Beyond the Mediterranean peninsulas: evidence of central European glacial refugia for a temperate forest mammal species, the bank vole (Clethrionomys glareolus)

This study details the phylogeographic pattern of the bank vole, Clethrionomys glareolus, a European rodent species strongly associated with forest habitat. We used sequences of 1011 base pairs of the mitochondrial DNA cytochrome b gene from 207 bank voles collected in 62 localities spread throughout its distribution area. Our results reveal the presence of three Mediterranean (Spanish, Italian and Balkan) and three continental (western, eastern and 'Ural') phylogroups. The endemic Mediterranean phylogroups did not contribute to the post-glacial recolonization of much of the Palaearctic range of species. Instead, the major part of this region was apparently recolonized by bank voles that survived in glacial refugia in central Europe. Moreover, our phylogeographic analyses also reveal differentiated populations of bank voles in the Ural mountains and elsewhere, which carry the mitochondrial DNA of another related vole species, the ruddy vole (Clethrionomys rutilus). In conclusion, this study demonstrates a complex phylogeographic history for a forest species in Europe which is sufficiently adaptable that, facing climate change, survives in relict southern and northern habitats. The high level of genetic diversity characterizing vole populations from parts of central Europe also highlights the importance of such regions as a source of intraspecific genetic biodiversity.     

Snow evens fragmentation effects and food determines overwintering success in ground-dwelling voles

Climate instability strongly affects overwintering conditions in organisms living in a strongly seasonal environment and consequently their survival and population dynamics. Food, predation and density effects are also strong during winter, but the effect of fragmentation of ground vegetation on ground-dwelling small mammals is unknown. Here, we report the results of a winter experiment on the effects of habitat fragmentation and food on experimental overwintering populations of bank voles Myodes glareolus. The study was conducted in large outdoor enclosures containing one large, two medium-sized or four small habitat patches or the total enclosure area covered with protective tall-grass habitat. During the stable snow cover of midwinter, only food affected the overwintering success, body condition, trappability and earlier onset of breeding in bank voles. However, after the snow thaw in spring, habitat fragmentation gained importance again, and breeding activities increased the movements of voles in the most fragmented habitat. The use of an open, risky matrix area increased along the habitat fragmentation. Our experiment showed that long-lasting stable snow cover protects overwintering individuals in otherwise exposed and risky ground habitats. We conclude that a stable winter climate and snow cover should even out habitat fragmentation effects on small mammals. However, along prolonged snow-free early winter and in an earlier spring thaw, this means loss of protection by snow cover both in terms of thermoregulation and predation. Thus, habitat cover is important for the survival of small ground-dwelling boreal mammals also during the non-breeding season.     

Vole mound effects and disturbance rate in a mediterranean plant community under different grazing and irrigation regimes

A factorial field experiment was used to assess the influence of soil-disturber mammals in the structure of a 9-year-old Mediterranean annual plant community subjected to different sheep grazing and irrigation regimes. We estimated the disturbance rate (mound building activity) by Mediterranean voles, their effects on vegetation and the mechanisms of these effects during a period of vole outbreak. The effects on vegetation were analysed at the levels of species, functional groups and plant community. Disturbance rate was high and voles can disturb the entire soil surface once every four or five years. The availability of certain trophic resources (perennial plants) appeared to drive vole expansion in the experimental plots and it was independent of the irrigation and grazing treatments. Mound building activities largely affected vegetation but conserved plot differences. Total vegetation cover, absolute cover of all functional groups, mean vegetation height and species richness were less on mounds than on undisturbed ground. These effects did not change the relative abundance of annuals, perennials, grasses and forbs. Only the relative abundance of small-seeded species decreased on mounds. As the proportion of these seeds was similar in both types of patches, we suggest that small-seeded species had more difficulties for germinating or emerging when they are buried during mound formation. Irrigation and sheep grazing promoted large changes in the vegetation parameters but these effects were, in general, similar on mounds and undisturbed ground. Our results show that the availability of germinable seeds may be the major limitation for mound revegetation, probably due to the scarcity of seeds existing at the depths from which soils are excavated. Our results also suggested a resource limitation on mounds. The results provide additional evidence that soil disturbances by small herbivore mammals exert relevant ecological effects on abandoned Mediterranean croplands. We discuss the ecological implications of vole mound-building activities for plant succession, plant species conservation and forage resource availability for livestock. This revised version was published online in June 2006 with corrections to the Cover Date.     

