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Maternal stress can have long-term adverse consequences on immunocompetence and disease risk of offspring, and winter survival is a crucial demographic parameter in the life-history of an individual that can substantially affect northern rodent population dynamics. An understanding of the effects of maternal stress on winter survival of offspring may help identify mechanisms driving population fluctuations of northern small mammals. Thus, we assessed the effects of maternal stress, resulting from high population densities, on winter survival of first generation (F(1)) and second generation (F(2)) in root voles Microtus oeconomus. Replicate high- and low-density enclosed parental populations were established, from which we obtained F(1) generation that were used to establish new enclosed, equal-density populations. The adults of the high-density parental populations had higher corticosterone levels, an indication of physiological stress, than did those of the low-density parental populations. Over-winter survival of the F(1) generation voles from the low-density parental populations was greater than that of those from the high-density parental populations. Over-winter survival of F(2) generation voles did not differ between the two treatments. Our results suggest that maternal stress affected over-winter survival of first generations but not second generations. Reduced immunocompetence, resulting from high population density stresses, transferred to offspring may be a factor in annual (winter) population declines. Because the effect is transitory, i.e. immunocompetence of F(2) voles is not affected, reduced immunocompetence resulting from high density stresses would not contribute to lengthy periods of low population densities that are characteristic of multi-annual population fluctuations.

Dispersal is one of the most important, yet least understood phenomena of evolutionary ecology. Triggers and consequences of dispersal are difficult to study in natural populations since dispersers can typically only be identified a posteriori. Therefore, a lot of work on dispersal is either of a theoretical nature or based on anecdotal observation. This is especially true for cryptic species such as small mammals. We conducted an experiment on the common vole, Microtus arvalis, in semi-natural enclosures and investigated the spatial and genetic establishment success of residents and dispersers in their natal and new populations. Our study uses genetic data on the reproductive success of 1255 individuals to measure the fitness trajectories of the residents and dispersing individuals. In agreement with past studies, we found that dispersal was highly male-biased, and was most probably induced by the agonistic encounters with conspecifics, suggesting it could act as an inbreeding avoidance mechanism. There was low breeding success of dispersers into new populations. Although nearly 26% of identified dispersers reproduced in their natal populations, only seven percent reproduced in the new populations. Settlement appeared to be a pre-requisite for reproduction in both sexes, and animals that did not spatially settle into a new population dispersed again, usually on the same day of immigration. In the event that dispersers reproduced in the new population, they did so at relatively low population densities. We also found age-related differences between the sexes in breeding success, and male dispersers that subsequently established in the new population were young individuals that had not reproduced in their natal population, whereas successful females had already reproduced in their natal population. In conclusion, with our detailed field data on establishment and substantial parentage assignments to understand breeding success, we were able to gain an insight into the fitness of dispersers, and how the two sexes optimise their fitness. Taken together, our results help to further understand the relative advantages and costs of dispersal in the common vole.

Competing species benefit from eavesdropping on each other's signals by learning about shared resources or predators. But conspicuous signals are also open to exploitation by eavesdropping predators and should also pose a threat to other sympatric prey species. In western Finland, sibling voles Microtus rossiameridionalis and field voles M. agrestis compete for food and space, and both species rely upon scent marks for intraspecific communication. Both vole species are prey to a range of terrestrial scent hunting predators such as least weasels, however, the competitively superior sibling voles are taken preferentially. We tested in large out-door enclosures whether field voles eavesdrop on the signals of its competitor, and whether they behave as though this eavesdropping carries a risk of predation. We presented field voles with scent marks from unknown conspecifics and sibling voles and measured their visitation, activity and scent marking behaviours at these scents under high (weasel present) and low (weasel absent) predation risk. Field voles readily visited both field and sibling vole scents under both high and low predation risk; however their activity at sibling vole scent marks declined significantly under increased predation risk. In contrast, predation risk did not affect field voles’ activity at conspecific scents. Thus, field voles were compelled to maintain eavesdropping on heterospecific scents under an increased risk of predation, however they compensated for this additional risk by reducing their activity at these risky scents. Scent marking rates declined significantly under high predation risk. Our results therefore reveal a hidden complexity in the use of social signals within multi-species assemblages that is clearly sensitive to the potential for increased predation risk. The predation risks of interspecific eavesdropping demonstrated here represents a significant generalisation of the concept of associational susceptibility.

We studied patterns of species co-occurrence in communities of ectoparasitic arthropods (ixodid ticks, mesostigmate mites and fleas) harboured by rodent hosts from South Africa (Rhabdomys pumilio), South America (Scapteromys aquaticus and Oxymycterus rufus) and west Siberia (Apodemus agrarius, Microtus gregalis, Microtus oeconomus and Myodes rutilus) using null models. We compared frequencies of co-occurrences of parasite species or higher taxa across host individuals with those expected by chance. When non-randomness of parasite co-occurrences was detected, positive but not negative co-occurrences of parasite species or higher taxa prevailed (except for a single sample of mesostigmate mites from O. rufus). Frequency of detection of non-randomness of parasite co-occurrences differed among parasite taxa, being higher in fleas and lower in mites and ticks. This frequency differed also among host species independent of parasite taxon, being highest in Microtus species and lowest in O. rufus and S. aquaticus. We concluded that the pattern of species co-occurrence in ectoparasite communities on rodent hosts is predominantly positive, depends on life history of parasites and may be affected to a great extent by life history of a host.

