Island area and species diversity in the southwest Pacific Ocean: is the lizard fauna of Vanuatu depauperate?

One island group suggested to be an exception to the species–area Relationship is the Vanuatu Archipelago, a group of 13 large and 80 small islands in the southwest Pacific Ocean. To test the hypothesis that the lizard fauna of the Vanuatu Archipelago does not meet the predictions of the species-area relationship, and thus is depauperate, we compare diversity among several island groups in the southwest Pacific: Fiji, the Loyalty Islands, New Caledonia, Samoa, the Solomon Islands, Tonga, and Vanuatu. We found that the lizard diversity of Vanuatu meets the pattern of diversity predicted by the species-area relationship. The Solomon Islands, the largest and least isolated oceanic archipelago considered, has the greatest species diversity and endemism of the oceanic islands. Inclusion or exclusion of island groups based on factors such as geologic history or faunal source affects the strength of the relationship between diversity, area, and history of emergence, and influences perceptions of diversity within individual archipelagos. In addition to island size, factors such evolutionary time scale, speciation, and archipelago complexity influence species richness on islands.     

Butterfly biogeography and endemism on tropical Pacific islands

Butterfly distributions on 26 tropical Pacific archipelagos were analysed to examine the effects of geography on diversity and endemism. The total butterfly fauna for each archipelago was divided into continental (found also on continental areas), Pacific (found within more than one archipelago but not outside of the study area), and endemic species (restricted to a single archipelago). Numbers and proportions of each species were related to eight geographic variables by stepwise multiple linear regression analysis. Total area of an archipelago and distance from other land masses were important predictors of the number of species within an archipelago. Proportions of butterfly species in each category were related differently to the geographic variables, with endemism being promoted by the number of large islands within an archipelago. Relative to birds, butterflies have been less successful in colonizing remote archipelagos and have much lower levels of endemism. Even if colonization is successful, butterfly speciation may be constrained by the mechanics of coevolution with available host plants.     

Dispersal and speciation of skinks among archipelagos in the tropical Pacific Ocean

Summary We examined the potential effects of geography on the distribution and speciation of skinks on tropical Pacific archipelagos. The entire tropical Pacific skink fauna was divided into continental (found also in continental areas), Pacific (endemic to the study area but found within more than one archipelago) and endemic (found within only one archipelago) species categories. The number and proportion of skinks within each species category were determined for each of the 27 archipelagos in the study area. Nine geographic variables reflecting archipelago size, isolation and elevation were estimated for each archipelago. Principal components analysis was used to reduce the nine variables to three uncorrelated composite variables that were interpreted as representing archipelago size, isolation and elevation. Numbers and proportions of skinks in each category within an archipelago were related to the composite geographic variables using multiple linear regression analysis. Archipelago size and isolation were important predictors of both skink diversity and endemism. Results were then compared to diversity and endemism of birds within the study area. Skinks showed an archipelago-wide level of endemism similar to that of birds. On an archipelago by archipelago basis, however, large differences between birds and skinks were evident. In particular, the New Caledonia skink fauna was much more endemic than that of birds. The bird faunas of Hawaii and the Marquesas were nearly completely endemic, while no endemic skinks occurred in these two archipelagos. These differences presumably reflect the relative dispersal powers of skinks and birds and, consequently, rates of colonization and speciation. Differences may also be due partly to morphological conservatism among isolated skink populations and the occurrence of cryptic species that have not yet been identified as separate species. The discovery of such cryptic species, however, is unlikely to increase the endemic skink fauna of Hawaii and other distant archipelagos to a level commensurate with that of birds. Differences in endemism between skinks and birds may also be due to unknown local ecological interactions.     

