Epigenetic polymorphism in populations of Apodemus sylvaticus and Apodemus flavicollis (Rodentia, Muridae)

The main object of this study was to use discontinuous variation as a means of measuring divergence between populations of A. sylvaticus (L.) and A. flavicollis (Melchior) from Britain and Eurasia. Samples from 17 populations (671 skulls) were each classified for 20 discontinuous morphological variants. The investigation has shown that certain epigenetic variants have a characteristic incidence in populations of the two species and that the relative incidence of the variants could be used as an aid to specific separation. The calculation of degree of divergence between the populations has made it possible to suggest the order in which populations are related to each other.     

Genetic variation and evolution in the genus Apodemus (Muridae: Rodentia)

Genetic variation was studied using protein electrophoresis of 28–38 gene loci in 1347 specimens of Apodemus agrar-ius, A. peninsulae, A. flavicollis, A. sylvaticus, A. alpicola, A. uralensis, A . cf. hyrcanicus, A. hermonensis, A. m. mystacinus and A. m. epimelas , representing 121 populations from Europe, the Middle East, and North Africa. Mean values of heterozygosity per locus for each species ranged from 0.02 to 0.04. Mean values of Nel's genetic distance ( D ) between the taxa ranged from 0.06 (between A. flavicollis and A. alpicola) to 1.34 (between A. uralensis and A. agrarius ). The highest values of D were found between A. agrarius and other Apodemus species (0.62-1.34). These values correspond to those generally observed between genera in small mammals. Our data show that A. agrarius and A. peninsulae are sister species, well-differentiated from other taxa. High genetic distance between A. m. mystacinus and A. m. epimelas leads us to consider them distinct species and sister taxa to other Western Palaearctic species of the subgenus Sylvaemus . The data also suggest a recent separation of members of the latter group from a common ancestor, and subsequent rapid radiation, making it difficult to infer phylogenetic relationships. Some taxonomic implications of the results are discussed further.© 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 395–419.     

Impact of geographical isolation on genetic differentiation in insular and mainland populations of Weigela coraeensis (Caprifoliaceae) on Honshu and the Izu Islands

Aim To provide insights into genetic differentiation between insular endemic Weigela coraeensis var. fragrans and its progenitor variety W. coraeensis var. coraeensis, the population genetic structure of both varieties was examined, and factors promoting genetic differentiation between the two taxa were explored. Location The natural range of W. coraeensis (sensu lato) throughout mainland Japan (Honshu) and the Izu Islands. Methods The analysis included 349 and 504 individuals across the mainland (Honshu) and the Izu Islands, respectively, using 10 allozyme and 10 microsatellite loci. The population genetic structure of W. coraeensis was assessed by analysing genetic diversity indices for each population, genetic differentiation among populations, model‐based Bayesian clustering or distance‐based clustering, and bottleneck tests. Results The level of genetic diversity in each of the populations on the Izu Islands was negatively correlated with geographical distance between each island and the mainland. The populations on the mainland and on the Izu Islands were genetically differentiated to a certain extent; however, the microsatellite analyses suggested that gene flow also occurred between the mainland and the islands, and among individual islands. These microsatellite analyses also suggested recent bottlenecks in several populations in both areas. Main conclusions The decrease in genetic diversity throughout the Izu Islands, which correlated with distance to the mainland, Honshu, may be the result of a repeated founder effect occurring at a series of inter‐island colonizations from north to south. The stepping stone‐like configuration of the islands may have played a role in the dispersal of the species. Geographical isolation by sea would effectively result in genetic differentiation of W. coraeensis between mainland Honshu and the Izu Islands, although some gene flow may still occur between Honshu and the northern Izu Islands. The differentiation process of the endemic plants on the Izu Islands is anagenetic but not completed, and the study of these plants will provide insightful knowledge concerning the evolution of insular endemics.     

