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

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

Multiple levels of allopatric divergence in the endemic Philippine fruit bat Haplonycteris fischeri (Pteropodidae)

As part of a larger comparative phylogeographical study of Philippine fruit bats, I used fragments of the mitochondrial genes cytochrome  b and ND2 to investigate phylogeography and diversification in Haplonycteris fischeri , a pteropodid bat endemic to the Philippines but widespread within the archipelago. Genetic diversity in H. fischeri was extremely high in these commonly studied genes, with 101 unique haplotypes in 123 sequenced individuals, although small, continuously isolated islands had less diversity than had large island complexes. Seven monophyletic groups and one paraphyletic group were restricted to individual islands, groups of islands, or parts of islands. Each Pleistocene island complex had a single resident monophyletic lineage; these five groups were separated by approximately 6–8% sequence divergence and apparently have been diverging for 4–6 Myr. Within island groups, monophyletic lineages on some individual islands suggest that current ocean channels have also been barriers to gene flow; in some cases, multiple allopatric clades were present on single islands. Basal divergence dates were estimated to be in the early Pliocene, and most diversification was apparently connected to the ongoing geological evolution of the Philippines. Geological history and current geography interact with ecology to cause substantial genetic differentiation within this primary forest-specialist species.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 329–349.     

How important were stepping stones in the colonization of Krakatau?

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

Both Recent and Pleistocene geography determine animal distributional patterns in the Tuscan Archipelago

Cluster analyses, Parsimony Analysis of Endemicity, species–area relationships, Mantel tests, partial Mantel tests and levels of endemism were used to investigate the influence of current geography and paleogeography on the composition of the fauna of the Tuscan Islands, a small archipelago in the Mediterranean. The composition of terrestrial mollusc and tenebrionid beetle faunas was prevalently influenced by Pleistocene conditions. The composition of butterfly (and possibly reptile) island fauna was influenced by both present (Holocene) and Pleistocene factors. Finally, chrysids were influenced only by present factors. On the whole, both present and historical factors concurred to determine present distributional patterns, but their relative influence varied according to the mobility of the group considered. Results provided here give some insights into the treatment of endemics to elucidate historical relationships among areas and add further momentum to a general model of biotic dynamics in landbridge archipelagos.     

Wallace’s line revisited: has vicariance or dispersal shaped the distribution of Malesian hawkmoths (Lepidoptera: Sphingidae)?

Large-scale biogeographical patterns of Sphingidae in the Malesian archipelago are investigated, using Geographic Information System-supported estimates of species ranges determined from a compilation of specimen-label databases. Distribution maps for all species and checklists for 114 islands are presented at http://www.sphingidae-sea.biozentrum.uni-wuerzburg.de . Phenetic analyses of island faunas reveal 'textbook'-like patterns of biogeographical associations in the region, with Wallace's line emerging as the main faunal discontinuity in the region. Further analyses, using partial Mantel statistics, indicate that historical features of geography (such as land bridges formed during periods of lower sea levels) are an important determinant of faunal similarity, but recent dispersal can also explain a significant portion of present day hawkmoth distributions. Faunal turnover in relation to distance between islands is steeper for Smerinthinae than for other subfamilies, indicating different dispersal abilities among higher taxonomic groups. These differences are discussed in the light of life-history differences between the subfamilies. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 455–468.     

Implications of contrasting patterns of genetic variability in two vespertilionid bats from the Indonesian archipelago

Wallacean island populations of two Vespertilioninae bats, Myotis muricola and Scotophilus kuhlii , which have similar geographical ranges, showed marked contrast in the amount and pattern of genetic variation. Within islands, genetic variation was on average much higher in M. muricola but declined from west to east, whereas all populations of S. kuhlii had uniformly low levels of genetic variation by mammalian standards. S. kuhlii showed little genetic differentiation between islands and estimates of gene flow were substantial whereas island populations of M. muricola differed markedly and there was a strong isolation-by-distance effect associated with the extent of the sea crossing between islands. Furthermore, the lower mean heterozygosity and small genetic distances between eastern island populations of M. muricola is evidence that there has been a bottleneck associated with the colonization of this area. The attenuation of genetic diversity to the east is also seen in some other mammalian species and may indicate limits to dispersal and have implications for species management. The patterns of variability in S. kuhlii may be a consequence of its strong dispersal capacity and close association with human activity, which, together with other factors, suggest a panmictic population.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 421–431.     