Molecular biogeography and diversification of the endemic terrestrial fauna of the Hawaiian Islands

Oceanic islands have played a central role in biogeography and evolutionary biology. Here, we review molecular studies of the endemic terrestrial fauna of the Hawaiian archipelago. For some groups, monophyly and presumed single origin of the Hawaiian radiations have been confirmed (achatinelline tree snails, drepanidine honeycreepers, drosophilid flies, Havaika spiders, Hylaeus bees, Laupala crickets). Other radiations are derived from multiple colonizations (Tetragnatha and Theridion spiders, succineid snails, possibly Dicranomyia crane flies, Porzana rails). The geographic origins of many invertebrate groups remain obscure, largely because of inadequate sampling of possible source regions. Those of vertebrates are better known, probably because few lineages have radiated, diversity is far lower and morphological taxonomy permits identification of probable source regions. Most birds, and the bat, have New World origins. Within the archipelago, most radiations follow, to some degree, a progression rule pattern, speciating as they colonize newer from older islands sequentially, although speciation often also occurs within islands. Most invertebrates are single-island endemics. However, among multi-island species studied, complex patterns of diversification are exhibited, reflecting heightened dispersal potential (succineids, Dicranomyia). Instances of Hawaiian taxa colonizing other regions are being discovered (Scaptomyza flies, succineids). Taxonomy has also been elucidated by molecular studies (Achatinella snails, drosophilids). While molecular studies on Hawaiian fauna have burgeoned since the mid-1990s, much remains unknown. Yet the Hawaiian fauna is in peril: more than 70 per cent of the birds and possibly 90 per cent of the snails are extinct. Conservation is imperative if this unique fauna is to continue shedding light on profound evolutionary and biogeographic questions.     

Review Oribatid mites of the Galápagos Islands – faunistics, ecology and speciation

The results from investigations on oribatid mites of the Galápagos archipelago during 10 years are presented. Samples were taken on all larger and most smaller islands of the archipelago, as well as in all vegetation zones and some special habitats such as grass or cactus litter and fumaroles. A total of 202 oribatid species belonging to 64 families were encountered; among them 81 species are new to science. The Oribatida occur from the littoral zone to the summit of the volcanoes. Diversity and abundance increases from the arid to the moister zones at higher elevations. Most species prefer moist habitats at medium to higher elevations of the islands, in some parts reaching remarkably high abundances (in the Scalesia zone of Santa Cruz approximately 18 000 individuals m–2). The species richness of an island depends on the altitude and number of available habitats rather than the area of the island. Many oribatid species on the Galápagos Islands have a wide biogeographical distribution. The majority originate from the Central and South American mainland, but several Pacific and even Holarctic elements were also found. In comparison with the species composition of the adjacent mainland, the oribatid mite fauna of the Galápagos Islands can be regarded as disharmonic. Sea surface transportation has been proved at least between the islands, which also applies to oribatid species living at higher elevations. Long distance dispersal can be assumed to be mainly hydrochorous. The oribatid fauna of the littoral and arid vegetation zones is presented in detail. Some species even inhabit such extreme habitats as warm fern litter in fumaroles or hot volcanic soils. Others were mainly found in or near agricultural areas, suggesting human introduction. Striking distribution patterns within the archipelago are discussed. The distribution of endemic forms in the genera Aeroppia, Scapheremaeus, Scheloribates and Cultrobates indicates both speciation on different islands, as well as speciation on the same island by occupying different habitats.     

Genetic structure of kestrel populations and colonization of the Cape Verde archipelago

Genetic diversity and population structure were studied in eight populations of the kestrel Falco tinnunculus to identify the genetic consequences of spatial distribution and to infer the colonization patterns of the Cape Verde archipelago. We studied genetic differentiation and gene flow among seven island populations and one mainland population using nine microsatellite loci. Within the archipelago, differentiation was strong and genetic diversity and heterozygosity were low but variable among populations. Two subspecies F. tinnunculus neglectus on the northwestern islands and F. tinnunculus alexandri on all the other islands were identified as genetically distinct units. F. t. alexandri could be further separated into two groups on eastern and southern islands. Populations are probably founded by birds originating from the mainland. Immigration is more likely to the eastern and southern populations, whereas the northwestern islands with the lowest genetic diversity and highest differentiation are likely to exhibit fewer founding events by immigrants. The number of founding events on each island may depend not only on geographical distance to neighbouring populations, but also on directional immigration due to the northeastern trade winds. This may explain differences in genetic differentiation and diversity between populations and subspecies and may enable allopatric speciation.     

