Phylogeography of the Russian flying squirrel (Pteromys volans): implication of refugia theory in arboreal small mammal of Eurasia

A phylogeographical study of the Russian (Siberian) flying squirrel (Pteromys volans) was carried out using the complete mitochondrial (mt) cytochrome b gene sequences with special reference to the refugia theory for the arboreal traits of this species. We examined 31 specimens from 24 localities, resulting in 28 haplotypes. One breeding specimen with a unique haplotype from Hokkaido, Japan was included in the phylogenetic analysis. There were three mtDNA lineages: Hokkaido, Far Eastern, and northern Eurasia. Divergence data among lineages demonstrated that the Hokkaido group separated from the other groups during the Holsteinian interglacial. The phylogeographical pattern of P. volans is different from that previously reported for terrestrial rodents associated with treeless habitats. Unlike grasslands, forests decreased during glaciation and moved southward because of the cold and arid environmental conditions. The glacial refugia of P. volans would have been associated with forest dynamics in the Pleistocene.     

Contrasting seasonal dynamics in fleas of the Siberian flying squirrel (Pteromys volans) in Finland

Abstract.  1. The seasonal and spatial variation of the adult flea fauna (Siphonaptera) was examined in connection with live-trapping studies of the Siberian flying squirrel ( Pteromys volans ) in three study areas in southern Finland between 1997 and 2005.
2. The numerically dominant flea species of the Siberian flying squirrel were Tarsopsylla octodecimdentata octodecimdentata and Ceratophyllus (Monopsyllus) indages indages (Ceratophyllidae); the latter being a host specialist of the Siberian flying squirrel. Tarsopsylla octodecimdentata , which also commonly occurs on the red squirrel ( Sciurus vulgaris ), infected a large proportion of the host population throughout the winter, whereas C. indages occurred predominantly during the summer and autumn, being practically the only flea species in nests during the flying squirrels' breeding season and on juveniles still inhabiting their natal nest.
3. The use of nest boxes by flying squirrels did not have any positive effects on flea populations, but it may have had a negative impact on T.   octodecimdentata .
4. The potentially vulnerable C.   i.   indages is a predictable, widespread component in the flea fauna of the Siberian flying squirrel at various spatial scales, and it is likely to occur in most of the permanent flying squirrel populations in Finland.     

黑龙江省张广才岭北部小飞鼠的遗传多样性与种群历史动态

小飞鼠 (Pteromys volans) 为树栖夜行滑行类啮齿动物,在森林种子传播和维持生态系统平衡等方面发挥着重要的生态学作用。本研究利用mtDNA Cytb、控制区和nDNA微卫星3种分子标记,对黑龙江省张广才岭北部的小飞鼠种群进行遗传多样性与历史动态分析。检测出Cytb全序列 (1 140 bp) 的平均单倍型多样性为0.909,平均核苷酸多样性为0.616%;控制区全序列 (1 066 bp) 的平均单倍型多样性为0.945,平均核苷酸多样性为1.698%;微卫星检测出种群平均等位基因数13.167个,观测杂合度0.727,期望杂合度0.864,近交系数0.159。结果表明,小飞鼠种群遗传多样性丰富,但存在一定程度的杂合度不足和近亲繁殖;未检测到种群近期遗传瓶颈效应,种群内无遗传分化。高比例的稀有单倍型 (≥ 60%) 、低频率等位基因与近亲繁殖,提示未来种群面临遗传多样性下降的风险,建议加大对该物种的关注和保护力度。基于Cyt b基因的系统进化关系结果表明,小飞鼠存在3个明显的遗传谱系：远东、欧亚大陆北部和日本北海道,本研究中张广才岭和大兴安岭的样本单倍型归属为远东谱系。     