Chromosomal variability of the field mouse Apodemus agarius (Rodentia, Muridae)

Chromosome sets of 114 Apodemus agrarius mice from 29 localities in Moldova, Ukraine, Siberia, and Far East were studied by means of G-, C-, and NOR-banding. In all populations studied, the Y chromosome was shown to be a medium-size acrocentric chromosome consisting of heterochromatin. Chromosome polymorphism observed in populations from Primorskii krai concerned (1) the morphology of the first two autosome pairs (variants A/A, A/ST, and ST/ST), (2) the number of metacentric chromosomes (from 6 to 8), and (3) heterochromatin localization in the pericentromeric regions of two metacentric chromosome pairs. A karyotype with an additional heterochromatic microchromosome found in all the metaphases studied was described in one mouse from a locality of western Primorye that has not been studied previously. In the karyotype of 15 mice from four populations of Primorye, the pool of nucleolus organizer regions is distributed over three autosome pairs rather than over four, as is the case A. agrarius from Europe. Based on the analysis of literature sources and our own data, the problem of chromosome polymorphism in the field mouse is discussed.     

Cumulative effects of founding events during colonisation on genetic diversity and differentiation in an island and stepping-stone model

This paper investigates the cumulative effect of founding events on the genetic differentiation and the within-population heterozygosity in a metapopulation increasing its size by colonisation. Two contrasting models are considered: first, an island model, where migrants and colonists are taken at random from the entire metapopulation, and second, a linear stepping-stone model, where migrants and colonists are sampled from a limited neighbourhood. The genetic consequences of a range expansion depend on the relative magnitudes of the number of colonists and migrants, in a way similar to extinction and colonisation processes (Wade and McCauley, 1988). The cumulative effect of founding events, resulting most often in a transient increase in genetic differentiation and a gradual loss of within-population heterozygosity, also depends on the age-structure that is established during colonisation. It is the highest when colonists are sampled from recently founded populations and migrants are exchanged among populations of similar ages. The genetic consequences of a range expansion are therefore far more pronounced and lasting in the linear stepping-stone model than in the island model. These two models, however, represent the two extremes between which real populations will fall.     

Origin and evolution of the Robertsonian populations of the house mouse (Rodentia, Muridae) in Tunisia based on allozyme studies

The central coastal region of Tunisia harbours two chromosomal races of the house mouse (2 n  = 22, 22Rb; 2 n  = 40, 40Std), which are genetically differentiated and show a high level of reproductive isolation. This study presents an allozyme analysis of house mice from the inland city of Kairouan 75 km from the coastal region. Results showed that the 22Rb and 40Std mice from Kairouan shared the same high degree of variability, and were not genetically differentiated. This contrasts with the genetic data registered for the two chromosomal races from the coastal towns, in which a particularly low genic diversity was observed in the 22Rb populations. As the two races in Kairouan show the same low hybridization rates as the populations in the coastal region, these results argue in favour of a local differentiation of the Rb race in Tunisia, most likely originating in Kairouan. The data further suggest that the decrease in variability observed in the coastal 22Rb populations is related to a founder effect or a bottleneck following the introduction of mice from Kairouan into one of these localities, prior to their dispersion throughout the coastal region.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 515–521.     

Role of interspecific competition in the coexistence ofApodemus argenteus andA. speciosus (Rodentia: Muridae)

Reciprocal-removal experiments with two replicates were conducted to test for the role of interspecific competition in the coexistence ofApodemus argenteus andA. speciosus. Population density, rate of appearance of new (unmarked) individuals, reproduction, survival rate and habitat use were monitored during pre- and removal periods. In both removal experiments, the removal ofA. argenteus had little effect onA. speciosus, while that ofA. speciosus affected several population characteristics ofA. argenteus. Namely, the removal ofA. speciosus shifted the distribution ofA. argenteus to the habitat with a denser shrub cover in one experiment. Also, the removal increased the population densities and appearance rates of new individuals ofA. argenteus in another experiment. Interspecific interactions between the two species appeared to be a one-way action fromA. speciosus toA. argenteus. In removal periods in both experiments, the rates of appearance of new individuals in each species were the highest on the grid where that species was removed. These results suggest that, though interspecific competition occurred between the two species, intraspecific competition had greater effects than interspecific competition on the abundance and habitat use ofA. argenteus andA. speciosus. This implies that the fundamental niches ofA. argenteus andA. speciosus differ potentially, which may play an important role in the coexistence of the two species.     

