The timid invasion: behavioural adjustments and range expansion in a non-native rodent.

Animal behaviour can moderate biological invasion processes, and the native fauna's ability to adapt. The importance and nature of behavioural traits favouring colonization success remain debated. We investigated behavioural responses associated with risk-taking and exploration, both in non-native bank voles (Myodes glareolus, N = 225) accidentally introduced to Ireland a century ago, and in native wood mice (Apodemus sylvaticus, N = 189), that decline in numbers with vole expansion. We repeatedly sampled behavioural responses in three colonization zones: established bank vole populations for greater than 80 years (2 sites), expansion edge vole populations present for 1-4 years (4) and pre-arrival (2). All zones were occupied by wood mice. Individuals of both species varied consistently in risk-taking and exploration. Mice had not adjusted their behaviour to the presence of non-native voles, as it did not differ between the zones. Male voles at the expansion edge were initially more risk-averse but habituated faster to repeated testing, compared to voles in the established population. Results thus indicate spatial sorting for risk-taking propensity along the expansion edge in the dispersing sex. In non-native prey species the ability to develop risk-averse phenotypes may thus represent a fundamental component for range expansions.

Top-down effects of foraging decisions on local, landscape and regional biodiversity of resources (DivGUD).

Foraging by consumers acts as a biotic filtering mechanism for biodiversity at the trophic level of resources. Variation in foraging behaviour has cascading effects on abundance, diversity, and functional trait composition of the community of resource species. Here we propose diversity at giving-up density (DivGUD), i.e. when foragers quit exploiting a patch, as a novel concept and simple measure quantifying cascading effects at multiple spatial scales. In experimental landscapes with an assemblage of plant seeds, patch residency of wild rodents decreased local α-DivGUD (via elevated mortality of species with large seeds) and regional γ-DivGUD, while dissimilarity among patches in a landscape (ß-DivGUD) increased. By linking theories of adaptive foraging behaviour with community ecology, DivGUD allows to investigate cascading indirect predation effects, e.g. the ecology-of-fear framework, feedbacks between functional trait composition of resource species and consumer communities, and effects of inter-individual differences among foragers on the biodiversity of resource communities.

Serologic Surveillance for SARS-CoV-2 Infection among Wild Rodents, Europe.

We report results from serologic surveillance for exposure to SARS-CoV-2 among 1,237 wild rodents and small mammals across Europe. All samples were negative, with the possible exception of 1. Despite suspected potential for human-to-rodent spillover, no evidence of widespread SARS-CoV-2 circulation in rodent populations has been reported to date.Esitämme tulokset serologisesta tutkimuksesta, jossa seulottiin SARS-CoV-2 tartuntojen varalta 1,237 luonnonvaraista jyrsijää ja piennisäkästä eri puolilta Eurooppaa. Kaikki näytteet olivat negatiivisia, yhtä näytettä lukuun ottamatta. SARS-CoV-2:n läikkymisen ihmisistä jyrsijöihin on arveltu olevan mahdollista, mutta todisteet viruksen laajamittaisesta leviämisestä jyrsijäpopulaatioissa puuttuvat.

My niche: individual spatial niche specialization affects within- and between-species interactions.

Intraspecific trait variation is an important determinant of fundamental ecological interactions. Many of these interactions are mediated by behaviour. Therefore, interindividual differences in behaviour should contribute to individual niche specialization. Comparable with variation in morphological traits, behavioural differentiation between individuals should limit similarity among competitors and thus act as a mechanism maintaining within-species variation in ecological niches and facilitating species coexistence. Here, we aimed to test whether interindividual differences in boldness covary with spatial interactions within and between two ecologically similar, co-occurring rodent species (Myodes glareolus, Apodemus agrarius). In five subpopulations in northeast Germany, we quantified individual differences in boldness via repeated standardized tests and spatial interaction patterns via capture-mark-recapture (n = 126) and automated VHF telemetry (n = 36). We found that boldness varied with space use in both species. Individuals of the same population occupied different spatial niches, which resulted in non-random patterns of within- and between-species spatial interactions. Behavioural types mainly differed in the relative importance of intra- versus interspecific competition. Within-species variation along this competition gradient could contribute to maintaining individual niche specialization. Moreover, behavioural differentiation between individuals limits similarity among competitors, which might facilitate the coexistence of functionally equivalent species and, thus, affect community dynamics and local biodiversity.