Biogeography and systematics of endemic island damselflies: The Nesobasis and Melanesobasis (Odonata: Zygoptera) of Fiji

The study of island fauna has greatly informed our understanding of the evolution of diversity. We here examine the phylogenetics, biogeography, and diversification of the damselfly genera Nesobasis and Melanesobasis, endemic to the Fiji Islands, to explore mechanisms of speciation in these highly speciose groups. Using mitochondrial (COI, 12S) and nuclear (ITS) replicons, we recovered Garli ‐part maximum likelihood and Mrbayes Bayesian phylogenetic hypotheses for 26 species of Nesobasis and eight species/subspecies of Melanesobasis. Biogeographical patterns were explored using Lagrange and Bayes ‐Lagrange and interpreted through beast relaxed clock dating analyses. We found that Nesobasis and Melanesobasis have radiated throughout Fiji, but are not sister groups. For Nesobasis, while the two largest islands of the archipelago—Viti Levu and Vanua Levu—currently host two distinct species assemblages, they do not represent phylogenetic clades; of the three major groupings each contains some Viti Levu and some Vanua Levu species, suggesting independent colonization events across the archipelago. Our Beast analysis suggests a high level of species diversification around 2–6 Ma. Our ancestral area reconstruction (Rasp ‐Lagrange ) suggests that both dispersal and vicariance events contributed to the evolution of diversity. We thus conclude that the evolutionary history of Nesobasis and Melanesobasis is complex; while inter‐island dispersal followed by speciation (i.e., peripatry) has contributed to diversity, speciation within islands appears to have taken place a number of times as well. This speciation has taken place relatively recently and appears to be driven more by reproductive isolation than by ecological differentiation: while species in Nesobasis are morphologically distinct from one another, they are ecologically very similar, and currently are found to exist sympatrically throughout the islands on which they are distributed. We consider the potential for allopatric speciation within islands, as well as the influence of parasitic endosymbionts, to explain the high rates of speciation in these damselflies.     

Contemporary geography dominates butterfly diversity gradients within the Aegean archipelago (Lepidoptera: Papilionoidea, Hesperioidea)

Aim We compare the influence of contemporary geography and historical influences on butterfly diversity for islands in the Aegean archipelago. Location The Aegean archipelago (Greece) and two islands (Cyprus and Megisti) in the Levantine Sea. Methods Thirty‐one islands were examined. Data are taken from own surveys (Coutsis and Olivier) and from the literature. Stepwise multiple regression is used to determine relationships between species richness, frequency, rarity and endemicity against potential geographical predictors. Stepwise logit regression is used to determine geographical predictors of species incidence on islands. Inter‐island and inter‐species associations have been examined using multivariate ordination and clustering techniques. Results The Aegean butterfly fauna is characterized by decreasing diversity and rarity, and increasing homogeneity, from the periphery to the present geographical centre of the archipelago (Cyclades). Diversity and rarity are shown to relate closely to species richness, and species richness, in turn, is largely explained by contemporary geography, particularly the degree of isolation from the nearest mainland sources of Greece or Turkey, and island dimensions. Islands towards the centre of the archipelago are characterized by a group of mobile species (n ≥ 20 species) with extensive ranges across Europe; species that would have recolonized Santorini (Thira) following the VI6 eruption there c. 1630 bc . Endemic components, indicative of autochthonous evolutionary events, are few (5% of species are endemic) compared to known sedentary organisms (molluscs and isopods), but exceed those for more mobile animals (i.e. birds); their distribution is mainly confined to large isolated islands along the Aegean arc (i.e. Kriti) and in the Dodecanese group. Main conclusions Contemporary geography, i.e. processes currently operating in ecological time, dominates butterfly diversity gradients (species richness, frequency, rarity and incidence) in the archipelago. Two reasons are suggested to account for the lack of endemism and the pattern of decreasing diversity into the Cyclades. First, relict butterfly elements may have become extinct on all but a few larger islands, particularly from environmental changes since the Neolithic (fire and overgrazing). Second, colonization from the continental landmasses is ongoing with more mobile species transferring even to the most isolated islands.     