Large predators may affect the hunting efficiency of smaller ones directly by decreasing their numbers, or indirectly by altering their behaviour. Either way this may have positive effects on the density of shared prey. Using large outdoor enclosures, we experimentally studied whether the presence of the Tengmalm's owl Aegolius funereus affects the hunting efficiency of the smallest member of the vole-eating predator guild, the least weasel Mustela nivalis, as measured by population responses of coexisting prey species, the field vole Microtus agrestis and the sibling vole M. levis. We compared the density and survival probability of vole populations exposed to no predation, weasel predation or combined predation by a weasel and an owl. The combined predation of both owl and weasel did not result in obvious changes in the density of sibling and field vole populations compared to the control populations without predators, while predation by least weasel alone decreased the densities of sibling voles and induced a similar trend in field vole densities. Survival of field voles was not affected by predator treatment while sibling vole survival was lower in predator treated populations than in control populations. Our results suggest that weasels are intimidated by avian predators, but without changing the effects of predators on competitive situations between the two vole species. Non-lethal effects of intraguild predation therefore will not necessarily change competitive interactions between shared prey species.

International audience; Population structure, in terms of the body mass, condition, sex and reproductive status of individuals, has been found to vary in phase with population density in cyclic populations of microtine rodents. Because sustained population cycles involve delayed density dependent changes in the population growth rate, we would expect at least some life history traits also to depend on past densities. Detailed, long-term studies of changes in vole life history traits are however few, and are largely restricted to northern Europe. In view of the uncertainty as to whether the cyclic microtine populations of western Europe represent the same phenomenon as those of northern Europe, we studied temporal variation in the structure of a clearly cyclic population of the common vole Microtus arvalis Pallas, in the cereal plains of mid-western France. Our data set contains seasonal, individual-level data from long-term, large-scale trapping covering four entire population cycles. We found considerable cyclic variation in population structure in spring (April), but less so in summer (June). In spring of post-peak years, animals were of low body weight and body condition (particularly females), litter sizes were smaller and there was a reduction in the proportion of breeders. All of these could be proximal drivers of increased mortality rates, or decreased birth rates, contributing to the population declines. Few life history traits, however, showed direct density dependent variation, and none of the traits studied here showed delayed density dependence. We have shown declines in the fecundity and body condition of voles from a western European population that coincides with, and may be a proximal cause of, cyclic declines in population density. Closer attention to proximal causes, by which ecological processes drive cycles, could clarify the extent to which microtine cycles across Europe represent a single phenomenon.

Interspecific competition is usually understood as different species competing directly with each other for limited resources. However, predators can alter such competitive interactions substantially. Predation can promote the coexistence of species in a situation where it would otherwise be impossible, for example if a tradeoff between the competitive abilities and predation resistance of the prey species exists. The field vole Microtus agrestis and the sibling vole M. rossiaemeridionalis are sympatric grassland species, which compete for the same resources. At the population level sibling voles are suggested to be superior competitors to field voles, yet more vulnerable to predation. We tested the effects of predation on the two species in 0.5 ha outdoor enclosures by exposing vole populations to radio-collared freely-hunting least weasels Mustela nivalis nivalis for three weeks. Lethal and non-lethal impacts of predation limited population densities of both species during and after the experimental period, but the effect was more pronounced in sibling voles in which population densities decreased markedly during the treatment period and even after that. Field vole population densities remained stable under weasel predation, while densities increased in controls. Survival in both species was lower in treatment populations compared to controls, but the effect tended to be more pronounced in sibling voles and in females of both species. The average mass of adults in both species declined in the treatment populations. These results suggest that predation by least weasels can limit vole populations locally, even during favourable summer conditions, and have extended negative effects on the dynamics of vole populations. In addition, predation alleviated interspecific competition between the vole species and is, therefore, a potential factor enabling the coexistence of them.

Vole–vegetation interactions in a predation-free taiga environment of northern Fennoscandia were studied by transferring vegetation from natural Microtus habitats into a greenhouse, where three hab ...

Lack of food resources has been suggested as a factor which limits the growth of cyclic vole populations. During peak phases of the cycle, vole population growth typically ceases during late autumn or early winter, and is followed by a decrease in density over the winter. To investigate whether this decrease is due to increased mortality induced by a depletion of food resources, we studied overwinter food consumption and physiological condition of field voles (Microtus agrestis) in western Finland in both an increase and a decrease phase of a three-year population cycle. The growth rate of vole populations was negatively related both to prevailing vole densities and to densities six months earlier. The condition index of voles, as well as their blood levels of haematocrit, proteins, free fatty acids and immunoglobulin G, were positively related to population growth rate when populations were declining. When populations were increasing, these parameters tended to be negatively related to population growth rate. The overall physiological condition of voles was lower in the winter of the decrease phase as compared to the increase phase. The return rate of voles, a proxy of survival, was also lower in the decrease than in the increase phase of the cycle and positively related to haematocrit levels. Almost 90% of all green vegetation shoots were consumed by voles during the winter of the decrease phase while only two thirds were eaten in the increase phase. Our results suggest that the winter decrease phase of cyclic vole populations is associated with both a deterioration in the physiological condition of voles and a significant depletion of winter food resources. This implies that malnutrition induces poor physiological condition in voles, which in turn may increase mortality either directly through starvation or indirectly through increased susceptibility to predators and pathogens.

