducted phylogenetic analyses and divergence time estimation using mitochondrial cytochome oxidase subunit I (COI) and nuclear 28S rDNA sequences. The island fauna consisted of four subgenera: Apterodela, Cicindina, Ifasina, and Cylindera. Apterodela is a flightless group with large bodies, whereas the others are fliers with small bodies. In Apterodela, the divergence among endemic species in Taiwan, Japan, and the mainland was ancient (2.1–4.7 Mya), as expected from their flightlessness. Their dispersal might have occurred across the extended landmass in East Asia during the Pliocene. In the subgenus Cicindina, Cylindera elisae has spread throughout East Asia, from which an endemic species, Cylindera bonina, was derived on the oceanic Bonin Islands during the early Pleistocene (0.9 Mya). This indicates the significance of Cylindera bonina, which is currently confined to a single island, for conservation. In the subgenus Ifasina, Cylindera kaleea is widely distributed in East Asia, and its sister species Cylindera humerula, endemic to Okinawa Island, diverged 1.0 Mya, whereas Cylindera psilica on Taiwan and the Yaeyama Islands diverged approximately 0.8 Mya. In the subgenus Cylindera, the colonization of Cylindera gracilis in Japan from the mainland occurred during the last glacial period. With the exception of C. bonina, which likely colonized new territories by flight or drifting, other dispersal events might have used land connections that occurred repeatedly during the Pliocene and Pleistocene. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 715–727.     

Molecular phylogeography reveals island colonization history and diversification of western Indian Ocean sunbirds (Nectarinia: Nectariniidae)

We constructed a phylogenetic hypothesis for western Indian Ocean sunbirds (Nectarinia) and used this to investigate the geographic pattern of their diversification among the islands of the Indian Ocean. A total of 1309 bp of mitochondrial sequence data was collected from the island sunbird taxa of the western Indian Ocean region, combined with sequence data from a selection of continental (African and Asian) sunbirds. Topological and branch length information combined with estimated divergence times are used to present hypotheses for the direction and sequence of colonization events in relation to the geological history of the Indian Ocean region. Indian Ocean sunbirds fall into two well-supported clades, consistent with two independent colonizations from Africa within the last 3.9 million years. The first clade contains island populations representing the species Nectarinia notata, while the second includes Nectarinia souimanga, Nectarinia humbloti, Nectarinia dussumieri, and Nectarinia coquereli. With respect to the latter clade, application of Bremer's [Syst. Biol. 41 (1992) 436] ancestral areas method permits us to posit the Comoros archipelago as the point of initial colonization in the Indian Ocean. The subsequent expansion of the souimanga clade across its Indian Ocean range occurred rapidly, with descendants of this early expansion remaining on the Comoros and granitic Seychelles. The data suggest that a more recent expansion from Anjouan in the Comoros group led to the colonization of Madagascar by sunbirds representing the souimanga clade. In concordance with the very young geological age of the Aldabra group, the sunbirds of this archipelago have diverged little from the Madagascar population; this is attributed to colonization of the Aldabra archipelago in recent times, in one or possibly two or more waves originating from Madagascar. The overall pattern of sunbird radiation across Indian Ocean islands indicates that these birds disperse across ocean barriers with relative ease, but that their subsequent evolutionary success probably depends on a variety of factors including prior island occupation by competing species.     

Genetic structure of house mouse (Mus musculus Linnaeus 1758) populations in the Atlantic archipelago of the Azores: colonization and dispersal

We analyzed the genetic structure and relationships of house mouse (Mus musculus) populations in the remote Atlantic archipelago of the Azores using nuclear sequences and microsatellites. We typed Btk and Zfy2 to confirm that the subspecies Mus musculus domesticus was the predominant genome in the archipelago. Nineteen microsatellite loci (one per autosome) were typed in a total of 380 individuals from all nine Azorean islands, the neighbouring Madeiran archipelago (Madeira and Porto Santo islands), and mainland Portugal. Levels of heterozygosity were high on the islands, arguing against population bottlenecking. The Azorean house mouse populations were differentiated from the Portuguese and Madeiran populations and no evidence of recent migration between the three was obtained. Within the Azores, the Eastern, Western, and Central island groups tended to act as separate genetic units for house mice, with some exceptions. In particular, there was evidence of recent migration events among islands of the Central island group, whose populations were relatively undifferentiated. Santa Maria had genetically distinctive mice, which may relate to its colonization history. © 2013 The Linnean Society of London     