Small mammals in the big city: Behavioural adjustments of non-commensal rodents to urban environments.

A fundamental focus of current ecological and evolutionary research is to illuminate the drivers of animals' success in coping with human-induced rapid environmental change (HIREC). Behavioural adaptations are likely to play a major role in coping with HIREC because behaviour largely determines how individuals interact with their surroundings. A substantial body of research reports behavioural modifications in urban dwellers compared to rural conspecifics. However, it is often unknown whether the observed phenotypic divergence is due to phenotypic plasticity or the product of genetic adaptations. Here, we aimed at investigating (a) whether behavioural differences arise also between rural and urban populations of non-commensal rodents; and (b) whether these differences result from behavioural flexibility or from intrinsic behavioural characteristics, such as genetic or maternal effects. We captured and kept under common environment conditions 42 rural and 52 urban adult common voles (Microtus arvalis) from seven subpopulations along a rural-urban gradient. We investigated individual variation in behavioural responses associated with risk-taking and exploration, in situ at the time of capture in the field and ex situ after 3 months in captivity. Urban dwellers were bolder and more explorative than rural conspecifics at the time of capture in their respective sites (in situ). However, when tested under common environmental conditions ex situ, rural individuals showed little change in their behavioural responses whereas urban individuals altered their behaviour considerably and were consistently shyer and less explorative than when tested in situ. The combination of elevated risk-taking and exploration with high behavioural flexibility might allow urban populations to successfully cope with the challenges of HIREC. Investigating whether the observed differences in behavioural flexibility are adaptive and how they are shaped by additive and interactive effects of genetic make-up and past environmental conditions will help illuminate eco-evolutionary dynamics under HIREC and predict persistence of populations under urban conditions.

Among-individual differences in foraging modulate resource exploitation under perceived predation risk.

Foraging is risky and involves balancing the benefits of resource acquisition with costs of predation. Optimal foraging theory predicts where, when and how long to forage in a given spatiotemporal distribution of risks and resources. However, significant variation in foraging behaviour and resource exploitation remain unexplained. Using single foragers in artificial landscapes of perceived risks and resources with diminishing returns, we aimed to test whether foraging behaviour and resource exploitation are adjusted to risk level, vary with risk during different components of foraging, and (co)vary among individuals. We quantified foraging behaviour and resource exploitation for 21 common voles (Microtus arvalis). By manipulating ground cover, we created simple landscapes of two food patches varying in perceived risk during feeding in a patch and/or while travelling between patches. Foraging of individuals was variable and adjusted to risk level and type. High risk during feeding reduced feeding duration and food consumption more strongly than risk while travelling. Risk during travelling modified the risk effects of feeding for changes between patches and resulting evenness of resource exploitation. Across risk conditions individuals differed consistently in when and how long they exploited resources and exposed themselves to risk. These among-individual differences in foraging behaviour were associated with consistent patterns of resource exploitation. Thus, different strategies in foraging-under-risk ultimately lead to unequal payoffs and might affect lower trophic levels in food webs. Inter-individual differences in foraging behaviour, i.e. foraging personalities, are an integral part of foraging behaviour and need to be fully integrated into optimal foraging theory.

Individuals in space: personality-dependent space use, movement and microhabitat use facilitate individual spatial niche specialization.

Personality-dependent space use and movement might be crucially influencing ecological interactions, giving way to individual niche specialization. This new approach challenges classical niche theory with potentially great ecological consequences, but so far has only scarce empirical support. Here, we investigated if and how consistent inter-individual differences in behavior predict space use and movement patterns in free-ranging bank voles (Myodes glareolus) and thereby contribute to individual niche specialization. Individuals were captured and marked from three different subpopulations in North-East Germany. Inter-individual differences in boldness and exploration were quantified via repeated standardized tests directly in the field after capture. Subsequently, space use and movement patterns of a representative sample of the behavioral variation (n = 21 individuals) were monitored via automated VHF telemetry for a period of four days, yielding on average 384 locations per individual. Bolder individuals occupied larger home ranges and core areas (estimated via kernel density analyses), moved longer distances, spatially overlapped with fewer conspecifics and preferred different microhabitats based on vegetation cover compared to shyer individuals. We found evidence for personality-dependent space use, movement, and occupation of individual spatial niches in bank voles. Thus, besides dietary niche specialization also spatial dimensions of ecological niches vary among individuals within populations, which may have important consequences for ecological interactions within- and between species.