Interspecific competition is assumed to generate negative effects on coexisting species, possibly including slower population growth and lower survival. The field vole (Microtus agrestis) and the sibling vole (M. rossiaemeridionalis) are sympatric close relatives which compete for similar resources. Previous non-experimental studies suggest that the smaller sibling vole is a superior competitor, yet more vulnerable to predation than the larger field vole. We studied the effects of coexistence on population densities, reproductive parameters, and survival in these two species by means of experimentation in large, predator-free outdoor enclosures. While populations of both species reached higher densities in the absence of the other, field voles appeared to suffer more from interspecific competition than sibling voles. The proportion of young individuals in the population was higher in the sibling vole than in the field vole at the end of the experiment. The presence of a coexisting species reduced the survival of field voles. Sibling voles, on the other hand, appeared to suffer more from intraspecific competition than interspecific competition. On a population level, the sibling vole seems to be a superior competitor in the absence of predators due to better survival and possibly a higher reproductive capacity. However, predation probably has a profound influence on the interspecific dynamics of these two species indicating that in natural surroundings apparent competition (i.e. competition via shared predators) is stronger than direct competition.

We conducted a field experiment that manipulated landscapes by mowing so that the amount of unfavorable habitat (low cover) for prairie voles (Microtus ochroeaster) increased while the number and size of favorable patches (high cover) remained constant. Distance between favorable patches increased as the amount of unfavorable habitat increased, so we could test two current hypotheses concerning the effect of habitat fragmentation on local populations: 1) increased distance between favorable habitat patches reduces successful per capita dispersal (emigration and immigration) because dispersers suffer greater exposure to predators (the predation hypothesis); and 2) per capita dispersal is inversely density dependent in voles because increased aggression at higher density inhibits movements (the social fence hypothesis). As predicted by the predation hypothesis, increased distance between favorable habitat patches led to decreased successful dispersal among patches and increased per capita mortality, particularly among subadult and adult males (the categories of voles most likely to emigrate). As predicted by the social fence hypothesis, dispersal was inversely density dependent, and dispersing voles displayed a greater frequency of wounding (an indication of increased aggressive interactions) than did residents. The amount of wounding in general did not increase with density, however, and, as distance between patches increased to 60 m, successful dispersal became rare and erratic. Nevertheless, our overall results supported current hypotheses regarding the effects of increased habitat fragmentation on patterns of dispersal and mortality. Examining the impact of these effects on local population dynamics within different landscapes will require longer periods of observation.

Three mechanisms have been proposed to induce spatial synchrony in fluctuations of small mammal populations: climate-related environmental effects, predation and dispersal. We conducted a field experiment in western Finland to evaluate the relative roles of these mechanisms in inducing spatial synchrony among cyclic populations of field voles Microtus agrestis. The study was conducted during the increase and peak phases of a vole population cycle on four agricultural field sites situated 1.5-7.0 km apart. Each field contained two 0.5-ha fenced enclosures and one 1-ha unfenced control area. One enclosure per field allowed access by small mustelid predators and the other by avian predators; all enclosures prevented the dispersal of voles. The unfenced control areas allowed access by all predators as well as dispersal by voles. Enclosed vole populations were in a treatment-wise asynchronous phase before the predator access treatments were applied. The growth rates of all enclosed populations were tightly synchronized during the course of the experiment. Conversely, synchrony both among the unfenced populations and between the fenced and unfenced populations was practically non-existent. During winter, in the increase phase of the cycle, vole populations in all treatments declined to low densities due to a seasonal effect of winter food depletion. During summer, in the peak year of the vole cycle, all populations fluctuated in synchrony. At this time, both small mustelids and birds of prey appeared to be abundant enough to induce synchrony. Dispersal was identified as a potential contributor to synchronization, but the magnitude of its effects could not be reliably discerned. Our results indicate that no single mechanism can account for the observed patterns of spatial synchrony among cyclic northern vole populations. Rather, spatial synchronization is induced by different mechanisms, namely seasonality and predation, acting successively during different seasons and phases of the vole cycle.

This paper reports the effects of food supply, predation and the interaction between them on the population dynamics of root voles, Microtus oeconomus, by adopting factorial experiments in field enclosures. This two-factor experiment proved the general hypothesis that food supply and predation had independent and additive effects on population dynamics of root voles. The experimental results proved the following predictions: (1) predation reduced population density and recruitment significantly; (2) food supply increased population density; (3) predation and food supply influenced spacing behavior of root voles separately and additively: Exposure to predation reduced long movements of root voles between trapping sessions; additional food supply reduced aggression level and home range size of root voles. Less movement of individuals that exposed to predators possibly reduced their opportunity of obtaining food and lessened population survival rate, which led population density to decrease. Smaller home range and lower aggression level could make higher population density tolerable. The interactive effect of predation and food on home range size was highly significant (P =0.0082

During 1991–95, mammalian predators (weasel, Mustela nivalis, stoat, M. erminea, mink, M. vison, and red fox, Vulpes vulpes) were excluded in late summers from a 2 ha piece of a north Norwegian mountain slope. The exclosure extended from an outpost of luxuriant sub-arctic birch forest to a typical arctic–alpine habitat complex, including productive willow scrublands, tundra heaths, dry ridges and snow-beds. The exclosure thus encompassed the entire range of habitat conditions encountered in a typical north Fennoscandian mountain and tundra landscapes. During 1991–95, the exclosure was predator-proof from late July to late September. In wintertime and in early summer, the exclosure was accessible to mammalian predators. Vole dynamics in the short-term exclosure were compared to dynamics in five reference areas with similar habitat conditions. In 1991, when vole densities were rising in the area, neither collective vole densities nor densities of individual vole species differed significantly between the exclosure and the replicated controls. Spring densities of voles were never significantly different between the exclosure and the controls. With respect to autumnal densities of voles, however, the exclosure was a statistical outlier in the peak year 1992 and throughout the gradual decline phase of 1993–95. In the peak year, the difference in collective vole densities was modest (30%), but increased to two-fold during the first two decline years and was almost four-fold in the crash year of 1995. The strongest response was displayed by field voles (Microtus agrestis), hypothesized to be the pivotal prey species of weasels, especially by females and young individuals, i.e. by the functional categories especially sensitive to mammalian predation. These results are consistent with the hypothesis that predation plays a pivotal role for the regulation of herbivorous mammals in relatively productive arctic–alpine habitats.