Spatio-temporal patterns of habitat use in voles and shrews modified by density, season and predators

1.?Although the intrinsic habitat preferences of a species can be considered to be fixed, the realized habitat use depends on the prevailing abiotic and biotic conditions. Often the core habitats are occupied by dense and stable populations, while marginal habitats become occupied only at times of high density. In a community of interacting species, habitat uses of different species become inter-related, for example an increased density of a strong competitor forcing a weaker competitor to use more marginal habitats. 2.?We studied the spatio-temporal distribution patterns of three common small mammal species, the bank vole Myodes glareolus; the field vole Microtus agrestis; and the common shrew Sorex araneus, in a 4-year trapping study carried out on six large islands, each containing a mixture of three main habitat types (forest, field and clear-cut). We experimentally released least weasels (Mustela n. nivalis) to some of the islands to see how the focal species respond to increased predation pressure. 3.?Both vole species were largely restricted to their core habitats (bank voles to forests and field voles to fields) at times of low population density. With increasing density, the relative habitat use of both species increased in the clear-cut areas. The common shrew was a generalist in its habitat use at all population densities. 4.?The release of the weasels changed the habitat use of all study species. 5.?The vole species showed a stronger aggregated pattern than the common shrew, especially at low population density. The vole aggregations remained in the same localities between seasons, except in the case of bank voles after the weasels were released. 6.?Bank voles and field voles avoided each other at high density. 7.?We conclude that intrinsically differential habitat requirements and flexibility to modify habitat use facilitate the coexistence of the two competing vole species in mosaic landscapes consisting of boreal forests and open habitats.     

A dynamic history of admixture from Mediterranean and Carpathian glacial refugia drives genomic diversity in the bank vole

Understanding the historical contributions of differing glacial refugia is key to evaluating the roles of microevolutionary forces, such as isolation, introgression, and selection in shaping genomic diversity in present‐day populations. In Europe, where both Mediterranean and extra‐Mediterranean (e.g., Carpathian) refugia of the bank vole (Clethrionomys glareolus) have been identified, mtDNA indicates that extra‐Mediterranean refugia were the main source of colonization across the species range, while Mediterranean peninsulas harbor isolated, endemic lineages. Here, we critically evaluate this hypothesis using previously generated genomic data (>6,000 SNPs) for over 800 voles, focusing on genomic contributions to bank voles in central Europe, a key geographic area in considering range‐wide colonization. The results provide clear evidence that both extra‐Mediterranean (Carpathian) and Mediterranean (Spanish, Calabrian, and Balkan) refugia contributed to the ancestry and genomic diversity of bank vole populations across Europe. Few strong barriers to dispersal and frequent admixture events in central Europe have led to a prominent mid‐latitude peak in genomic diversity. Although the genomic contribution of the centrally located Carpathian refugium predominates, populations in different parts of Europe have admixed origins from Mediterranean (28%–47%) and the Carpathian (53%–72%) sources. We suggest that the admixture from Mediterranean refugia may have provisioned adaptive southern alleles to more northern populations, facilitating the end‐glacial spread of the admixed populations and contributing to increased bank vole diversity in central Europe. This study adds critical details to the complex end‐glacial colonization history of this well‐studied organism and underscores the importance of genomic data in phylogeographic interpretation.     

Spatiotemporal dynamics of Puumala hantavirus in suburban reservoir rodent populations

The transmission of pathogens to susceptible hosts is dependent on the vector population dynamics. In Europe, bank voles (Myodes glareolus) carry Puumala hantavirus, which causes nephropathia epidemica (NE) in humans. Fluctuations in bank vole populations and epidemics in humans are correlated but the main factors influencing this relationship remain unclear. In Belgium, more NE cases are reported in spring than in autumn. There is also a higher incidence of human infections during years of large vole populations. This study aimed to better understand the link between virus prevalence in the vector, vole demography, habitat quality, and human infections. Three rodent populations in different habitats bordering Brussels city, Belgium, were studied for two years. The seroprevalence in voles was influenced first by season (higher in spring), then by vole density, vole weight (a proxy for age), and capture site but not by year or sex. Moreover, voles with large maximal distance between two captures had a high probability for Puumala seropositivity. Additionally, the local vole density showed similar temporal variations as the number of NE cases in Belgium. These results showed that, while season was the main factor influencing vole seroprevalence, it was not sufficient to explain human risks. Indeed, vole density and weight, as well as the local habitat, were essential to understanding the interactions in these host‐pathogen dynamics. This can, in turn, be of importance for assessing the human risks.     