Phylogeography of the land snail Albinaria hippolyti (Pulmonata: Clausiliidae) from Crete, inferred from ITS-1 sequences

The polytypic Cretan land snail Albinaria hippolyti has a range that is partly fragmented and partly subdivided by hybrid zones. For this reason, it has served as a model species for investigating speciation and radiation in Mediterranean Clausiliidae. The first internal transcribed spacer ( ITS-1 ) of the nuclear ribosomal DNA was sequenced in 20 populations of A. hippolyti and phylogenetically analysed using maximum parsimony. We employed a novel method involving logarithmic weighting of gaps and topological constraints based on bootstrap values. The resulting phylogeography suggests that the species has undergone a recent cycle of range expansion and range reduction. Speciation cannot be linked to major geological vicariance events in the Miocene and Pliocene, as has been suggested previously. The subspecies A. h. arthuriana appears unrelated to other A. hippolyti subspecies, which supports recent suggestions, based on morphology, to regard it as a separate species.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 317–326.     

The biogeography of land snails in the islands of the Gulf of Guinea

Data on land snail diversity in the Gulf of Guinea islands is presented and the biogeography of the Gulf of Guinea fauna is discussed with reference to island snail faunas in the other areas of the world. Although the land snail faunas of the four islands clearly have west and central African affinities, speciation events following rare colonizations have led to high rates of endemism at the species and generic levels. The influence of island size, altitude, isolation and other factors on the development of the land snail faunas is discussed. The effect of land use changes on the land snail fauna over the last 150 years is evaluated. Current threats include recent changes in agricultural practice leading to the destruction of snail habitats in both forest and plantation areas. The potential threat posed by introduced species is also discussed. The Gulf of Guinea snail faunas appear to be in relatively good health in comparison to many other island snail faunas. However, the uniqueness of the faunas makes it imperative that further ecological and taxonomic research is undertaken both to understand the processes by which they developed and to evaluate the current status of many species in terms of distribution and threats.     

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.     

Comparative phylogeography of six herpetofauna species in Cyprus: late Miocene to Pleistocene colonization routes

The colonization patterns of oceanic islands are often interpreted through transmarine dispersal. However, in islands with intense human activities and unclear geological history, this inference may be inappropriate. Cyprus is such an island, whose geotectonic evolution has not been clarified yet to the desired level for biogeographical reconstructions, leaving the questions of ‘how the Cypriote biota arrived’ and ‘does the dispersal have the formative role in patterns of its diversification’ unanswered. Here, we address these issues through a reconstruction of the evolutionary history of six herptiles (Ablepharus budaki, Ophisops elegans, Acanthodactylus schreiberi, Telescopus fallax, Pelophylax cf. bedriagae, and Hyla savignyi) by means of mitochondrial DNA (cytochrome b and 16S rRNA), applying a Bayesian phylogenetic, biogeographical, and chronophylogenetic analyses. The phylogeographical analyses show that the colonization history of those species in Cyprus started in the late Miocene and extended into the Pliocene and Pleistocene, with geodispersal, transmarine dispersal, and human‐mediated dispersal having their share in shaping the diversification of Cypriote herptiles. The revealed patterns could be divided into three biogeographical categories: old colonizers that arrived in Cyprus during the late Miocene or early Pliocene either by a land bridge (geodispersal) which connected Cyprus with the mainland or by transmarine dispersal, younger colonizers that reached the island through transmarine dispersal from the Middle East, and new settlers that arrived through human‐induced (voluntary or not) introductions. This work advances our knowledge of the biogeography of Cyprus and highlights the need to consider both geo‐ and transmarine dispersal when dealing with islands whose associations do not have a straightforward interpretation. © 2013 The Linnean Society of London     

Molecular phylogeny of the endemic Philippine rodent Apomys (Muridae) and the dynamics of diversification in an oceanic archipelago

We analysed the phylogenetic relationships of ten of the 13 known species of the genus Apomys using DNA sequences from cytochrome b . Apomys, endemic to oceanic portions of the Philippine archipelago, diversified during the Pliocene as these oceanic islands arose de novo . Several of the speciation events probably took place on Luzon or Mindanao, the two largest, oldest, and most topographically complex islands. Only one speciation event is associated with vicariance due to Pleistocene sea-level fluctuation, and a Pleistocene diversification model in which isolation is driven by sea-level changes is inconsistent with the data. Tectonic vicariance is nearly absent from the Philippines, in which tectonic coalescence plays a significant role. Most speciation events (about two-thirds) are associated with dispersal to newly developed oceanic islands. The data imply that the species have persisted for long periods, measured in millions of years after their origins; further implications therefore are that faunal turnover is very slow, and persistence over geological time spans is more prominent than repeated colonization and extinction. Neither the equilibrium nor the vicariance model of biogeography adequately encompasses these results; a model incorporating colonization, extinction, and speciation is necessary and must incorporate long-term persistence to accommodate our observations.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80 , 699–715.     