Barriers that dissect continuous habitat, such as roads, initiate the process of habitat fragmentation and can filter or eliminate animal movement through otherwise pristine areas. A severe road barrier appears to exist in Banff National Park where the Trans-Canada Highway (TCH) is known to impede the movement of some animals, but the effects of this barrier relative to natural barriers and continuous habitat have not been assessed. To compare animal movement in response to these three contexts, we translocated three species of murid rodents (meadow voles, Microtus pennsylvanicus, deer mice, Peromyscus maniculatus, and red-backed voles, Clethrionomys gapperi) across the TCH (an artificial barrier), a forested strip in the median of the TCH (a natural barrier), in the adjacent highway verge (continuous grassy) and in nearby forest (continuous forested). We coated murids with fluorescent dye, released them at standardized distances, and followed the resulting trails to obtain detailed information on movement paths. Overall, individuals were 20% less successful crossing the TCH than natural (forested) barriers and 10% less successful crossing natural barriers than continuous habitat, but exhibited marked variation among species. Path tortuosity was negatively related to return success; more convoluted paths were associated with lower success across all treatments. Deer mice returned to their home-ranges more often than the other two species, perhaps because of their generalist habitat preferences, nocturnal activities, or larger home ranges. In all treatments, the fractal dimensions of movement paths were very qualitatively similar to those observed for return success and tortuosity, indicating that the responses of these species were not influenced by spatial scale. Together, our results suggest that (a) murid species differ in the movement limitation imposed by both natural and artificial barriers, perhaps as a consequence of differences in habitat preferences and activity patterns, and (b) murid responses to a road barrier are only moderately different from responses to the natural barrier of a forest edge in the context of translocation experiments.

In small mammal populations with multiannual oscillations in density, observational data have revealed cyclic changes in population structure, reproduction, and individual quality, but mechanisms inducing these changes have remained an open question. We analysed data collected during a 3-year predator reduction experiment to find out the effects of predators on population structure, reproductive parameters, and individual quality of Microtus voles (the field vole M. agrestis and the sibling vole M. rossiaemeridionalis) in western Finland. Voles were collected by snap trapping in April, June, August, and October during 1997-1999. The yearly reduction of predators from April to October had a clear positive effect on the abundance of sibling voles but did not significantly affect the densities of field voles. Predator reduction apparently also affected the age ratio and mean body size in late summer, as well as pancreatic weights of voles. However, all observed differences between predator reduction and control areas, except those in abundance, were small and may mainly reflect a generally higher survival leading to higher densities of voles in predator reduction areas. Our results also indicated a relative lack of high quality food at population peaks but not because of reduced foraging activity in the presence of predators. We conclude that the indirect effects of vole-eating predators on the population growth of main prey are small compared to the detrimental direct effects on prey survival. In the case of less preferred prey, indirect effects of predation through reduced interspecific competition may play a role at high densities.

Predation impacts by introduced predators are predicted to be most intense in island ecosystems, and also variable depending on environmental conditions, but large-scale experimental field testing is rare. In this study we examine the factors that determine the distribution and abundance of vole metapopulations preyed upon by feral American mink Mustela vison in the outer Finnish archipelago of the Baltic Sea. Specifically, we follow the dynamics of field voles Microtus agrestis and bank voles Clethrionomys glareolus on 40 small islands under variable rainfall as part of a large-scale mink removal experiment. For both vole species occupancy rates were negatively influenced by island isolation, as were extinction events for field voles. High summer rainfall in 1998 corresponded to large vole populations where mink were absent, populations that then crashed in 1999 and 2000 when below average rains fell during the summer breeding season. Where mink were present however, vole abundance remained more constant between years with no boom-bust apparent. We conclude that weather and predation may drive vole abundance whereas habitat patchiness and metapopulation processes more strongly drive vole distributions. There may also be potential for interaction between these factors: because feral mink prevent rapid vole population growth after good summer rains, and vole dispersal is influenced by population size, feral mink may be changing vole dispersal patterns to disrupt the natural metapopulation dynamic. Hence this indirect impact of mink could lead to gradual erosion of vole populations in the outer archipelago by reducing recolonisation processes.

Habitat fragmentation is a common feature of modern landscapes, with significant impacts on the population densities of and space use by animals. A frequent model system for studying these effects is that of voles (Microtus spp.) and other rodents in experimentally fragmented grasslands. Design of these study sites is motivated by landscape theory, and often assumes that the matrix between discrete experimental habitat patches is inhospitable to focal species. Our census of M. ochrogaster runways in an experimentally fragmented landscape in eastern Kansas indicates that voles make substantial use of the mowed matrix, and that such use increases with degree of habitat fragmentation. Census season (winter, summer) was also an important predictor of observed runway distributions. Our results suggest that voles may have home ranges that include multiple small fragments and also the intervening matrix, leading to overestimates of population density. While the assumption that the matrix is inhospitable in experimentally fragmented habitats may be justified in many cases, studies using the model system of Microtus spp. or ecologically similar rodents in grasslands should incorporate methods that account for use of the matrix.