Female-biased natal dispersal in the Siberian flying squirrel

Natal dispersal is usually sex biased in birds and mammals.Female-biased natal dispersal is the prevailing pattern in birdsbut is rare among mammals. Hypotheses explaining sex bias indispersal include the mate-defense mating hypothesis, whichpredicts male-biased dispersal, the resource-defense hypothesispredicting female-biased dispersal, and the competition hypothesis,which predicts that if dispersal is caused by competition forresources between sexes, then the subdominant sex will disperse.We studied natal dispersal of Siberian flying squirrels Pteromysvolans using radio telemetry in Southern Finland in 1996–2004.Of 86 juveniles that survived over the dispersal period, almostall young females dispersed from the natal site, whereas almost40% of males were philopatric. Dispersal was farther for femalesthan males. Females began dispersal on average 2 weeks earlierthan males and were lighter in mass at the onset of dispersalthan later dispersing males. No mate- or resource-defense matingsystem could be found among males, but females seemed to defendnest and apparently food resources, in contrast to the expectationof dispersal bias in resource-defense systems. Competition forresources between sexes does not explain female bias either:in the flying squirrel, the female seems to be the dominantsex. We propose that young females are subordinate to theirmothers and have to disperse to find a vacant, suitable sitefor reproduction.     

High inbreeding and low connectivity among Ambystoma texanum populations in fragmented Ohio forests

Habitat loss and fragmentation negatively impact the size and diversity of many natural populations. Woodland amphibians require connected aquatic and terrestrial habitats to complete their life cycle, and often rely on metapopulation structure for long‐term persistence. Wetland loss and deforestation fragment amphibian populations, which may result in population isolation and its negative effects. The aim of this research was to analyze the population genetic structure of small‐mouthed salamanders (Ambystoma texanum) in western Ohio, where agriculture is now the dominant land use. Salamander tail tissue was collected from eight breeding pools. Three pools occur in the same forest; the other five are in forest patches at distances ranging from 250 m to 20 km from one another. Eight microsatellite loci were amplified by PCR and genotyped for allele size. Observed heterozygosities were lower than expected in all sampled populations; the two most isolated sites (Ha1, Ha2) had the highest inbreeding coefficients. Ha2 also had the lowest mean number of alleles and was found to be genetically differentiated from populations to which our data analysis indicates it was historically connected by gene flow. The most distant site (Ha1) had the highest number of private alleles and showed genetic differentiation from other populations both historically and currently. Geographic distance between pools was strongly correlated with the number of private alleles in a population. The results suggest that population isolation results in decreased genetic diversity and that a breakdown of metapopulation structure due to landscape change may contribute to differentiation between once‐connected populations.     

An RAPD (random amplified polymorphic DNA) analysis of genetic population structure of Balea biplicata (Gastropoda: Clausiliidae) in fragmented floodplain forests of the Elster/Saale riparian system

Eight German populations of the land snail Balea biplicata(Mollusca: Clausiliidae) were studied using the randomly amplified polymorphic DNA-polymerase chain reaction and morphometrics (principal component and discriminant analysis) to examine population structure and gene flow patterns in a fragmented landscape mosaic along the Elster/Saale riparian system, Germany. A variety of population genetic analyses targeting either more on the geographic scale of gene flow (genetic distances, F statistics, Mantel test) or on local genotypic structure (heterozygosity, linkage disequilibrium, bottleneck probability) showed that (1) the population system in total is governed by high gene flow independent of geographic distance, (2) genetic structure on the narrower sampling scale is mainly determined by stochastic processes due to genetic drift in small isolated and frequently recolonized populations, and (3) the morphometrical variation of the populations was related neither to habitat nor to genetic heterogeneity. The potentials for active and passive dispersal capacity of the snails and possible environmental impacts on their population structure are discussed.     

Inbreeding avoidance in cunningham's skinks (Egernia cunninghami) in natural and fragmented habitat

Habitat fragmentation/alteration has been proposed as a distinct process threatening the viability of populations of many organisms. One expression of its impact may be the disruption of core population processes such as inbreeding avoidance. Using the experimental design outlined in our companion paper, we report on the impact of habitat alteration (deforestation) on inbreeding in the rock-dwelling Australian lizard Egernia cunninghami. Ten microsatellite loci were used to calculate relatedness coefficients of potential and actual breeding pairs, and to examine mate-choice and heterozygosity. Despite significantly less dispersal and higher within-group relatedness between potential mates in deforested than in natural habitats, this did not result in significantly more inbred matings. Average relatedness amongst breeding pairs was low, with no significant difference between natural and fragmented populations in relatedness between breeding pairs, or individual heterozygosity. Active avoidance of close kin as mates was indicated by the substantially and significantly lower relatedness in actual breeding pairs than potential ones. These facts, and heterozygote excesses in all groups of immature lizards from both habitats, show that E. cunninghami maintained outbreeding in the face of increased accumulation of relatives.     