Biogeographically dynamic genetic structure bridging two continents in the monotypic Central American rodent Ototylomys phyllotis

Central America is an ideal region in which to study patterns of historical divergence and population genetic differentiation, because of its extraordinarily dynamic biogeographical, tectonic, and climatic history. The rodent Ototylomys phyllotis is the only extant species of the genus Ototylomys and is distributed within this region from the Isthmus of Tehuantepec, Mexico, to central Costa Rica, offering an excellent opportunity to study spatial and temporal patterns of population structure of the species and to explain the ecological and evolutionary processes responsible for those patterns. We estimated the genetic diversity and structure within and between populations of O. phyllotis, times of divergence, and migration patterns using mitochondrial DNA and a comprehensive combination of phylogenetic and phylogeographical computational analyses. Our results support monophyly of the genus Ototylomys. We identified three major phylogeographical lineages within O. phyllotis that are linked to its diversification and coincide with the main geological features that shaped Middle America. The origin of the genus was before 3.35 Mya, prior to the Great American Biotic Interchange (GABI), and its initial occurrence was near the centre of its current distribution (Honduras/El Salvador), from which it later spread (3.20–2.84 Mya) following a series of GABI pulses. The species showed an initial northward dispersal to the Chiapas and Guatemala highlands (2.27 Mya) followed by diversification. A later dispersal (1.82 Mya) occurred toward both the south (Nicaragua, Costa Rica) and the north (Belize). The Yucatan peninsula was colonized (0.8 Mya) by individuals from Belize. Extremely high radiation and range expansion occurred throughout the entire range, the highest of which was in the Yucatan peninsula (0.125 Mya). © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 107 , 593–610.     

Phylogeographical study of the alpine plant Cassiope lycopodioides (Ericaceae) suggests a range connection between the Japanese archipelago and Beringia during the Pleistocene

Given that East Asia is located south‐west of Beringia and was less glaciated during the Pleistocene, species at higher latitudes were able to expand their range in this region during climate cooling. Although southward migration is an inevitable colonization process, the biogeographical history of the disjunct ranges of higher‐latitude species in East Asia has been investigated less extensively. Here, we assess whether their disjunct distributions in the Japanese archipelago connected sufficiently with Beringia or persisted in isolation following their establishment. Sequences of nine nuclear loci were determined for Cassiope lycopodioides (Ericaceae) from the Japanese archipelago as well as its surrounding areas, Kamchatka and Alaska. According to the geographical pattern of genetic diversity, the northern populations from Kamchatka to the northern part of the Japanese archipelago were similar genetically and were differentiated from populations in central Japan. Our study suggested that the distribution of C. lycopodioides was connected between the northern part of the Japanese archipelago and south‐western Beringia due to Pleistocene climate cooling. Conversely, central Japan harboured a disjunct range after its establishment. These inferences suggest that widespread range expansion in northern East Asia was plausible for species distributed in Beringia. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 497–509.     

Comparative phylogeography of the centipedes <Emphasis Type="Italic">Cryptops pictus</Emphasis> and <Emphasis Type="Italic">C. niuensis</Emphasis> (Chilopoda) in New Caledonia,Fiji and Vanuatu

The South Pacific is a biodiverse region of extreme evolutionary importance because it harbors ancient lineages and recent radiations. However, few population-level studies of genetic variation have been conducted in the land masses of this region. Likewise, the number of population-level studies using myriapods as models is extremely small. In this article, we compare the genetic structure of two species of centipedes in the genus Cryptops endemic to the South Pacific, one from a continental island, the other from oceanic islands. The level of genetic diversity and structure in C. pictus, a species endemic to New Caledonia, is much higher than in C. niuensis in Fiji and Vanuatu, despite the fact that C. niuensis is spread across two different archipelagos and several islands. The most likely explanation is the relatively young age of the remnants of the Vitiaz Arc (Fiji and Vanuatu) compared to New Caledonia. Using the emergence of Fiji-Vanuatu as a calibration point, C. pictus is estimated to have diverged by 23.4 Mya (upper 95% confidence interval) with a mean estimate of 11.7 Mya versus the 9.7 Mya of C. niuensis. Considering the absence of shared sequences between specimens from different sampling sites and the high genetic structuring within populations, C. pictus appears to be an ideal candidate to assess historical processes at a micro-evolutionary scale in New Caledonia.     