Reversal of fortune: plant suppression and recovery after vole herbivory

It is not clear how plant species preferred as forage by rodents persist in prairie vegetation. To test permanence of suppression of wet-mesic prairie vegetation by vole (Microtus pennsylvanicus) herbivory in synthetic experimental communities, access treatments were reversed after 9 years of vole exclusion or access. Between 1996 and 2004, rye grass Elymus virginicus (Poaceae) and tick-trefoil Desmodium canadense (Fabaceae) achieved mean cover of up to 30 and 25%, respectively, in plots where voles were excluded, but disappeared from plots where voles had access. To determine whether these species remained vulnerable to vole herbivory as established adults, and to determine whether the species could recover if vole herbivory were removed, access treatments were reversed at the end of the 2004 growing season and monitored through 2007. Repeated measures ANOVA demonstrated dramatic vole suppression of established E. virginicus, but not D. canadense, indicating continuing vulnerability of the grass but not of the adult legume. Release from vole herbivory resulted in re-growth of rye, but not tick-trefoil, which was apparently suppressed by established vegetation. Two additional common planted species did not respond to treatment reversal, nor did 11 much less common planted species that comprised a minor portion of the vegetation. Dominant perennial black-eyed Susan Rudbeckia subtomentosa (Asteraceae) did not change in plant numbers by year or treatment, but expanded or contracted in stems per plant and cover as E. virginicus was suppressed or released by vole herbivory or its absence. Results indicate that preferred food plants may persist through capacity to quickly recover during periods of relative vole scarcity, or reach a refuge in maturity.     

Winter herbivory by voles during a population peak: the relative importance of local factors and landscape pattern

1. We studied the relative role of local habitat variables and landscape pattern on vole–plant interactions in a system with grey-sided voles ( Clethrionomys rufocanus (Sund.)) and their favourite winter food plant, bilberry ( Vaccinium myrtillus L.). The study was conducted during a vole peak year (1992–93) in a tundra area in northern Norway.
2. Using Mantel statistics we were able to separate the direct effects of the spatial patterning of habitats and the indirect effects due to spatial aggregations of similar habitats.
3. Results indicate that knowledge about the explicit spatial patterning of patches does not improve our understanding of the system. Instead, two local factors, vegetation height and bilberry biomass, explained more than 50% of the variation in cutting intensity in winter (defined as the proportion of above-ground shoots cut). Increasing vegetation height increased, and increasing bilberry biomass decreased, the cutting intensity.
4. The conclusion that grey-sided voles are able to distribute themselves relative to habitat quality was also partially supported by our estimated over-winter persistence by voles in the various habitats. Vole persistence was uncorrelated with vegetation height, the important predictor of autumn vole density, but tended to correlate with the deviation from the relation between vegetation height and autumn vole density. This conforms to the expectations from the theory of ideal-free habitat distribution.
5. The cue for vole habitat choice, i.e. vegetation height, indicates that either predation or freezing risk is important for voles when selecting over-wintering habitat.     

European map of alien plant invasions based on the quantitative assessment across habitats