Australian parasitic Ogyris butterflies: east–west divergence of highly‐specialized relicts

Not all butterflies are innocuous plant‐feeders. A small number of taxa in the family Lycaenidae have graduated from mutualistic partnerships with ants to predatory or parasitic associations. These highly‐specialized life histories, involving butterfly larvae living inside ant colonies, are often associated with rarity and vulnerability to extinction. In the present study, we examined the evolutionary relationships of a poorly‐known group of seven taxa herein referred to as the idmo‐group within the Australian lycaenid genus Ogyris. The idmo‐group has a relictual distribution across southern Australia and includes taxa with highly‐specialized phytophagous and myrmecophagous life histories. A phylogeny based on mitochondrial DNA (cytochrome oxidase I and cytochrome b] and the nuclear DNA locus elongation factor 1α (EF1α), generally agrees with current taxonomy and supports the recent elevation of endangered taxon Ogyris halmaturia to full species status. The transition to myrmecophagy was dated to the mid‐Miocene (approximately 16 Mya), when southern Australia experienced a humid climate and extensive mesic biome. The arid Nullarbor Plain, a major biogeographical feature of central southern Australia, divides the remnants of this mesic biome into south‐eastern and south‐western isolates. Late‐Miocene to Pliocene divergence estimates for polytypic Ogyris species across the Nullarbor were older than estimates made for similarly distributed birds, butterflies, mammals, and reptiles, which mostly date to the Pleistocene. The concept of highly‐specialized life histories as evolutionary dead‐end strategies is well exemplified by the idmo‐group. Data compiled on the known extant subpopulations for idmo‐group taxa show that all of these extraordinary butterflies are scarce and several face imminent threat of extinction. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 473–484.     

Species richness, rarity and endemicity on Italian offshore islands: complementary signals from island-focused and species-focused analyses

Aims To investigate the relative explanatory power of source faunas and geographical variables for butterfly incidence, frequency, richness, rarity, and endemicity on offshore islands. Location The western Italian offshore islands (Italy and Malta). Methods Thirty‐one islands were examined. Data were taken from our own field surveys and from the literature. Two approaches were undertaken, described as island‐focused and species‐focused, respectively. Offshore islands were allocated to their neighbouring source landmasses (Italian Peninsula, Sicily and Sardinia–Corsica) and compared with each other for faunal attributes, source and island geography. Generalized linear and stepwise multiple regression models were then used to determine the relationships of island species richness, rarity and endemicity with potential geographical predictors and source richness, rarity, and endemicity (island‐focused). Species frequency and incidence were assessed in relation to geographical and source predictors using stepwise linear and logistic regression, and inter‐island associations were examined using K‐Means clustering and non‐metric scaling (species‐focused). Results The analysis reveals firm evidence for the influence of the nearest large landmass sources on island species assemblages, richness, rarity and endemicity. A clear distinction in faunal affinities occurs between the Sardinian islands and islands lying offshore from the Italian mainland and Sicily. Islands neighbouring these three distinct sources differ significantly in richness, rarity and endemicity. Source richness, rarity, and endemicity have explanatory power for island richness, rarity, and endemicity, respectively, and together with island geography account for a substantial part of the variation in island faunas (richness 59%, rarity 60% and endemicity 64%). Source dominates the logistic regression parameters predicting the incidence of island species [13 (38%) of 34 species that could be analysed]; three ecological factors (source frequency, flight period and maximal altitude at which species live) explained 75% of the variation in the occurrence of species on the islands. Species found more frequently on islands occurred more frequently at sources, had longer flight periods, and occurred at lower altitudes at the sources. The incidence of most species on islands (84%) is correctly predicted by the same three variables. Main conclusions The Italian region of the Mediterranean Sea has a rich butterfly fauna comprising endemics and rare species as well as more cosmopolitan species. Analysis of island records benefited from the use of two distinct approaches, namely island‐focused and species‐focused, that sift distinct elements in island and source faunas. Clear contemporary signals appear in island–source relationships as well as historical signals. Differences among faunas relating to sources within the same region caution against assuming that contemporary (ecological) and historical (evolutionary) influences affect faunas of islands in different parts of the same region to the same extent. The implications of source–island relationships for the conservation of butterflies within the Italian region are considered, particularly for the long‐term persistence of species.     