We studied the predation rate and prey selection of the least weasel (Mustela nivalis nivalis) on its two most common prey species in boreal environments, the bank vole (Clethrionomys glareolus) and the field vole (Microtus agrestis), in large outdoor enclosures. We also studied the response of weasels to odours of the two species in the laboratory. The enclosure experiment was conducted using constant vole densities (16 voles/ha) but with varying relative abundance of the two species. Weasels showed higher predation rates on bank voles, and males had higher predation rate than females. Females killed disproportionately more of the more abundant prey species, but they preferred bank voles to field voles when both were equally available. Overall, the predation rate also increased with increasing abundance of bank voles. Therefore our results are in agreement with earlier laboratory results showing preference for bank voles, even if no intrinsic preference for odours of either species was observed in our laboratory study. We suggest that the least weasel hunts according to prey availability, prey aggregation and suitability of hunting habitat, and that this causes the observed dependence of least weasels on field voles and emphasises the role of the field vole in the vole-weasel interaction in cyclic vole populations. Furthermore, our results suggest that predation by weasels may facilitate the coexistence of the two vole species via predator switching, and that it may cause the observed synchrony in dynamics between vole species.

Meadow voles, Microtus pennsylvanicus, affect the species composition, distribution, and succession of plants in grassland ecosystems, but the effects of voles on herbaceous plants when grasslands are bordered by wooded edges is not known. We investigated the impact of wooded edges on vole distribution and herbivory of relatively palatable and unpalatable native prairie plant species by studying five reconstructed tallgrass prairies with wooded edges in central Iowa. A 50 x 50 m trapping grid at each site was established to determine the proportion of voles captured at various distances from the edge. We found that meadow voles were less abundant at wooded edges and, in general, increased in number toward the prairie interior. Seedlings of purple prairie clover (Dalea purpurea) and Illinois bundleflower (Desmanthus illinoensis), a relatively palatable and unpalatable species, respectively, were transplanted onto simulated gopher mounds 2, 5, 10, 20 and 30 m from the edge. The number of plants grazed per species per mound was determined 1 week and 4 weeks after planting. The amount of herbivory on both species was significantly different by distance, with fewer plants eaten 2 m from the edge. Interestingly, the amount of herbivory on relatively unpalatable plants did not differ from more palatable plants. Herbivory on both plant species also varied by site, such that sites with lower vole density tended to have lower amounts of herbivory. These results indicate that wooded edges do have an effect on meadow vole distribution and native prairie seedling herbivory. Because voles avoid wooded edges, seedlings of any species may experience a small refuge from herbivory along wooded edges.

Studies on competing mammalian species in the past have focused mainly on the competitive exclusion of one species from the preferred habitat of the other. Investigations on effects of competition and coexistence on individual fitness are rare. In this study we were able to measure effects of interspecific competition on major fitness components, using a system with two vole species in asymmetric competition. Survival, reproduction and space use of bank vole Clethrionomys glareolus females were monitored in 32 enclosed populations over four replicates of eight parallel run enclosures. Into half of the enclosures we introduced an additional number of field voles Microtus agrestis, a dominant competitor. Survival of bank vole females was lower under competitive conditions. Total number of breeding females was lower in populations coexisting with competitors. Territory size of bank vole females decreased. Females body weight and litter size bank vole litters conceived during the experiment were not affected by interspecific competition. These characteristics should respond to differences in food resources, and territory size should increase if food was scarce, thus we found no indication of direct exploitation competition between the two species. Space use was overlapping between the species, but individuals of both species were never caught together in the same trap, indicating avoidance behaviour. We conclude that adult bank vole females do suffer fitness consequences through interference competition with field voles, probably basing on increased number of aggressive encounters in the presence of the dominant species. Our results suggest, that direct interference rather than indirect exploitation competition may be the cause for observed fitness decrease in bank vole females.

Coexisting prey species interact indirectly via their shared predators when one prey type influences predation rates of the second prey type. In a temperate system where the predominant shared predator is a generalist, I studied the indirect effects of rodent populations on waterfowl nest success, both within the nesting season among sites and among years. Among six to ten upland fields (14 to 27 ha), mallard (Anas platyrhynchos) nest success was positively correlated with rodent abundance in all three years of the study. After removing year effects, mallard nest success remained positively correlated with the relative abundance of rodents. Of the rodent species present, California voles (Microtus californicus) were the most important coexisting prey type influencing nest success. Among years, mallard nest success was positively correlated with vole abundance; the asymptotic relationship suggests a threshold response to vole abundance, beyond which predators become satiated and additional voles do little to affect nest success. I tested and rejected three alternative explanations for the observed positive correlation between mallard nest success and rodent abundance that do not involve an indirect effect of coexisting prey populations. The influences of dense nesting cover, nesting density, and predator activity did not explain the observed patterns of nest success. These results suggest that rodent populations buffer predation on waterfowl nests, both within and among years, via the behavioral responses of shared predators to coexisting prey.

Within Fennoscandia, two well-studied groups of herbivores exhibit clear geographical gradients in their population dynamics. Populations of a forest lepidopteran (Epirrita autumnata, the autumnal moth) and voles of the genera Microtus and Clethrionomys show pronounced multi-annual cycles in the north but become more stable towards the south. Here we review empirical and theoretical studies on these species, mainly regarding the biological mechanisms that are assumed to generate the pattern of population dynamics in both systems. We conclude that the specialist/generalist predation hypothesis offers a common explanation for the population cycles and their geographical gradients irrespective of whether a herbivorous insect or small mammals are concerned. According to this hypothesis, originally developed for the Fennoscandian voles, but now applied also to E. autumnata, population cycles are generated by specialist natural enemies (predators for the voles and parasitoids for E. autumnata). Furthermore, the dynamic shift from cycles to stability is assumed to be caused by an increase in the density and diversity of generalist natural enemies from north to south in Fennoscandia.