The Bruce effect revisited: is pregnancy termination in female rodents an adaptation to ensure breeding success after male turnover in low densities?

Pregnancy termination after encountering a strange male, the Bruce effect, is regarded as a counterstrategy of female mammals towards anticipated infanticide. While confirmed in caged rodent pairs, no verification for the Bruce effect existed from experimental field populations of small rodents. We suggest that the effect may be adaptive for breeding rodent females only under specific conditions related to populations with cyclically fluctuating densities. We investigated the occurrence of delay in birth date after experimental turnover of the breeding male under different population composition in bank voles (Myodes glareolus) in large outdoor enclosures: one-male-multiple-females (n = 6 populations/18 females), multiple-males-multiple-females (n = 15/45), and single-male-single-female (MF treatment, n = 74/74). Most delays were observed in the MF treatment after turnover. Parallel we showed in a laboratory experiment (n = 205 females) that overwintered and primiparous females, the most abundant cohort during population lows in the increase phase of cyclic rodent populations, were more likely to delay births after turnover of the male than year-born and multiparous females. Taken together, our results suggest that the Bruce effect may be an adaptive breeding strategy for rodent females in cyclic populations specifically at low densities in the increase phase, when isolated, overwintered animals associate in MF pairs. During population lows infanticide risk and inbreeding risk may then be higher than during population highs, while also the fitness value of a litter in an increasing population is higher. Therefore, the Bruce effect may be adaptive for females during annual population lows in the increase phases, even at the costs of delaying reproduction.

Puumala hantavirus infections in bank vole populations: host and virus dynamics in Central Europe.

BACKGROUND: In Europe, bank voles (Myodes glareolus) are widely distributed and can transmit Puumala virus (PUUV) to humans, which causes a mild to moderate form of haemorrhagic fever with renal syndrome, called nephropathia epidemica. Uncovering the link between host and virus dynamics can help to prevent human PUUV infections in the future. Bank voles were live trapped three times a year in 2010-2013 in three woodland plots in each of four regions in Germany. Bank vole population density was estimated and blood samples collected to detect PUUV specific antibodies. RESULTS: We demonstrated that fluctuation of PUUV seroprevalence is dependent not only on multi-annual but also on seasonal dynamics of rodent host abundance. Moreover, PUUV infection might affect host fitness, because seropositive individuals survived better from spring to summer than uninfected bank voles. Individual space use was independent of PUUV infections. CONCLUSIONS: Our study provides robust estimations of relevant patterns and processes of the dynamics of PUUV and its rodent host in Central Europe, which are highly important for the future development of predictive models for human hantavirus infection risk.

Evidence for selection maintaining MHC diversity in a rodent species despite strong density fluctuations.

Strong spatiotemporal variation in population size often leads to reduced genetic diversity limiting the adaptive potential of individual populations. Key genes of adaptive variation are encoded by the immune genes of the major histocompatibility complex (MHC) playing an essential role in parasite resistance. How MHC variation persists in rodent populations that regularly experience population bottlenecks remains an important topic in evolutionary genetics. We analysed the consequences of strong population fluctuations on MHC class II DRB exon 2 diversity in two distant common vole (Microtus arvalis) populations in three consecutive years using a high-throughput sequencing approach. In 143 individuals, we detected 25 nucleotide alleles translating into 14 unique amino acid MHC alleles belonging to at least three loci. Thus, the overall allelic diversity and amino acid distance among the remaining MHC alleles, used as a surrogate for the range of pathogenic antigens that can be presented to T-cells, are still remarkably high. Both study populations did not show significant population differentiation between years, but significant differences were found between sites. We concluded that selection processes seem to be strong enough to maintain moderate levels of MHC diversity in our study populations outcompeting genetic drift, as the same MHC alleles were conserved between years. Differences in allele frequencies between populations might be the outcome of different local parasite pressures and/or genetic drift. Further understanding of how pathogens vary across space and time will be crucial to further elucidate the mechanisms maintaining MHC diversity in cyclic populations.