Parallel evolution in molar outline of murine rodents: the case of the extinct Malpaisomys insularis (Eastern Canary Islands)

The lava mouse Malpaisomys was part of Pleistocene and Holocene faunas of the eastern Canary Islands; it became extinct during historical times. In order to evaluate the evolutionary processes of this endemic species, we set out to identify its relationships with possible mainland relatives. Its dental morphology was compared to a set of fossil and modern murine rodents from various phylogenetic groups, characterized by different diets and dental patterns, using a quantitative method based on a Fourier analysis of the outline of the first upper and lower molars. This morphometric analysis identified different evolutionary grades that are independent of the phylogenetic group. The first cluster is associated with primitive, asymmetric dental outlines, the second with intermediate forms linked to a more herbivorous diet. Highly differentiated forms diverge not only from the second cluster but also from each other. Our investigations reveal Malpaisomys to have had an intermediate dental pattern, a result which confirms previous palaeoecological interpretations of this taxon. However, conclusions about its closest mainland relative remain tentative. Based on a comparison of dental size and shape, as well as geographical considerations, a possible mainland ancestor could be either Paraethomys (North-African Pliocene) or Occitanomys (South-western European Pliocene). Such results support the hypothesis of a Pliocene colonization event, in which case Malpaisomys would display a normal evolutionary rate in dental size and shape, in spite of the insular context. In contrast, a late colonization by a modern representative of the African fauna would imply exceptionally high evolutionary rates compared to the background morphological evolution and a decrease in size that is unlikely under insular conditions.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 555–572.     

Enhanced colour polymorphisms in island populations of the land snail Euhadra peliomphala

Variations in visible genetic polymorphisms are assumed to decrease in populations on small islands because of intense founder effects, genetic drift and inbreeding. However, we have found evidence of a marked enhancement of colour polymorphisms within populations on small oceanic islands that were colonized from the mainland. The source populations on the mainland of the land snail Euhadra peliomphala in four oceanic islands were estimated by phylogenetic analysis of mitochondrial DNA sequences. Diversity of shell colour was higher in the island populations than in the source populations on the mainland. In addition, the shell colour morphs differed not only among populations from different islands but also between the island populations and the source populations on the mainland. By contrast, no mtDNA variations were found in any of the island populations, even though the source populations possessed high mtDNA diversity. Thus, components of colour morphs changed in the island populations after their colonization, and colour polymorphisms are enhanced in these islands despite the loss of genetic variation. The above findings suggest that ecological mechanisms such as morphological release owing to a release from competition may overcome the tendency toward reduced genetic variation in islands to enhance the colour polymorphism.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 417–425.     

Fruits and seeds of Ruppia (Potamogetonaceae) from the Pliocene of Yushe Basin, Shanxi, northern China and their ecological implications

Numerous fruits and seeds of Ruppia are reported from the Upper Pliocene (2.3–3.5 Myr ago) Zhangcun Formation in Yushe Basin, Shanxi, northern China. They are the first fossil Ruppia from China and demonstrate the importance of fruit and seed fossils in recording genera not represented by fossil leaves. These Ruppia are characterized by possessing a small oval endocarp, smooth endocarp surface, distinct elliptical external depressions, distinct apical mucro, slightly curved seed shape and conspicuous globose hilum. A new species, R. yushensis Zhao, Collinson and Li, is described from these endocarp and seed features. Comparison with the two European Miocene species, R. palaeomaritima Negru and R. maritime-miocenica Szafer, indicates the existence at that time of three different geographical and stratigraphical species. R. yushensis constitutes the first Pliocene record of Ruppia and extends the range of fossils of this genus from Europe to eastern Asia. R. yushensis is the only aquatic plant in the uppermost middle part of the Zhangcun Formation. This monotypic occurrence indicates a brackish, clear, tranquil and shallow lake in this region in the Late Pliocene. The smooth endocarp surface further suggests a warm temperate or temperate palaeoclimate.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 317–329.     