Factors involved in causing cyclic vole populations to decline, and in preventing populations from recovering during the subsequent low density phase have long remained unidentified. The traditional view of self-regulation assumes that an increase in population density is prevented by a change in the quality of individuals within the population itself, but this is still inadequately tested in the field. We compared the population growth of wild field voles (Microtus agrestis) from the low phase (conducted in 1998) with that of voles from the increase phase (conducted in 1999) in predator-proof enclosures (each 0.5 ha) in western Finland. Within a few months, enclosed vole populations increased to high density, and the realised per capita rate of change over the breeding season did not differ between the populations from different cycle phases. This implies that the recovery of populations from the low phase was not hindered by an impoverishment in quality of individual voles. Accordingly, we suggest that population intrinsic factors (irrespective of the mechanisms they are based on) are unlikely to play a significant role in the generation of cyclic density fluctuations of voles. Instead, we discovered direct density-dependent regulation in the vole populations. Accurate estimates of population growth and the observed density dependence provide important information for empirically based models on population dynamics of rodents.

Cyclic changes in population growth rate are caused by changes in survival and/or reproductive rate. To find out whether cyclic changes in reproduction are an important part of the mechanism causing cyclic fluctuations in small mammal populations, we studied changes in the population structure and reproduction of field voles (Microtus agrestis), sibling voles (M. rossiaemeridionalis), bank voles (Clethrionomys glareolus), and common shrews (Sorex araneus) in western Finland during 1984-1992, in an area with 3-yr vole cycles. We also modelled the population growth of voles using parameter values from this study. The animals studied were collected by snap trapping in April, May, June, August, September, and, during 1986-1990, also in October. We found several phase-related differences in the population structure (age structure, sex ratio, proportion of mature individuals) and reproduction (litter size, length of the breeding season) of voles. In non-cyclic common shrews, the only significant phase-related difference was a lower proportion of overwintered individuals in the increase phase. According to the analyses and the vole model, phase-related changes in litter size had only a minor impact on population growth rate. The same was true for winter breeding in the increase phase. The length and intensity of the summer breeding season had an effect on yearly population growth but this impact was relatively weak compared to the effect of cyclic changes in survival. The population increase rates of Microtus were delayed dependent on density (8-12-month time lag). Our results indicate that cyclic changes in reproduction are not an important part of the mechanism driving cyclic fluctuations in vole populations. Low survival of young individuals appeared to play an important role in the shift from the peak to the decline phase in late summer and early autumn.

Reproductive traits of voles vary with the phases of the population density fluctuations. We sought to determine whether the source of this variation resides in the individuals or in their environment. Overwintering field voles (Microtus agrestis) from two cyclic out-of-phase populations (increase and peak phases) were sampled in early spring and bred in the laboratory for two generations under standardised conditions with ambient light and temperature. Monitoring of the source populations by capture-mark-recapture showed large differences in reproductive performance. In the increase area, reproduction started six weeks earlier, the probability of maturation of young-of-the-year was more than ten times higher during mid-summer, and reproduction continued nearly two months later in the autumn than in the peak area. These differences were not found to be associated with a difference in age structure of overwintered animals between the two areas (assessed by the distribution of eye lens masses from autopsy samples). Although the population differences in reproductive traits were to some degree also present among the overwintered animals in the laboratory, we found no difference in reproductive traits in the laboratory-born generations. There was a strongly declining seasonal trend in probability of sexual maturation both in the field and in the laboratory under ambient light conditions. However, in the field there were large population differences in the steepness of the seasonal decline that were not seen under the standardised laboratory conditions. We conclude that seasonal decline in maturation rates is governed by change in photoperiod, but that the population level variation in the shape of the decline is caused by a direct response to the environment and not due to variation in any intrinsic state of the individuals reflecting the environment experienced by the previous generation(s).

Corridors have become a key issue in the discussion of conservation planning; however, few empirical data exist on the use of corridors and their effects on population dynamics. The objective of this replicated, population level, capture-recapture experiment on meadow voles was to estimate and compare population characteristics of voles between (1) corridor-linked fragments, (2) isolated or non-linked fragments, and (3) unfragmented areas. We conducted two field experiments involving 22 600 captures of 5700 individuals. In the first, the maintained corridor study, corridors were maintained at the time of fragmentation, and in the second, the constructed corridor study, we constructed corridors between patches that had been fragmented for some period of time. We applied multistate capture-recapture models with the robust design to estimate adult movement and survival rates, population size, temporal variation in population size, recruitment, and juvenile survival rates. Movement rates increased to a greater extent on constructed corridor-linked grids than on the unfragmented or non-linked fragmented grids between the pre- and post-treatment periods. We found significant differences in local survival on the treated (corridor-linked) grids compared to survival on the fragmented and unfragmented grids between the pre- and post-treatment periods. We found no clear pattern of treatment effects on population size or recruitment in either study. However, in both studies, we found that unfragmented grids were more stable than the fragmented grids based on lower temporal variability in population size. To our knowledge, this is the first experimental study demonstrating that corridors constructed between existing fragmented populations can indeed cause increases in movement and associated changes in demography, supporting the use of constructed corridors for this purpose in conservation biology.