Population genetic structure and dispersal across a fragmented landscape in cerulean warblers (<Emphasis Type="Italic">Dendroica cerulea</Emphasis>)

Cerulean warblers (Dendroica cerulea) have experienced significant declines across their breeding range and presently exist in disjunct populations, largely because of extensive loss and fragmentation of their breeding and wintering habitat. Despite this overall decline, a recent north-eastern expansion of the breeding range has been proposed, and some researchers have suggested that the eastern Ontario population may be acting as a source population maintaining sink populations elsewhere. However, little is known about either the geographic distribution of genetic variation or dispersal in these birds. We assayed variation in five microsatellite loci and a 366 base-pair fragment of the mitochondrial control region among 154 cerulean warblers from five populations throughout the breeding range. No evidence of population genetic structure was found. Assignment tests suggested that six individuals were either inter-population migrants or descendants of recent migrants. The lack of population genetic structure is probably due to a combination of historical association and contemporary dispersal. Population decline does not appear to have reduced genetic variation yet. Overall results suggest that cerulean warblers from Ontario, Illinois, Arkansas and Tennessee should be considered a single genetic management unit for conservation.     

Spatial and temporal variation in flight morphology in the butterfly Melitaea cinxia (Lepidoptera: Nymphalidae)

Some studies on insects have found a relationship between habitat structure and investment in dispersal-related traits. In this study we compared the morphology of the butterfly Melitaea cinxia from five sites on the large Baltic island Öland that differed markedly in degree of fragmentation and size. Both wild-caught adults and individuals reared in a split-plot design were compared. We found significant site differences in size-adjusted thorax mass and total body mass. Male thorax mass was on average larger among the sites with the highest degree of habitat fragmentation. However, due to significant sex–site interactions, males and females may have adapted differently to the habitat fragmentation. Using museum specimens, we also analysed changes in morphology, finding an increase in size (measured as head and thorax width) over time. Thorax width appears to have increased among females and decreased among males. Possible explanations include increasing fragmentation of the landscape and changes in population density.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 445–453.     

Population structure of spotted salamanders (Ambystoma maculatum) in a fragmented landscape

Understanding the impacts of landscape-level processes on the population biology of amphibians is critical, especially for species inhabiting anthropogenically modified landscapes. Many pond-breeding amphibians are presumed to exist as metapopulations, but few studies demonstrate the extent and consequences of this metapopulation structure. Gene flow measures may facilitate the construction of more realistic models of population structure than direct measures of migration. This is especially true for species that are cryptic, such as many amphibians. We used eight polymorphic microsatellite loci to determine the genetic population structure of spotted salamanders ( Ambystoma maculatum ) breeding at 17 ponds in northeastern Ohio, a landscape fragmented by roads, agriculture, urban areas and the Cuyahoga River. Using a variety of analyses (Bayesian clustering, F -statistics, AMOVA) we generated a model of salamander population genetic structure. Our data revealed patterns of genetic connectivity that were not predicted by geographical distances between ponds (no isolation by distance). We also tested for a relationship between population structure and several indices of landscape resistance, but found no effect of potential barriers to dispersal on genetic connectivity. Strong overall connectivity among ponds, despite the hostile habitat matrix, may be facilitated by a network of riparian corridors associated with the Cuyahoga River; however, high gene flow in this system may indicate a general ability to disperse and colonize beyond particular corridors.     

Movement patterns of Rhyssomatus lineaticollis Say (Coleoptera: Curculionidae) within and among Asclepias syriaca (Asclepiadaceae) patches in a fragmented landscape