Aim Recent studies using vegetation plots have demonstrated that habitat type is a good predictor of the level of plant invasion, expressed as the proportion of alien to all species. At local scale, habitat types explain the level of invasion much better than alien propagule pressure. Moreover, it has been shown that patterns of habitat invasion are consistent among European regions with contrasting climates, biogeography, history and socioeconomic background. Here we use these findings as a basis for mapping the level of plant invasion in Europe. Location European Union and some adjacent countries. Methods We used 52,480 vegetation plots from Catalonia (NE Spain), Czech Republic and Great Britain to quantify the levels of invasion by neophytes (alien plant species introduced after ad 1500) in 33 habitat types. Then we estimated the proportion of each of these habitat types in CORINE land‐cover classes and calculated the level of invasion for each class. We projected the levels of invasion on the CORINE land‐cover map of Europe, extrapolating Catalonian data to the Mediterranean bioregion, Czech data to the Continental bioregion, British data to the British Isles and combined Czech–British data to the Atlantic and Boreal bioregions. Results The highest levels of invasion were predicted for agricultural, urban and industrial land‐cover classes, low levels for natural and semi‐natural grasslands and most woodlands, and the lowest levels for sclerophyllous vegetation, heathlands and peatlands. The resulting map of the level of invasion reflected the distribution of these land‐cover classes across Europe. Main conclusions High level of invasion is predicted in lowland areas of the temperate zone of western and central Europe and low level in the boreal zone and mountain regions across the continent. Low level of invasion is also predicted in the Mediterranean region except its coastline, river corridors and areas with irrigated agricultural land.     

Niche differentiation in a postglacial colonizer,the bank vole Clethrionomys glareolus

Species‐level environmental niche modeling has been crucial in efforts to understand how species respond to climate variation and change. However, species often exhibit local adaptation and intraspecific niche differences that may be important to consider in predicting responses to climate. Here, we explore whether phylogeographic lineages of the bank vole originating from different glacial refugia (Carpathian, Western, Eastern, and Southern) show niche differentiation, which would suggest a role for local adaptation in biogeography of this widespread Eurasian small mammal. We first model the environmental requirements for the bank vole using species‐wide occurrences (210 filtered records) and then model each lineage separately to examine niche overlap and test for niche differentiation in geographic and environmental space. We then use the models to estimate past [Last Glacial Maximum (LGM) and mid‐Holocene] habitat suitability to compare with previously hypothesized glacial refugia for this species. Environmental niches are statistically significantly different from each other for all pairs of lineages in geographic and environmental space, and these differences cannot be explained by habitat availability within their respective ranges. Together with the inability of most of the lineages to correctly predict the distributions of other lineages, these results support intraspecific ecological differentiation in the bank vole. Model projections of habitat suitability during the LGM support glacial survival of the bank vole in the Mediterranean region and in central and western Europe. Niche differences between lineages and the resulting spatial segregation of habitat suitability suggest ecological differentiation has played a role in determining the present phylogeographic patterns in the bank vole. Our study illustrates that models pooling lineages within a species may obscure the potential for different responses to climate change among populations.     

Environmental change and disease dynamics: effects of intensive forest management on Puumala hantavirus infection in boreal bank vole populations

Intensive management of Fennoscandian forests has led to a mosaic of woodlands in different stages of maturity. The main rodent host of the zoonotic Puumala hantavirus (PUUV) is the bank vole (Myodes glareolus), a species that can be found in all woodlands and especially mature forests. We investigated the influence of forest age structure on PUUV infection dynamics in bank voles. Over four years, we trapped small mammals twice a year in a forest network of different succession stages in Northern Finland. Our study sites represented four forest age classes from young (4 to 30 years) to mature (over 100 years) forests. We show that PUUV-infected bank voles occurred commonly in all forest age classes, but peaked in mature forests. The probability of an individual bank vole to be PUUV infected was positively related to concurrent host population density. However, when population density was controlled for, a relatively higher infection rate was observed in voles trapped in younger forests. Furthermore, we found evidence of a "dilution effect" in that the infection probability was negatively associated with the simultaneous density of other small mammals during the breeding season. Our results suggest that younger forests created by intensive management can reduce hantaviral load in the environment, but PUUV is common in woodlands of all ages. As such, the Fennoscandian forest landscape represents a significant reservoir and source of hantaviral infection in humans.     