Social differences in the subcaudal scent gland secretion of Apodemus sylvaticus (Rodentia: Muridae)

This paper presents a preliminary chemical analysis of the secretion of the subcaudal scent gland of the Woodmouse Apodemus sylvaticus . A statistical analysis of samples from various sites in the United Kingdom and Eire shows that there are individual differences in the secretion, differences between the sexes, differences between juveniles and adults and differences between populations.     

Population structure of the large Japanese field mouse, Apodemus speciosus (Rodentia: Muridae), in suburban landscape, based on mitochondrial D-loop sequences

Genetic structure of the large Japanese field mouse populations in suburban landscape of West Tokyo, Japan was determined using mitochondrial DNA control region sequence. Samples were collected from six habitats linked by forests and green tract along the Tama River, and from two forests segregated by urban areas from those continuous habitats. Thirty-five haplotypes were detected in 221 animals. Four to eight haplotypes were found within each local population belonging to the continuous landscape. Some haplotypes were shared by two or three adjacent local populations. On the other hand, two isolated habitats were occupied by one or two indigenous haplotypes. Significant genetic differentiation between all pairs of local populations, except for one pair in the continuous habitats, was found by analysis of molecular variance (amova). The geographical distance between habitats did not explain the large variance of pairwise F(ST)-values among local populations. F(ST)-values between local populations segregated by urban areas were higher than those between local populations in the continuous habitat, regardless of geographical distance. The results of this study demonstrated quantitatively that urban areas inhibit the migration of Apodemus speciosus, whereas a linear green tract along a river functions as a corridor. Moreover, it preserves the metapopulation structure of A. speciosus as well as the corridors in suburban landscape.     

The phylogeny of the Praomys complex (Rodentia: Muridae) and its phylogeographic implications

Among the African Murinae (Rodentia, Muridae), the Praomys complex, whose systematics has been studied by different approaches, has raised numerous taxonomic problems. Different taxa, namely Praomys, Mastomys , Myomys and Hylomyscus have been considered either as separate genera or subgenera of Praomys . In order to understand the relationships within the Praomys complex and to test the monophyly of the genus Praomys , a cladistic analysis was conducted, based on morpho-anatomical factors involving different species of Praomys , Mastomys , Myomys and Hylomyscus. The results indicate that the Praomys complex is monophyletic, as are the genera Hylomyscus , Mastomys and Myomys , whereas the genus Praomys appears paraphyletic. Indeed, a group of species including Praomys jacksoni was found to be more closely related to the genera Mastomys and Myomys than to a Praomys tullbergi -group. The biotopes and the distribution areas of the species were mapped on the phylogeny. It appears that the different clades each present a relative ecological cohesion and are arranged according to a gradient from closed to open habitats. From there, an evolutionary scenario is proposed for the emergence of the different clades and species of the genus Praomys sensu stricto.     

Pattern of Breeding-System Change in the Izu Islands in Campanula punctata: Bumblebee-Absence Hypothesis

Abstract Bagging experiments of Campanula punctata were carried out in natural populations on the six of the Izu Islands and mainland Honshu to investigate distribution of self-incompatibility. Additional observations on pollinators of Campanula supplementary to the previous study (Inoue and Amano, 1986) were made. Most of the plants on Honshu and Oshima Island showed self-incompatibility: while almost all plants in the Izu Islands except Oshima were self-compatible. Bumblebees did not exist in the Izu Islands except Oshima and the dominant pollinators of Campanula in these islands were halictid bees. These results support a hypothesis that a decrease in pollination efficiency in the absence of bumblebees results in the evolution of breeding systems in Campanula.     

History vs. habitat type: explaining the genetic structure of European nine‐spined stickleback (Pungitius pungitius) populations