Combining molecular, morphological and ecological data to infer species boundaries in a cryptic tropical pitviper

Few operational methods exist for delimiting species boundaries, and these usually require sampling strategies that are unrealistic for widespread organisms that occur at low densities. Here we apply molecular, morphological and ecological species delimitation criteria to a wide-ranging, fragmented group of Asian green pitvipers, the Popeia popeiorum complex. A mitochondrial DNA phylogeny for the group indicates two well-differentiated clades, corresponding mainly to northern and southern parts of its range. Strong phylogeographical structure within each clade suggests isolation in forest refugia during the Pliocene and a southward colonization of the Sunda islands during the Pleistocene. Multivariate analysis of morphological characters reveals a generally conserved pattern of geographical variation, incongruent with the recovered phylogenetic history. We compare groups delineated by mtDNA variation to morphological and ecological divisions in the complex, and discuss the implications of these for the taxonomy of the group. Discordance between species boundaries inferred from different criteria suggests that combining independent sources of data provides the most reliable estimation of species boundaries in organisms that are difficult to sample in large numbers.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 343–364.     

Palaeoclimatic variation, adaptation and biogeography of inversion polymorphisms in natural populations of Drosophila robusta

Studies of natural and sexual selection in wild populations of Drosophila have historically provided strong inference for the maintenance of inversion polymorphism. Analysis of geographical variation in the Drosophila robusta chromosomal data collected over more than 50 years from 133 natural populations across eastern North America has confirmed several north–south and east–west clines in the frequencies of some gene arrangements and linked arrangement combinations. Patterns of geographical variation, including several north–south clines, revealed by regression and spatial autocorrelation analyses are concordant with palaeoclimatic shifts, Pleistocene glaciations and historical changes in the composition of North American forest communities. Because D. robusta is a sap-breeder, using the microbe-infested sap exudates of a number of deciduous tree species in which they carry out their life cycle, shifts in climate and palaeovegetation types since the formation of the eastern deciduous forests in the Miocene are hypothesized to be major factors influencing patterns of inversion polymorphisms across the range of this drosophilid species. In areas where sharp deviations in frequencies have been observed, particularly in the mid-western and western portions of the range, these divisions parallel historical geographical disjunctions in the species range that have yet to promote divergence and species formation despite the long history of D. robusta in North America.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 395–411.     

Taxonomy and sexual dimorphism of a new species of Loxoconcha (Podocopida: Ostracoda) from the Pleistocene of the Japan Sea

A new ostracod, Loxoconcha kamiyai sp. nov. in the Family Loxoconchidae, is described from the Pleistocene Omma Formation of Japan. Its geological and geographical distributions suggest that this species was once endemic to the Japan Sea, where it would have evolved until the Pliocene. Since the early Pleistocene, this species would have become extinct within this marginal sea during glacial maxima, probably due to its narrower salinity tolerances and geographical distributions than those of extant species inhabiting the euryhaline environments in other seas. The distributional patterns of pore systems in this species strongly suggest its closest phylogenetic affinities to a living species, Loxocorniculum mutsuense . These two species show a unique adult sexual dimorphism in the anterior element of the hingement. Taking the female hingement morphology as a standard, the male hingement can be explained in terms of heterochrony, i.e. paedomorphosis. Sexual hingement dimorphism with paedomorphosis occurs in only one phylogenetic group of the genus Loxoconcha , which is distinguished by the ontogenetic distributional patterns of pore systems. This morphology may represent relict primitive characters of ancient ostracods and could be an important character for evaluating the history of sexual dimorphism in ostracods since the Palaeozoic.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 153 , 239–251.     

The colonization of Scottish islands by the common shrew, Sorex araneus (Eulipotyphla: Soricidae)

In this paper we apply molecular methods to study the colonization of islands off the west coast of Scotland by the common shrew ( Sorex araneus L.), and current gene flow. We collected 497 individuals from 13 islands of the Inner Hebrides and Clyde Island groups and six mainland regions. Individuals were genotyped at eight microsatellite loci, and the mitochondrial cytochrome b sequence (1140 base pairs) was obtained for five individuals from each island/mainland region. Based on these molecular data, island colonization apparently proceeded directly from the mainland, except for Islay, for which Jura was the most likely source population. Raasay may also have been colonized by island hopping. Most island populations are genetically very distinct from the mainland populations, suggesting long periods of isolation. Two exceptions to this are the islands of Skye and Seil, which are geographically and genetically close to the mainland, suggesting in each case that there has been long-term gene flow between these islands and the mainland. We consider possible methods of island colonization, including human-mediated movement, swimming, and land and ice bridges.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 797–808.