High genetic structuring of Tula hantavirus.

Tula virus (TULV) is a vole-associated hantavirus with low or no pathogenicity to humans. In the present study, 686 common voles (Microtus arvalis), 249 field voles (Microtus agrestis) and 30 water voles (Arvicola spec.) were collected at 79 sites in Germany, Luxembourg and France and screened by RT-PCR and TULV-IgG ELISA. TULV-specific RNA and/or antibodies were detected at 43 of the sites, demonstrating a geographically widespread distribution of the virus in the studied area. The TULV prevalence in common voles (16.7 %) was higher than that in field voles (9.2 %) and water voles (10.0 %). Time series data at ten trapping sites showed evidence of a lasting presence of TULV RNA within common vole populations for up to 34 months, although usually at low prevalence. Phylogenetic analysis demonstrated a strong genetic structuring of TULV sequences according to geography and independent of the rodent species, confirming the common vole as the preferential host, with spillover infections to co-occurring field and water voles. TULV phylogenetic clades showed a general association with evolutionary lineages in the common vole as assessed by mitochondrial DNA sequences on a large geographical scale, but with local-scale discrepancies in the contact areas.

How to account for behavioral states in step-selection analysis: a model comparison.

Step-selection models are widely used to study animals' fine-scale habitat selection based on movement data. Resource preferences and movement patterns, however, often depend on the animal's unobserved behavioral states, such as resting or foraging. As this is ignored in standard (integrated) step-selection analyses (SSA, iSSA), different approaches have emerged to account for such states in the analysis. The performance of these approaches and the consequences of ignoring the states in step-selection analysis, however, have rarely been quantified. We evaluate the recent idea of combining iSSAs with hidden Markov models (HMMs), which allows for a joint estimation of the unobserved behavioral states and the associated state-dependent habitat selection. Besides theoretical considerations, we use an extensive simulation study and a case study on fine-scale interactions of simultaneously tracked bank voles (Myodes glareolus) to compare this HMM-iSSA empirically to both the standard and a widely used classification-based iSSA (i.e., a two-step approach based on a separate prior state classification). Moreover, to facilitate its use, we implemented the basic HMM-iSSA approach in the R package HMMiSSA available on GitHub.

Genomic signatures of climate adaptation in bank voles.

Evidence for divergent selection and adaptive variation across the landscape can provide insight into a species' ability to adapt to different environments. However, despite recent advances in genomics, it remains difficult to detect the footprints of climate-mediated selection in natural populations. Here, we analysed ddRAD sequencing data (21,892 SNPs) in conjunction with geographic climate variation to search for signatures of adaptive differentiation in twelve populations of the bank vole (Clethrionomys glareolus) distributed across Europe. To identify the loci subject to selection associated with climate variation, we applied multiple genotype-environment association methods, two univariate and one multivariate, and controlled for the effect of population structure. In total, we identified 213 candidate loci for adaptation, 74 of which were located within genes. In particular, we identified signatures of selection in candidate genes with functions related to lipid metabolism and the immune system. Using the results of redundancy analysis, we demonstrated that population history and climate have joint effects on the genetic variation in the pan-European metapopulation. Furthermore, by examining only candidate loci, we found that annual mean temperature is an important factor shaping adaptive genetic variation in the bank vole. By combining landscape genomic approaches, our study sheds light on genome-wide adaptive differentiation and the spatial distribution of variants underlying adaptive variation influenced by local climate in bank voles.

Decreased external skeletal robustness due to reduced physical activity?