Gene conversion in the mitochondrial genome on interspecific hybridization in voles of the <Emphasis Type="Italic">Clethrionomys</Emphasis> genus

The phenomenon of interspecific hybridization accompanied by transfer of the mitochondrial genome from the northern red-backed vole (Clethrionomys rutilus) to the bank vole (Cl. glareolus) in northeastern Europe is well known already for 25 years. However, the possibility of recombination between homologous segments of maternal and paternal mtDNAs of the voles during fertilization was not previously studied. Analysis of data on variability of nucleotide sequences of the mitochondrial gene for cytochrome b in populations of red-backed and bank voles in the area of their sympatry has shown that as a result of interspecific hybridization, the mitochondrial gene pool of bank voles contains not only mtDNA haplotypes of red-backed vole females, but also mtDNA haplotypes of bank voles bearing short nucleotide tracts of red-backed vole mtDNA. This finding supports the hypothesis that an incomplete elimination of red-backed vole paternal mtDNA during the interspecific hybridization between bank vole females and red-backed vole males leads to the gene conversion of bank vole maternal mtDNA tracts by homologous ones of mtDNA of red-backed vole males.     

Natural infections of small mammals with blood parasites on the borderland of boreal and temperate forest zones

Blood parasites of small mammals living in Białowieża Forest (eastern Poland) were investigated between 1996 and 2002. The following haemoparasite species were found:Trypanosoma (Herpetosoma) evotomys in bank voleClethrionomys glareolus; T. (H.) microti in root voleMicrotus oeconomus; Babesia microti in root vole;Hepatozoon erhardovae in bank vole andHepatozoon sp. in root vole. Some non-identifiedBartonella species were found in bank vole, root vole, field voleMicrotus agrestis, yellow-necked mouseApodemus flavicollis, common shrewSorex araneus, Eurasian water shrewNeomys fodiens, and Mediterranean water shrewN. anomalus. The prevalence and diversity of blood parasites were lower in shrews than small rodents. Totally, 52.0% of bank voles, 50.0% of root voles, 32.5% of common shrews, and 41.2% of Eurasian water shrews were infected with any of the blood parasites. Mixed infections were seldom observed in bank vole (17.3% of investigated individuals) and root vole (14.7%). No animals were infected with three or four parasites simultaneously. Infection of Białowieża small mammals with haemoparasites seemed to be similar to those described in other temperate forest regions rather than boreal ones. Infection rates of rodent species seem to be higher in their typical habitats: for bank vole it was the highest in mixed forest, whereas for root vole in sedge swamp. The results suggest that Arvicolidae play a greater role than Muridae or Soricidae in maintenance ofBabesia andHepatozoon foci in natural environments of central Europe.     

Competition, predation and interspecific synchrony in cyclic small mammal communities

Although competition and predation are considered to be among the most important biotic processes influencing the distribution and abundance of species in space and time, the relative and interactive roles of these processes in communities comprised of cyclically fluctuating populations of small mammals are not well known. We examined these processes in and among populations of field voles, sibling voles, bank voles and common shrews in western Finland, using spatially replicated trapping data collected four times a year during two vole cycles (1987–1990 and 1997–1999). Populations of the four species exhibited relatively strong interspecific temporal synchrony in their multiannual fluctuations. During peak phases, we observed slight deviations from close temporal synchrony: field vole densities peaked at least two months earlier than those of either sibling voles or bank voles, while densities of common shrews peaked even earlier. The growth rates of all four coexisting small mammal species were best explained by their own current densities. The growth rate of bank vole populations was negatively related to increasing densities of field voles in the increase phase of the vole cycle. Apart from this, no negative effects of interspecific density, direct or delayed, were observed among the vole species. The growth rates of common shrew populations were negatively related to increasing total rodent (including water voles and harvest mice) densities in the peak phase of the vole cycle. Sibling voles appeared not to be competitively superior to field voles on a population level, as neither of these Microtus voles increased disproportionately in abundance as total rodent density increased. We suggest that interspecific competition among the vole species may occur, but only briefly, during the autumn of peak years, when the total available amount of rodent habitat becomes markedly reduced following agricultural practices. Our results nonetheless indicate that interspecific competition is not a strong determinant of the structure of communities comprised of species exhibiting cyclic dynamics. We suggest that external factors, namely predation and shortage of food, limit densities of vole populations below levels where interspecific competition occurs. Common shrews, however, appear to suffer from asymmetric space competition with rodents at peak densities of voles; this may be viewed as a synchronizing effect.     