Phylogenetic analyses of Tolpis Adans. (Asteraceae) reveal patterns of adaptive radiation,multiple colonization and interspecific hybridization

The plant genus Tolpis (Asteraceae) has been the subject of several investigations on the evolution of oceanic island plants. Its insular species were utilized in studies of artificial hybrid fertility, testing the validity of Baker’s law, the application of DNA barcodes, and the phylogenetic utility of inter‐simple sequence repeat markers. Despite this considerable interest in Tolpis, little is known about its phylogenetic history. Past investigations were unable to resolve most of the interspecific relationships, especially within the Canary Islands, where the genus is particularly diverse. Incomplete taxon sampling, the use of ambiguous outgroups and the limited utility of slowly evolving chloroplast DNA markers precluded detailed reconstructions. The present investigation presents a comprehensive molecular phylogeny of Tolpis. By utilizing highly variable nuclear DNA markers and a comprehensive taxon set, we have resolved the majority of interspecific relationships in the genus. Evaluations of competing tree topologies and ancestral area reconstructions complemented the analyses. Our results highlight the presence of three dominant mechanisms of island plant evolution—island colonization, adaptive radiation and interspecific hybridization—in Tolpis: (i) the extant distribution of the genus is the result of two independent colonization pathways, (ii) Tolpis has colonized at least one archipelago multiple times, (iii) the present insular diversity is the product of adaptive radiation, (iv) potential hybridization was detected between species now inhabiting different islands and archipelagoes, indicating sympatric historical distributions, and (v) several undescribed species await taxonomic recognition.     

Genetic structure of Micromeria (Lamiaceae) in Tenerife,the imprint of geological history and hybridization on within‐island diversification

Geological history of oceanic islands can have a profound effect on the evolutionary history of insular flora, especially in complex islands such as Tenerife in the Canary Islands. Tenerife results from the secondary connection of three paleo‐islands by a central volcano, and other geological events that further shaped it. This geological history has been shown to influence the phylogenetic history of several taxa, including genus Micromeria (Lamiaceae). Screening 15 microsatellite markers in 289 individuals representing the eight species of Micromeria present in Tenerife, this study aims to assess the genetic diversity and structure of these species and its relation with the geological events on the island. In addition, we evaluate the extent of hybridization among species and discuss its influence on the speciation process. We found that the species restricted to the paleo‐islands present lower levels of genetic diversity but the highest levels of genetic differentiation suggesting that their ranges might have contracted over time. The two most widespread species in the island, M. hyssopifolia and M. varia, present the highest genetic diversity levels and a genetic structure that seems correlated with the geological composition of the island. Samples from M. hyssopifolia from the oldest paleo‐island, Adeje, appear as distinct while samples from M. varia segregate into two main clusters corresponding to the paleo‐islands of Anaga and Teno. Evidence of hybridization and intraspecific migration between species was found. We argue that species boundaries would be retained despite hybridization in response to the habitat's specific conditions causing postzygotic isolation and preserving morphological differentiation.     

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.     

Genetic diversification of the Kanehira bitterling Acheilognathus rhombeus inferred from mitochondrial DNA,with comments on the phylogenetic relationship with its sister species Acheilognathus barbatulus