Abstract.  1. Dispersal capabilities of organisms are critical in determining the landscape population structure of species as well as their likelihood of survival in fragmented landscapes. Using mark–recapture techniques on the monophagous weevil Rhyssomatus lineaticollis Say (Curculionidae), within- and between-patch dispersal capabilities, landscape level population structure, and the role of beetle density and host patch characteristics in setting distances, amounts, and timing of dispersal were studied.
2. The data indicate that R. lineaticollis is sedentary, with 50% of recaptured beetles moving < 1 m and the maximum distance moved < 1 km. Within- and between-patch movement of beetles was unrelated to host plant patch characteristics and beetle densities.
3. Despite limited dispersal, R. lineaticollis probably functions as a patchy population in east-central Iowa, U.S.A. because dispersals between patches are common and because all host patches surveyed contained this herbivore, indicating a lack of suitable vacant patches, a prerequisite for metapopulation structure.
4. Between-patch distances are well within the dispersal capabilities of R. lineaticollis , although this may be the result of an increase in the density of patches of its host, Asclepias syriaca , in the landscape over the last 150 years as a result of human disturbance and this species' weedy habit.
5. Metapopulation structure in monophagous prairie herbivores may be most likely in species whose non-weedy host plants form highly predictable resources in space and time, but which are now widely scattered in habitat fragments.     

Cold-induced photoinhibition and growth of seedling snow gum (Eucalyptus pauciflora) under differing temperature and radiation regimes in fragmented forests

Fluorescence characteristics and growth of seedling snow gum ( Eucalyptus pauciflora Sieb. ex Spreng.) during autumn and winter were related to variation in radiation and temperature regime in a fragmented forest. Seedlings were planted in four treatments along transects perpendicular to tree island edges to characterize plant responses to microclimates ranging from those of cleared areas to those beneath forest canopies. Three-dimensional mapping of seedling leaf display, in combination with information retrieved from hemispherical photographs about shading from overstory canopies, were used to calculate the intercepted amounts of direct radiation energy for unit area of leaves on clear days (IDRE) . IDRE was highest on the outside, most variable at the edges and lowest well inside the tree islands. Minimum temperature decreased with increasing view of the sky. Photoinhibition, measured as decrease in F _v/ F _m, was correlated with spatial and seasonal differences in weekly minimum temperature and IDRE . Seedlings in the open and under the most canopy cover, with low variability in IDRE in a scale of weeks, exhibited less variability in photoinhibition than those growing along forest edges. Seedlings in the open tended to be most photoinhibited and grew the most. The combination of increased IDRE with reduced minimum temperatures resulted in persistent and strong photoinhibition as the season progressed. Results are discussed in relation to the potential for seedling establishment following forest fragmentation.     

Preliminary Information on the Home Range and Movement Patterns of Sturnira lilium (Phyllostomidae) in a Naturally Fragmented Landscape in Bolivia

Here, we used radiotelemetry to determine home range and movement patterns of the frugivorous bat Sturnira lilium in a naturally fragmented landscape in Beni, Bolivia. Four females were monitored for up to 7 d. Based on 575 radiotelemetry positions, kernel home ranges were between 36.5 and 90.7 ha. During each night, bats visited most of their home range, which included forest islands and continuous forest; however activity was concentrated in core-use areas representing 6–12 percent of the home ranges. There was moderate overlap in home range and core-use areas among the four females. The mean long axis across the kernel home ranges was 1324 m. The results reported here show that S. lilium is able to move freely among forest elements in the landscape while crossing a savanna matrix. Hence, S. lilium may contribute to maintain connectivity and ecological processes in a fragmented landscape.     

Population structure and genetic diversity of black redhorse (Moxostoma duquesnei) in a highly fragmented watershed

Dams have the potential to affect population size and connectivity, reduce genetic diversity, and increase genetic differences among isolated riverine fish populations. Previous research has reported adverse effects on the distribution and demographics of black redhorse (Moxostoma duquesnei), a threatened fish species in Canada. However, effects on genetic diversity and population structure are unknown. We used microsatellite DNA markers to assess the number of genetic populations in the Grand River (Ontario) and to test whether dams have resulted in a loss of genetic diversity and increased genetic differentiation among populations. Three hundred and seventy-seven individuals from eight Grand River sites were genotyped at eight microsatellite loci. Measures of genetic diversity were moderately high and not significantly different among populations; strong evidence of recent population bottlenecks was not detected. Pairwise F_ST and exact tests identified weak (global F_ST = 0.011) but statistically significant population structure, although little population structuring was detected using either genetic distances or an individual-based clustering method. Neither geographic distance nor the number of intervening dams were correlated with pairwise differences among populations. Tests for regional equilibrium indicate that Grand River populations were either in equilibrium between gene flow and genetic drift or that gene flow is more influential than drift. While studies on other species have identified strong dam-related effects on genetic diversity and population structure, this study suggests that barrier permeability, river fragment length and the ecological characteristics of affected species can counterbalance dam-related effects.     