Spatial autocorrelation is ubiquitous in ecological systems. However, consideration of this phenomenon in seed and seedling predation studies has been limited, perhaps explaining some of the variation observed between studies and the difficulty in generalizing about the impacts of herbivores on successional processes. I examined the prevalence and patterns of spatial autocorrelation in seed and seedling survival between neighboring tree propagules in old-fields. All experiments were conducted within enclosures in which I manipulated the density of meadow voles (Microtus pennsylvanicus). Voles, and competitively inferior white-footed mice (Peromyscus leucopus), are the dominant consumers of tree seedlings and seeds in old-fields in the northeastern United States, respectively. An intermediate level of spatial autocorrelation was present in seed and seedling predation trials. Furthermore, the frequency of spatial autocorrelation increased under low-vole density conditions and in study sites with greater heterogeneity in vegetation cover. Survival of neighboring tree propagules was generally positively autocorrelated, suggesting that foraging by mice and voles contributes to a clumped pattern of tree propagule establishment in old-fields. A review of published predation studies found that only 8.3% attempted to test for spatial autocorrelation and that the minimum average spacing used between replicates is well below the average autocorrelation distance of 8.1 ± 6.3 m observed in the current study, Patterns of autocorrelation appear somewhat predictable based on knowledge of the ecology of mice and voles and to provide insights into the factors influencing their foraging behavior and potential impacts on old-field succession.

We used depletable food patches to determine the effect of microhabitat (mowed versus unmowed adjacent grasslands) and time (day versus night) on the foraging behavior of common voles (Microtus arvalis). The food remaining after 12-h periods (giving-up density, GUD) measured the vole's habitat selection under predation risk. In accord with several other rodent species and the effects of avian predators, voles had significantly lower GUDs in the unmowed than mowed portion of the grassland. GUDs in patches along the border between adjacent habitats were more similar to the risky mowed grassland than the safe unmowed grass. Time interacted strongly with microhabitat. In the mowed grass, voles had significantly higher GUDs at night than day. Whereas in the unmowed grass, GUDs were significantly higher during the day than night. Vole GUDs did not vary with time along the boundary. This suggests that predators are more abundant or effective in the mowed grass at night (owls?), and in the unmowed grass during the day (weasels?). In terms of predation risk, the voles perceived the mowed grass at night as the riskiest and the unmowed grass at night as the safest. Voles may have difficulties assessing resources under high predation risk: GUDs among patches were well equalized in the unmowed microhabitat whereas in the mowed grass only day GUDs did not vary significantly among patches. We linked these results to the vole's day-night-activity and life span. For the 533 voles live-trapped at the study area, the ratio of day versus night captures for each individual served as an activity index and the span between first and last capture measured minimum life span. In accord with higher GUDs at night, very few individuals behaved selectively towards the night, but individual life expectancy increased with temporal opportunism. Microhabitat differences in GUDs reflect short-term strategies of predator avoidance and the trapping data reflect long-term patterns of anti-predator behavior.

Theoretical models predict that delayed density-dependent processes with a time-lag of approximately nine months are sufficient to generate regular 3-5 year fluctuations in densities of northern small rodents. To examine whether this time-lag could be generated by plant-herbivore interactions, we studied delayed effects of overgrazed food plants on voles. We introduced field voles (Microtus agrestis) in four large predator-proof enclosures that had suffered heavy grazing during the preceding autumn and winter, and compared them with voles introduced to previously ungrazed control areas. We found no detrimental effects of previous grazing on population growth, reproduction or body condition of voles. Chemical analyses did not show consistent effects of grazing on nutritional components of common food plants (grasses). These results suggest that short-term population cycles of Microtus voles in grassland habitats are not primarily driven by delayed effects of plant-herbivore interactions.

We investigated the functional response of the least weasel (Mustela nivalis nivalis) in a series of experiments conducted in large outdoor enclosures (0.5 ha). Radiocollared Microtus voles were released in the enclosures at different densities (4, 8, 16 and 100/ha) three days before the release of a radiocollared weasel. During the three-day experiment every vole killed was replaced with another one as soon as possible to retain constant prey density. The results demonstrated type II functional response with the predation rate reaching 50% of the asymptotic rate at a vole density of 15 individuals per ha. More voles were killed at the highest densities than would be expected from the known energy demands of weasels. Female and male voles were killed in proportion to their abundances in the enclosures, and no difference in predation risk was detected between voles released in the enclosure before the weasel (residents) and during the experiment (transients).

The meadow vole, Microtus pennsylvanicus, is the most widely distributed Microtus species in North America. Across its range, it shows marked demographic differences, experiences a large range of climatic conditions, and varies considerably in body size and life-history characteristics. To study the genetic basis of the geographic variation in size and life history of this species, we subjected three populations, one from central Canada and two from eastern Canada, to quantitative genetic analysis in the lab. We studied the variance and covariance of several size and growth variables as well as age and size at maturity by means of population crosses, full-sib analysis, and parent-offspring regressions. We found that the phenotypic differences among these populations are almost entirely due to environmental effects. However, within populations, additive genetic and maternal effects explain most of the variation. We discuss possible explanations for the lack of genetic differences among the populations and speculate that a similar reaction norm is maintained in all populations through heterogeneity in the temporal or spatial environment that the populations experience. The heterogeneity may be mediated through population density fluctuations, climatic variation, or variation in site productivity. Thus, we hypothesize that M. pennsylvanicus has evolved to be the best in all possible worlds rather than in one actual world. This study highlights the crucial importance of maternal and environmental effects on the size, growth, and life history of small rodents.