The dynamics and regulation of small rodent populations in the woodland ecosystems of Killarney, Ireland

Populations of Wood mice Apodemus sylvaticus and Bank voles Clelhrionomys glareolus were studied by mark-recapture techniques in five different woodlands but concentrated on an oak Quercus petraea wood and a yew Taxus baccata wood, where food supply was also investigated. Much of the yew wood was on limestone pavement which was greatly creviced thus providing static and quantifiable cover. Indices of the intensity with which food was being searched for were obtained on all areas by seed-removal experiments.
Trappability varied within the populations and seasonally, being highest in summer. Unusually high densities of mice were recorded in the yew wood but not elsewhere; vole densities were always low. There was an inverse relationship between range length and density: mean range length in the yew wood was exceptionally low. Males generally ranged further than females, voles further than mice. Females outnumbered males in the oak wood, where cover was minimal, when the population was low. In the two areas with most cover, males outnumbered females. In a further area, where numbers were initially exceedingly low, the population consisted entirely of males for almost a year. Sexes differed in habitat preference but numbers, especially of males, were significantly correlated with cover. Voles usually bred for shorter periods than mice. Overwinter breeding and increased overwinter survival followed good seed years. Individuals recruited when food was plentiful lived longest. In seed-removal experiments rate of removal increased with shortage of natural seed.
Food and intraspecific aggression were the probable major factors in regulating population size, of mice at least. Cover probably modifies intrinsic mechanisms by reducing numbers of encounters between individuals.     

Habitat selection by Southern water vole (Arvicola sapidus) in riparian environments of Mediterranean mountain areas: a conservation tool for the species

The Southern water vole, Arvicola sapidus, is endemic to the Iberian Peninsula and France. Despite being catalogued as vulnerable, our current knowledge of this species is not sufficient to establish measures for its conservation and recovery, particularly in riparian zones of Mediterranean mountain areas. The aim of this study was to identify factors related to habitat configuration that determine the presence or absence of the species in the Montsant River. Specifically, we associated the presence/absence of this arvicolid rodent with composition of vegetation, river bank morphology, and watercourse characteristics. The results suggest that, in this area, the most favorable places for the species are those with a high degree of cover of herbaceous plants (mainly helophytes) and moderate to low levels of tree and shrub cover; gently sloping banks and a soft substrate; and the presence of water, with moderate to high stream widths and depths. In addition, we developed a classification method which allowed us to classify and characterize habitat conditions: the optimal scenario (preferential for arvicoline establishment), the suboptimal scenario (whose use is related to opportunities to find best scenarios), and the hostile scenario (not acceptable for use). In such riparian areas, the results revealed that the Southern water vole is a specialist in terms of habitat selection, but behaves as a generalist in terms of occupancy. Its ability to adapt to suboptimal conditions widens the options for managing Southern water vole populations, and indicates that the maintenance and rehabilitation of habitat along continuous stretches of river is the most effective approach to achieving self-sustaining populations.     

Reproductive investment under fluctuating predation risk: Microtine rodents and small mustelids

Summary We studied the reproductive investment of microtine rodents (bank vole (Clethrionomys glareolus),Microtus epiroticus andMicrotus agrestis) in western Finland under predation risk from small mustelids. During 1984–1992, the yearly mean litter size of overwintered bank voles was smaller at high least weasel and stoat densities than at low densities (close to 3 versus 4–5). In addition, the annual mean litter size of young bank voles was negatively correlated to the least weasel density. In youngM. agrestis voles, the yearly late summer litter size was negatively associated with the autumn density of small mustelids. In the crash phase of the vole cycle (1989 and 1992), we removed small mustelids (mainly least weasels) from four unfenced areas in late April to late May and studied the reproduction of voles in four removal and comparable control areas (each 2–4 km²). Reduction of small mustelids significantly increased the proportion of pregnant bank vole females, but not that of pregnantMicrotus vole females. We conclude that predation risk apparently reduced reproductive investment of free-living bank vole females; these voles appear to trade their current parental investment against future survival and reproductive prospects. Accordingly, the presence of small mustelids (or their scent) may slow down the reproductive rate of voles. As antipredatory behaviours occurred on a large scale, our results add evidence to the hypothesis that crashes in multiannual vole cycles are driven by small mustelid predators.     