Landscape genetic structure of coastal tailed frogs (Ascaphus truei) in protected vs. managed forests

Habitat loss and fragmentation are the leading causes of species’ declines and extinctions. A key component of studying population response to habitat alteration is to understand how fragmentation affects population connectivity in disturbed landscapes. We used landscape genetic analyses to determine how habitat fragmentation due to timber harvest affects genetic population connectivity of the coastal tailed frog (Ascaphus truei), a forest-dwelling, stream-breeding amphibian. We compared rates of gene flow across old-growth (Olympic National Park) and logged landscapes (Olympic National Forest) and used spatial autoregression to estimate the effect of landscape variables on genetic structure. We detected higher overall genetic connectivity across the managed forest, although this was likely a historical signature of continuous forest before timber harvest began. Gene flow also occurred terrestrially, as connectivity was high across unconnected river basins. Autoregressive models demonstrated that closed forest and low solar radiation were correlated with increased gene flow. In addition, there was evidence for a temporal lag in the correlation of decreased gene flow with harvest, suggesting that the full genetic impact may not appear for several generations. Furthermore, we detected genetic evidence of population bottlenecks across the Olympic National Forest, including at sites that were within old-growth forest but surrounded by harvested patches. Collectively, this research suggests that absence of forest (whether due to natural or anthropogenic changes) is a key restrictor of genetic connectivity and that intact forested patches in the surrounding environment are necessary for continued gene flow and population connectivity.     

Isolation and characterization of microsatellite loci in the dark bush cricket,Pholidoptera griseoaptera (Tettigoniidae)

Twelve novel polymorphic microsatellite loci are presented for the dark bush cricket, Pholidoptera griseoaptera. All loci are polymorphic, with up to 37 alleles per locus. These microsatellites will be useful tools for studying the influence of landscape structure and land use intensity in agricultural landscapes on genetic diversity within and among populations of P. griseoaptera.     

Genetic diversity and population differentiation in the endangered Siberian flying squirrel (<Emphasis Type="Italic">Pteromys volans</Emphasis>) in a fragmented landscape

Siberian flying squirrel (Pteromys volans) has declined in Finland and it is considered an endangered species. We studied microsatellite variation in four flying squirrel populations in a fragmented landscape in Finland to determine the amount of gene flow and genetic diversity in the populations. Demographic data from these areas suggest that the populations are declining. All the populations are significantly differentiated (F _ST = 0.23). The most notable result is the high degree of differentiation between two adjacent populations (F _ST = 0.11) and low genetic variability (number of alleles 3.0) in one of the populations. These findings suggest problems in dispersal and possible fragmentation effects in the landscape where only 10–20% of habitat favorable for the flying squirrel is left. Conservation ensuring dispersal should be urgently considered. Future studies should concentrate on the modeling of the population viability and on the effects of inbreeding in these small populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Constraints on home range behaviour affect nutritional condition in urban house sparrows (Passer domesticus)

In human‐dominated landscapes (semi)natural habitats are typically embedded in tracts of unsuitable habitat. Under such conditions, habitat characteristics and grain size of the surrounding landscape may affect how much food, and at what cost, is available for sedentary species with low home‐range plasticity. Here we combine behavioural radio‐tracking, feather ptilochronology, and landscape analysis to test how nutritional condition varies with home range size in 13 house sparrow [Passer domesticus (Linnaeus, 1758)] populations along an urban gradient. Urban individuals occupied smaller home ranges than conspecifics from rural areas, most distinctly if key cover was highly scattered. In urban plots, patch connectivity, home range sizes, and activity areas were positively correlated, indicating that individual ranging behaviour was related to the spatial distribution of suitable habitat. Urban House sparrows also showed the smallest feather growth bars, which were positively related to home range size at plot level. In contrast, growth bar widths and home range sizes were negatively related in rural populations, whereas in suburban populations, both variables varied independently. We conclude that individuals from progressively more built‐up areas show a restricted ability to adjust their daily ranging behaviour to the scattered distribution of critical resources. This may complement other putative causes of the widespread population decline of urban house sparrows. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 41–50.