Patterns of cladogenesis in the venomous marine gastropod genus Conus from the Cape Verde islands

Isolated oceanic archipelagos are excellent model systems to study speciation, biogeography, and evolutionary factors underlying the generation of biological diversity. Despite the wealth of studies documenting insular speciation, few of them focused on marine organisms. Here, we reconstruct phylogenetic relationships among species of the marine venomous gastropod genus Conus from the Cape Verde archipelago. This small island chain located in the Central Atlantic hosts 10% of the worldwide species diversity of Conus. Analyses were based on mtDNA sequences, and a novel nuclear marker, a megalin-like protein, member of the low-density lipoprotein receptor gene family. The inferred phylogeny recovered two well-defined clades within Conus. One includes Cape Verde endemic species with larger shells, known as the "venulatus" complex together with C. pulcher from the Canary Islands. The other is composed of Cape Verde endemic and West Africa and Canary Island "small" shelled species. In both clades, nonendemic Conus were resolved as sister groups of the Cape Verde endemics, respectively. Our results indicate that the ancestors of "small" and "large" shelled lineages independently colonized Cape Verde. The resulting biogeographical pattern shows the grouping of most Cape Verde endemics in monophyletic island assemblages. Statistical tests supported a recent radiation event within the "small shell" clade. Using a molecular clock, we estimated that the colonization of the islands by the "small" shelled species occurred relatively close to the origin of the islands whereas the arrival of "large" shelled Conus is more recent. Our results suggest that the main factor responsible for species diversity in the archipelago may be allopatric speciation promoted by the reduced dispersal capacity of nonplanktonic lecithotrophic larvae.     

Habitat, isolation and the evolution of Madeiran Iandsnails

The Madeiran archipelago consists of Madeira itself, Porto Santo and the Deserta islands. On Madeira, the forest arid the coastal floral associations are so different that their faunas are effectively isolated and have undergone largely independent development. There are different faunal associations on the eastern peninsula and in the SE coastal region, which may have been separated from each other in the past. On Porto Santo, western and eastern hills have different faunas. Most observations on the fauna are compatible with evolution by allopatric speciation, consequent upon isolation on different islands or mountains, as opposed to parapatric or sympatric processes following disruptive selection. Some cases where the taxonomy is difficult to unravel may, however, indicate parapatric speciation; examples belong to the genera Discula and Heterostoma (Helicidae) and Amphorella (Ferussaciidae). Most evidence relating to species composition in communities is compatible with a balance of random immigration and extinction, rather than selective interaction, allowing clusters of similar sympatric species to accumulate. However, this impression may indicate that test procedures are insufficiently sensitive to detect interactions, and detailed ecological studies are required. Questions about speciation and distribution would be clarified if dates of divergence were established.     

Rotifers from the Balearic archipelago

Ecological and historical factors of the Balearic archipelago (Western Mediterranean Sea) result is some peculiarities to its rotifer fauna. A group of warm-stenothermous species are frequent seasonally in the islands. Among them the occurrence of Keratella procurva (Thorpe, 1912), a pantropical species, is remarkable, and this is found more or less abundantly on each island. We briefly describe the different habitat categories of the archipelago. Clustering the collected species we relate each community type to its habitat; in this sense we distinguish 4 community type in the islands corresponding to different environments. This work also summarizes all the obtainable data on the rotifer fauna of the Balearic archipelago, adding the results of the investigations done in the last survey (spring, 1989); a checklist, including 100 species from the archipelago, is shown (69 Majorca, 72 Minorca, 24 Ibiza, 25 Formentera). Some examples of pantropical distribution are given, and the biotic and abiotic factors which determine the colonization of the islands by these species are discussed.     

Evolution on oceanic islands: molecular phylogenetic approaches to understanding pattern and process

By their very nature oceanic island ecosystems offer great opportunities for the study of evolution and have for a long time been recognized as natural laboratories for studying evolution owing to their discrete geographical nature and diversity of species and habitats. The development of molecular genetic methods for phylogenetic reconstruction has been a significant advance for evolutionary biologists, providing a tool for answering questions about the diversity among the flora and fauna on such islands. These questions relate to both the origin and causes of species diversity both within an archipelago and on individual islands. Within a phylogenetic framework one can answer fundamental questions such as whether ecologically and/or morphologically similar species on different islands are the result of island colonization or convergent evolution. Testing hypotheses about ages of the individual species groups or entire community assemblages is also possible within a phylogenetic framework. Evolutionary biologists and ecologists are increasingly turning to molecular phylogenetics for studying oceanic island plant and animal communities and it is important to review what has been attempted and achieved so far, with some cautionary notes about interpreting phylogeographical pattern on oceanic islands.     