The genetic structure of contemporary populations can be shaped by both their history and current ecological conditions. We assessed the relative importance of postglacial colonization history and habitat type in the patterns and degree of genetic diversity and differentiation in northern European nine‐spined sticklebacks (Pungitius pungitius), using mitochondrial DNA (mtDNA) sequences and 12 nuclear microsatellite and insertion/deletion loci. The mtDNA analyses identified – and microsatellite analyses supported – the existence of two historically distinct lineages (eastern and western). The analyses of nuclear loci among 51 European sites revealed clear historically influenced and to minor degree habitat dependent, patterns of genetic diversity and differentiation. While the effect of habitat type on the levels of genetic variation (coastal > freshwater) and differentiation (freshwater > coastal) was clear, the levels of genetic variability and differentiation in the freshwater sites were independent of habitat type (viz. river, lake and pond). However, levels of genetic variability, together with estimates of historical effective population sizes, decreased dramatically and linearly with increasing latitude. These geographical patterns of genetic variability and differentiation suggest that the contemporary genetic structure of freshwater nine‐spined sticklebacks has been strongly impacted by the founder events associated with postglacial colonization and less by current ecological conditions (cf. habitat type). In general, the results highlight the strong and persistent effects of postglacial colonization history on genetic structuring of northern European fauna and provide an unparalleled example of latitudinal trends in levels of genetic diversity.     

Island hopping across the central Pacific: mitochondrial DNA detects sequential colonization of the Austral Islands by crab spiders (Araneae: Thomisidae)

Aim Phylogenetic studies concerning island biogeography have been concentrated in a fraction of the numerous hot‐spot archipelagos contained within the Pacific Ocean. In this study we investigate relationships among island populations of the thomisid spider Misumenops rapaensis Berland, 1934 across the Austral Islands, a remote and rarely examined southern Pacific hot‐spot archipelago. We also assess the phylogenetic position of M. rapaensis in relation to thomisids distributed across multiple Polynesian archipelagos in order to evaluate the proposed hypothesis that thomisid spiders colonized Polynesia from multiple and opposing directions. The data allow an examination of genetic divergence and species accumulation in closely related lineages distributed across four Polynesian archipelagos. Location The study focused on four Polynesian hot‐spot archipelagos: the Austral, Hawaiian, Marquesan and Society islands. Methods Mitochondrial DNA sequences comprising c. 1400 bp (portions of cytochrome oxidase subunit I, ribosomal 16S and NADH dehydrogenase subunit I) were obtained from thomisid spiders (64 specimens, representing 33 species) collected in the Australs, the Hawaiian Islands, the Society Islands, the Marquesas, Tonga, Fiji, New Zealand, New Caledonia and North and South America. Phylogenetic analyses using parsimony, maximum‐likelihood and Bayesian approaches were employed to resolve relationships of M. rapaensis to other Polynesian Misumenops and across the Austral Islands. Results Rather than grouping with other Misumenops spp. from the archipelagos of the Society Islands, Marquesas and Hawaiian Islands, M. rapaensis appears more closely related to Diaea spp. from Tonga, Fiji, New Zealand and New Caledonia. Phylogenetic analyses strongly support M. rapaensis as monophyletic across the Austral Islands. Misumenops rapaensis sampled from the two older islands (Rurutu and Tubuai) form reciprocally monophyletic groups, while individuals from the younger islands (Raivavae and Rapa) are paraphyletic. Across the Austral Islands, M. rapaensis exhibits a surprising level of genetic divergence (maximally 11.3%), an amount nearly equivalent to that found across the 16 examined Hawaiian species (14.0%). Main conclusions Although described as a single morphologically recognized species, our results suggest that M. rapaensis comprises multiple genetically distinct lineages restricted to different Austral Islands. Phylogenetic relationships among the island populations are consistent with sequential colonization of this lineage down the Austral archipelago toward younger islands. Analyses support the hypothesis that thomisid spiders colonized the central Pacific multiple times and suggest that M. rapaensis arrived in the Austral Islands from a westward direction, while Misumenops found in neighbouring archipelagos appear to be more closely related to New World congeners to the east. Finally, our data detect asymmetrical rates of morphological evolution and species diversification following colonization of four different Polynesian archipelagos.     

Population dynamics of the wood mouse Apodemus sylvaticus (Rodentia: Muridae) in a Sitka spruce successional mosaic

A population of wood mice, Apodemus sylvaticus (L.), was studied by capture-mark-recapture, in five 0.81 ha grids within a habitat mosaic produced by the felling of a plantation of mature Sitka spruce ( Picea sitchensis Carriere). From May 1990 to June 1992, we estimated population sizes, survival (total and by weight classes), recruitment, and home-range sizes, usually at monthly intervals, for the study area taken as a whole. Inter-grid movements were frequent for both sexes, supporting the assumption that the population could be analysed in this way. Wood mice were more abundant in 1991 than in 1990, associated with increased seed supply. In both years, numbers were low in spring and increased gradually towards an autumn peak, which was of similar size in both years. Male home-range areas varied seasonally, and were negatively correlated with population sizes, while female home ranges showed little seasonal variation. Survival and recruitment both declined sharply in late autumn, apparently in a densitydependent way. Except for this latter result, the population dynamics in a heterogeneous coniferous woodland were consistent with those described previously for this species in more homogeneous deciduous woodlands, suggesting that the same regulatory processes are operating in both habitats.     