The Kanehira bitterling, Acheilognathus rhombeus, is a freshwater fish, discontinuously distributed in western Japan and the Korean Peninsula. Unusually among bitterling it is an autumn-spawning species and shows developmental diapause. Consequently, the characterization of its evolutionary history is significant not only in the context of the fish assemblage of East Asia, but also for understanding life-history evolution. This study aimed to investigate the phylogeography of A. rhombeus and its sister species Acheilognathus barbatulus, distributed in China, using a mitochondrial analysis of the ND1 gene from 311 samples collected from 50 localities in Japan and continental Asia. Phylogenetic analysis revealed that A. barbatulus is included in A. rhombeus and genetically closer to Japanese A. rhombeus than to Korean A. rhombeus. Divergence of Korean A. rhombeus and A. barbatulus from Japanese A. rhombeus was estimated to be from the late Pliocene (3.44 Mya) and the early Pleistocene (1.98 Mya), respectively. Each event closely coincided with the time of the Japan Sea opening. Japanese A. rhombeus comprised seven lineages: three in Honshu and four in Kyushu. One lineage in central Kyushu was genetically closer to the Honshu lineages than to other lineages in northern Kyushu. Divergence of Japanese lineages was estimated to be from the early to middle Pleistocene (0.55–0.93 Mya), during a period of geological and paleoclimatic change, including volcanic activity. Population expansion in the late Pleistocene (<0.10 Ma) was suggested in many of the lineages, which accords with other freshwater fishes. Biogeographically the ancestral A. rhombeus/A. barbatulus was likely to have repeatedly colonized Japan from the continent through land bridges in the late Pliocene and the early Pleistocene. However, the close genetic relationship between Japanese A. rhombeus and A. barbatulus suggests another possibility, with the second colonization occurring in reverse, from Japan to China. The small genetic distance between them indicates that the colonization occurred later than colonization events of other freshwater fishes, including other bitterling species.     

Les environnements du Vanuatu de l’Holocène à nos jours : un état des lieux des connaissances

The first Vanuatu archaeological site discovery dates back to the 1960s, then the scientific and archaeological knowledge of this archipelago have increased. Nevertheless, the human activities–palaeoenvironmental changes relation in Vanuatu is still a matter of debate. In order to better determine this relation, reviewing the literature on past and actual environment is needed. This paper presents here a state of knowledge on Vanuatu geology, climate, biodiversity and archaeology, from the mid-Holocene to our present-day. The young archipelago results from strong volcanic and tectonic activity, due to the active subduction zone between Australian and Pacific plates. The atmosphere–ocean interactions determine the wet and dry seasons. Fauna and flora are principally derived from Southeast Asia. The occurrence of several species and subspecies endemic to the archipelago, if not to some islands, is probably due to the Vanuatu isolated location, the size of the islands and the climatic gradient between north and south. The first human populations reached the archipelago around 3200 yr BP. The Lapita culture, characterized by decorated potteries, is shared across the whole Vanuatu between 3200 and 2900 yr BP. After 2900 yr BP, cultures differ from one island to another. After 600 yr BP, the Polynesian culture dominates in Vanuatu.     

Large‐scale introduction of the Indo‐Pacific damselfish Abudefduf vaigiensis into Hawai'i promotes genetic swamping of the endemic congener A. abdominalis

Hybridization in the ocean was once considered rare, a process prohibited by the rapid evolution of intrinsic reproductive barriers in a high‐dispersal medium. However, recent genetic surveys have prompted a reappraisal of marine hybridization as an important demographic and evolutionary process. The Hawaiian Archipelago offers an unusual case history in this arena, due to the recent arrival of the widely distributed Indo‐Pacific sergeant (Abudefduf vaigiensis), which is hybridizing with the endemic congener, A. abdominalis. Surveys of mtDNA and three nuclear loci across Hawai'i (N = 396, Abudefduf abdominalis and N = 314, A. vaigiensis) reveal that hybridization is significantly higher in the human‐perturbed southeast archipelago (19.8%), tapering off to 5.9% in the pristine northwest archipelago. While densities of the two species varied throughout Hawai'i, hybridization was highest in regions with similar species densities, contradicting the generalization that the rarity of one species promotes interspecific mating. Our finding of later generation hybrids throughout the archipelago invokes the possibility of genetic swamping of the endemic species. Exaptation, an adaptation with unintended consequences, may explain these findings: the endemic species has transient yellow coloration during reproduction, whereas the introduced species has yellow coloration continuously as adults, in effect a permanent signal of reproductive receptivity. Haplotype diversity is higher in Hawaiian A. vaigiensis than in our samples from the native range, indicating large‐scale colonization almost certainly facilitated by the historically recent surge of marine debris. In this chain of events, marine debris promotes colonization, exaptation promotes hybridization, and introgression invokes the possible collapse of an endemic species.