中街山列岛岩相潮间带大型底栖动物的群落格局

于2006年9月和2007年7月在中街山列岛4个有居民海岛岩相潮间带设立8个断面,对大型底栖动物的群落结构进行了调查。共获得大型底栖动物34种,其中腔肠动物1种,环节动物3种,甲壳动物10种,软体动物20种。优势种有日本笠藤壶(Tetraclita japonica)、紫贻贝(Mytilus edulis)和节蝾螺(Turb articulatus)等4种。以季节和站位为因子对多样性指数进行双因素方差分析,结果表明只有夏、秋季丰富度指数(P=0.028)存在显著差异,其他指数的方差分析没有显著差异(P>0.05)。3个潮带4个岛屿共12个站位的聚类分析和MDS标序分析表明,12个群落可分为3组(Ⅰ、Ⅱ和Ⅲ组)。大型底栖动物ABC曲线分析表明底栖动物群落尚未受到干扰。     

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

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

Colonization and diversification of Galápagos terrestrial fauna: a phylogenetic and biogeographical synthesis

Remote oceanic islands have long been recognized as natural models for the study of evolutionary processes involved in diversification. Their remoteness provides opportunities for isolation and divergence of populations, which make islands remarkable settings for the study of diversification. Groups of islands may share a relatively similar geological history and comparable climate, but their inhabitants experience subtly different environments and have distinct evolutionary histories, offering the potential for comparative studies. A range of organisms have colonized the Galápagos Islands, and various lineages have radiated throughout the archipelago to form unique assemblages. This review pays particular attention to molecular phylogenetic studies of Galápagos terrestrial fauna. We find that most of the Galápagos terrestrial fauna have diversified in parallel to the geological formation of the islands. Lineages have occasionally diversified within islands, and the clearest cases occur in taxa with very low vagility and on large islands with diverse habitats. Ecology and habitat specialization appear to be critical in speciation both within and between islands. Although the number of phylogenetic studies is continuously increasing, studies of natural history, ecology, evolution and behaviour are essential to completely reveal how diversification proceeded on these islands.     

How old is the Hawaiian biota? Geology and phylogeny suggest recent divergence.

This study quantifies long-term landscape changes in the Hawaiian archipelago relating to dispersal, speciation and extinction. Accounting for volcano growth, subsidence and erosion, we modelled the elevations of islands at time intervals of 0.5 Myr for the last 32 Myr; we also assessed the variation in the spacing of volcanoes during this period. The size, spacing and total number of volcanic islands have varied greatly over time, with the current landscape of large, closely spaced islands preceded by a period with smaller, more distantly spaced islands. Considering associated changes in rates of dispersal and speciation, much of the present species pool is probably the result of recent colonization from outside the archipelago and divergence within contemporary islands, with limited dispersal from older islands. This view is in accordance with abundant phylogenetic studies of Hawaiian organisms that estimate the timing of colonization and divergence within the archipelago. Twelve out of 15 multi-species lineages have diverged within the lifetime of the current high islands (last 5 Myr). Three of these, and an additional seven (mostly single-species) lineages, have colonized the archipelago within this period. The timing of colonization of other lineages remains uncertain.     

Determinants of alpha and beta vascular plant diversity in Mediterranean island systems: the Ionian islands,Greece

The Ionian archipelago is the second largest Greek archipelago after the Aegean, but the factors driving plant species diversity in the Ionian islands are still barely known. We used stepwise multiple regressions to investigate the factors affecting plant species diversity in 17 Ionian islands. Generalized dissimilarity modelling was applied to examine variation in the magnitude and rate of species turnover along environmental gradients, as well as to assess the relative importance of geographical and climatic factors in explaining species turnover. The values of the residuals from the ISAR log₁₀‐transfomed models of native and endemic taxa were used as a measure of island floristic diversity. Area was confirmed to be the most powerful single explanatory predictor of all diversity metrics. Mean annual precipitation and temperature, as well as shortest distance to the nearest island are also significant predictors of vascular plant diversity. The island of Kalamos constitutes an important plant diversity hotspot in the Ionian archipelago. The recent formation of the islands, the close proximity to the mainland source and the relatively low dispersal filtering of the Ionian archipelago has resulted in islands with a flora principally comprising common species and a low proportion of endemics. Small islands keep a key role in conservation of plant priority sites.     

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.