Phylogeographic structure and regional history of Lemniscomys striatus (Rodentia: Muridae) in tropical Africa

Aim This study aims to elucidate the phylogeography of the murid rodent Lemniscomys striatus and to evaluate the relative roles of ecological change, habitat patchiness, rivers and geological barriers in structuring patterns of diversity. Location Sub‐Saharan Africa. Methods The extent of phylogeographic patterns and molecular genetic diversity (cytochrome b gene) were addressed in a survey of 128 individuals of L. striatus from 42 localities. Using maximum parsimony, maximum likelihood, Bayesian, network and genetic structure analyses, we inferred intraspecific relationships and tested hypotheses for historical patterns of gene flow within L. striatus. Results Our results identified four major geographical clades within L. striatus: a West African clade, a Benin‐Nigeria clade, a Central African clade, and an East African clade. Several subclades were identified within these four major clades. Restricted gene flow with isolation by distance was recorded, which is congruent with the low dispersal ability of such a small murid rodent. No clear signal of population expansion was detected within clades or subclades. Main conclusions The western rift system and the Volta and Niger rivers may have acted as long‐term extrinsic barriers to gene flow, resulting in the emergence of the four main clades of L. striatus. The observed pattern of mitochondrial variation observed within each clade probably results from late Pleistocene climatic and vegetation changes: during adverse conditions (forest expansion), L. striatus may have survived only in refugia, and then experienced range expansion under favourable conditions (savanna expansion).     

Evolution of Campanula punctata Lam. in the Izu Islands: Changes of Pollinators and Evolution of Breeding Systems

Abstract The pollination and breeding systems of Campanula punctata both in the Izu Islands and the mainland (Honshu) adjacent to the islands were investigated. The predominant pollinators of campanulas are: 1) Bombus diversus in Honshu, 2) Bombus ardens and halictid bees on Oshima Island, and 3) halictid bees on the islands of Niijima, Kodzushma, and Hachijo. The absence of bumble bees except B. ardens on Oshima Island, and the abundance of halictid bees represent a kind of island effect. Flowers of the campanula in the Islands were found to be smaller than those of the mainland. This might have resulted from adaptation to the small pollinators. Pollination experiments with plants grown from seeds revealed the following differences in breeding system: 1) most of the campanulas from Honshu were highly self-incompatible (SI) and a few partially self-compatible (SC), 2) one half of the plants from Oshima Island were more or less SC and the remaining highly SI, 3) those from Hachijo Island were SC and potentially autogamous. The breakdown of dichogamy co-operated with self-compatibility in the development of self-fertilizing ability in Hachijo plants. Several possible causes for the change in the breeding system including the change in pollinators in the Islands are discussed.     

Effects of acorn abundance on density dependence in a Japanese wood mouse (Apodemus speciosus) population

We analysed the effects of Quercus crispula acorn abundance on the density dependence of the large Japanese wood mouse Apodemus speciosus using time series data (1992–2007). The data were obtained in a forest in northern Hokkaido, Japan, by live-trapping rodents and directly counting acorns on the ground. Acorn abundance in one year clearly influenced the abundance of wood mice in the following year in all models examined based on the Gompertz and Ricker model; in addition, the abundance of wood mice had effects on the population. Acorn abundance influenced the strength of density dependence (intraspecific competition) of the wood mouse population. When the abundance of acorns was high, density dependence was relaxed, and as a result the equilibrium density at which the population growth rate decreased to zero became higher. Those effects of acorn abundance were regarded as a nonlinear perturbation effect (sensu Royama 1992). The nonlinearity of density dependence was also detected; higher densities had stronger effects on population growth rates. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.