Small mammal responses to moose supplementary winter feeding

Supplementary feeding of wild large herbivores is a widespread practice in North America and Europe. The presence of feeding stations may have ecological consequences through changes to animal distributions, patterns of herbivory and a net nutrient input into the ecosystem. In Fennoscandia, supplementary feeding of moose in winter (Alces alces) is increasing. Although it has been shown to affect bird communities, its effects on small mammal communities were unknown. Here, we studied the effects of moose supplementary feeding stations on plants and on abundance, reproduction, and biomass of small mammals in years with low and high vole abundance. We sampled small mammals with snap traps and conducted surveys of the field layer vegetation, at varying distances from moose supplemental feeding stations. Due to the vegetation changes induced by feeding stations, abundance of common shrews (Sorex araneus) and Microtus voles were positively affected by long-term moose winter feeding, while bank voles (Myodes glareolus) were not affected. Moose feeding stations did not affect reproduction, individual body mass, or the total biomass of small mammals. Moose winter-feeding stations have impacts on nontarget species, providing islands of preferred grass and forb habitat for Microtus spp. and common shrews, allowing them to penetrate into a matrix of less preferred forest habitat.     

Local and Regional Determinants of Colonisation-Extinction Dynamics of a Riparian Mainland-Island Root Vole Metapopulation

The role of local habitat geometry (habitat area and isolation) in predicting species distribution has become an increasingly more important issue, because habitat loss and fragmentation cause species range contraction and extinction. However, it has also become clear that other factors, in particular regional factors (environmental stochasticity and regional population dynamics), should be taken into account when predicting colonisation and extinction. In a live trapping study of a mainland-island metapopulation of the root vole (Microtus oeconomus) we found extensive occupancy dynamics across 15 riparian islands, but yet an overall balance between colonisation and extinction over 4 years. The 54 live trapping surveys conducted over 13 seasons revealed imperfect detection and proxies of population density had to be included in robust design, multi-season occupancy models to achieve unbiased rate estimates. Island colonisation probability was parsimoniously predicted by the multi-annual density fluctuations of the regional mainland population and local island habitat quality, while extinction probability was predicted by island population density and the level of the recent flooding events (the latter being the main regionalized disturbance regime in the study system). Island size and isolation had no additional predictive power and thus such local geometric habitat characteristics may be overrated as predictors of vole habitat occupancy relative to measures of local habitat quality. Our results suggest also that dynamic features of the larger region and/or the metapopulation as a whole, owing to spatially correlated environmental stochasticity and/or biotic interactions, may rule the colonisation – extinction dynamics of boreal vole metapopulations. Due to high capacities for dispersal and habitat tracking voles originating from large source populations can rapidly colonise remote and small high quality habitat patches and re-establish populations that have gone extinct due to demographic (small population size) and environmental stochasticity (e.g. extreme climate events).     

The combined effect of environmental factors and neighbouring populations on the distribution and abundance of Arvicola terrestris. An approach using rule-based models

We investigated how the interplay between environmental factors and presence of neighbouring populations determines the distribution and abundance of a small, endangered rodent, the water vole ( Arvicola terrestris ). We studied thriving and non-fragmented populations of water voles in the absence of their main predator, the introduced American mink ( Mustela vison ). A low degree of population fragmentation, such as the one characterizing the studied populations, was probably typical of water voles before their decline started. We found that under these conditions water voles' distribution is mainly determined by three environmental factors: presence of freshwater, adequate food, and cover. Variance in other factors is well tolerated by water voles. We obtained this result by the use of rule-based models in two separate areas. The two models correctly classified 81% and 83% of the observed cases, respectively. When optimised on one area and cross-validated on the other area the performance of the models was still high (73% and 79%) indicating that the models were robust and generalizable. We also found that the density of animals was lower in sub-optimal than in optimal habitat. We then tested the hypothesis that the number of neighbouring colonies determines the probability of finding voles in a given section. We found that the presence of nearby colonies was an important factor in determining the presence of water voles in sub-optimal habitat, while isolated patches of suitable habitat were less likely to host water voles. These observations suggest the possible presence of a source-sink dynamic, where an optimal habitat acts as a source for populating sub-optimal habitats that may be considered a sink habitat. These findings are discussed in the context of water vole conservation.