Population densities, food and predators were examined for bank voles, Clethrionomys glareolu and field voles, Microtus agrestis, during the period 1971-1998 in one northern (Stromsund) and one south-central (Uppsala) region in Sweden to see if trophic relationships persisted in the presence of large changes in landscape composition, and rodent dynamics, in the northern region. The main work was done on early reforestations. Whereas the limited forest areas at Uppsala were consistently situated on poor soil, forests were extensively clearcut at Stromsund in the 1960-1980s, continuously leaving poorer and fragmented old forests and extensive young thicket forests that were of little use to small rodents. Population densities and amplitudes were larger at Stromsund in the 1970-1980s than in the 1990s and densities at Stromsund reached higher peaks than at Uppsala. The proportion of M. agrestis in the rodent communities was consistently larger at Stromsund. Densities of C. glareolus were higher in forests than on reforestations but there was a continuous exchange of individuals between the two habitats. M. agrestis on abandoned fields, a main habitat for this species, did not show any temporal or regional variation and were more common than on reforestations, except at Stromsund in the 1970-1980s. Consumption of artificial seed supplies did not differ annually at Uppsala but did so at Stromsund in 1994-96; almost all supplies were there consumed at a vole peak year, particularly inside forests. Consumption of an artificial bark supply on reforestations was much larger in the 1980s than in the 1990s at Stromsund; it was consistently low at Uppsala. There was not any difference between regions in such bark consumption on abandoned fields. Vole damage to forestry seedlings was important in the 1960-1970s, of less importance in the 1980s, and did not occur at all in the surveyed years of the 1990s. Mustelid numbers (particularly Mustela erminea) were high at Stromsund after vole peaks in the 1980s but were generally low in the 1990s; they were consistently low at Uppsala. Fox tracks did not vary in any consistent way. Following interpretation emerged: Sheltering snow cover generated cycles. Peak C. glareolus numbers were generally limited by winter food, particularly by pendulous lichens that decreased during the study period. Abundant seed or fruit supply in increase autumns can irregularly cause particularly high C. glareolus peaks. Massive invasion of northern reforestations by C. glareolus from forests appears to have buffered predation on M. agrestis, permitting extensive cycles of the latter species on reforestations in the 1960-1980s. Limited numbers of C. glareolus on reforestations permitted only small M. agrestis populations there and generalist predators as foxes or raptors may then have terminated minor increases already in summer. Trophic interactions are intensified and spatially extended at pronounced peak numbers. More generally, regional changes in fluctuation patterns of small rodents can be related to varying use of managed landscapes, where habitat quality changes both in time and space.

Presence and stability of populations of Microtus ochrogaster, M. pennsylvanicus and Blarina brevicauda were studied in tallgrass prairie habitat in central Illinois for 25 yr: 52 yr of autumn data were available for M. ochrogaster. M. ochrogaster displayed erratic, very low amplitude population fluctuations in tallgrass; it was absent from the study site 49% of the time. Multiannual cycles were not obvious. Population fluctuations in adjacent alfalfa and bluegrass sites were higher, more frequent and in synchrony with those in tallgrass. Densities recorded in the autumn from 1944 to 1996 revealed distinct, erratic multiannual population cycles. M. pennsylvanicus populations were continuously present at relatively high densities in tallgrass; neither annual nor multiannual fluctuations were evident. Infrequent annual cycles were recorded in the other two habitats. B. brevicauda was persistent and displayed a synchronous annual cycle in the three habitats.

We determined life expectancy (to within approximately 3 d) of 1457 prairie voles. Microtus ochrogaster, in a free-living population over a period of 6 yr. During this time the population underwent four cycles of abundance, with peak densities of 160, 250, 350, and 624 individuals per ha. Life expectancy was calculated based on the interactions of season, phase of the population cycle, population density, type of natal social group, and weather conditions 21 and 30 d before and after birth. Life expectancy was greatest for animals born (1) in autumn, (2) during the increase phase of the population cycle, (3) at population densities >100/ha, and (4) into communal groups. The combination of these factors appears to result in a surge in population density in late autumn. The short life expectancy of animals born during the peak and decline phases of the population cycle may accelerate the rate of the decline phase. During periods of high temperatures before birth in summer, life expectancy increased with increased precipitation. There was no such interaction of precipitation and temperature after birth. In both summer and autumn, high precipitation after birth resulted in lower life expectancy, possibly reflecting increased mortality of young forced to leave flooded underground nests.

We studied the possible role of food quality in cyclic vole populations by using the size of internal organs as an indicator of forage quality. We trapped field (Microtus agrestis) and sibling (M. rossiaemeridionalis) voles during two increase and two decline phases of the 3-yr population cycle in western Finland, recorded total body mass and weighed their pancreas and liver. There were no obvious differences in the relative size of organs related to phase of the cycle, which suggests voles from declining populations do not exhibit pancreatic hypertrophy. These results do not support the hypothesis that proteinase inhibitors in food plants that have adverse effects on metabolic processes of herbivores, and which are induced by grazing, cause regular cycles in small rodent populations.

We developed an empirically based model of density-dependent vole population growth based on experimental data on population dynamics of Microtus pennsylvanicus in large field enclosures. Statistical analysis of the data indicated that both density dependent regulation and seasonal effects were important in influencing vole population growth. Together, these two factors explained approximately one-half of variance in the realized per capita rate of change exhibited by experimental vole populations. A population model assuming simple functional forms (linear for population density and sine for seasonality) provided an adequate description of the data, with more complex functional forms leading to at best minimal improvements. The natural rate of population increase, averaged over all seasons, was estimated as (mean ± SE) r max = 6.0(±0.4) yr -1 . This estimate suggests an impressive power of population increase, implying that each female vole could be replaced by about 400 daughters a year later (assuming density-independent growth). A survey of literature, however, indicates that this is by no means the largest rate of increase observed in a vole population.