OBJECTIVES: Childhood obesity is a global problem, e.g., due to physical inactivity. External skeletal robustness (Frame-Index) has decreased in German schoolchildren. An association between Frame-Index and physical activity was assumed. Further often body mass index (BMI) is analyzed without reference to bone structure. Therefore, we analyze relationships between Frame-Index, BMI, % body fat, and physical activity. METHODS: In a cross-sectional study, 691 German children aged 6-10 years were investigated. BMI, % body fat, Frame-Index, total steps p.w., sports club rate p.w., training time p.d., and TV-time p.d. were determined. RESULTS: Total steps (P < 0.001), BMI (P < 0.001), and % body fat (P = 0.024) are positively linked to Frame-Index. Total steps (P < 0.001), sports club rate (P = 0.001), and training time (P < 0.001) are negatively associated with % body fat. Total steps (P = 0.017) are negatively linked to BMI. TV-time is positively related to BMI (P < 0.001) and % body fat (P < 0.001). % Body fat is affected by age (P < 0.001), sex (P = 0.028), and total steps (P = 0.002). BMI is influenced by age (P < 0.001), and Frame-Index by sex (P < 0.001) and total steps (P = 0.029). Principal component analysis indicates an association between BMI and TV-time and Frame-Index and total steps. CONCLUSIONS: We demonstrate an association between external skeletal robustness and physical activity, which is not captured by in BMI measurements. Children should be physically active in order to maintain skeletal robustness.

Seasonal variation in the behaviour of a short-lived rodent.

BACKGROUND: Short lived, iteroparous animals in seasonal environments experience variable social and environmental conditions over their lifetime. Animals can be divided into those with a "young-of-the-year" life history (YY, reproducing and dying in the summer of birth) and an "overwinter" life history (OW, overwintering in a subadult state before reproducing next spring).We investigated how behavioural patterns across the population were affected by season and sex, and whether variation in behaviour reflects the variation in life history patterns of each season. Applications of pace-of-life (POL) theory would suggest that long-lived OW animals are shyer in order to increase survival, and YY are bolder in order to increase reproduction. Therefore, we expected that in winter and spring samples, when only OW can be sampled, the animals should be shyer than in summer and autumn, when both OW and YY animals can be sampled.We studied common vole (Microtus arvalis) populations, which express typical, intra-annual density fluctuation. We captured a total of 492 voles at different months over 3 years and examined boldness and activity level with two standardised behavioural experiments. RESULTS: Behavioural variables of the two tests were correlated with each other. Boldness, measured as short latencies in both tests, was extremely high in spring compared to other seasons. Activity level was highest in spring and summer, and higher in males than in females. CONCLUSION: Being bold in laboratory tests may translate into higher risk-taking in nature by being more mobile while seeking out partners or valuable territories. Possible explanations include asset-protection, with OW animals being rather old with low residual reproductive value in spring. Therefore, OW may take higher risks during this season. Offspring born in spring encounter a lower population density and may have higher reproductive value than offspring of later cohorts. A constant connection between life history and animal personality, as suggested by the POL theory, however, was not found. Nevertheless, correlations of traits suggest the existence of animal personalities. In conclusion, complex patterns of population dynamics, seasonal variation in life histories, and variability of behaviour due to asset-protection may cause complex seasonal behavioural dynamics in a population.

Consistency in boldness, activity and exploration at different stages of life.

BACKGROUND: Animals show consistent individual behavioural patterns over time and over situations. This phenomenon has been referred to as animal personality or behavioural syndromes. Little is known about consistency of animal personalities over entire life times. We investigated the repeatability of behaviour in common voles (Microtus arvalis) at different life stages, with different time intervals, and in different situations. Animals were tested using four behavioural tests in three experimental groups: 1. before and after maturation over three months, 2. twice as adults during one week, and 3. twice as adult animals over three months, which resembles a substantial part of their entire adult life span of several months. RESULTS: Different behaviours were correlated within and between tests and a cluster analysis showed three possible behavioural syndrome-axes, which we name boldness, exploration and activity. Activity and exploration behaviour in all tests was highly repeatable in adult animals tested over one week. In animals tested over maturation, exploration behaviour was consistent whereas activity was not. Voles that were tested as adults with a three-month interval showed the opposite pattern with stable activity but unstable exploration behaviour. CONCLUSIONS: The consistency in behaviour over time suggests that common voles do express stable personality over short time. Over longer periods however, behaviour is more flexible and depending on life stage (i.e. tested before/after maturation or as adults) of the tested individual. Level of boldness or activity does not differ between tested groups and maintenance of variation in behavioural traits can therefore not be explained by expected future assets as reported in other studies.

So many choices, so little time: Food preference and movement vary with the landscape of fear.

Spatial and temporal variation in perceived predation risk is an important determinant of movement and foraging activity of animals. Foraging in this landscape of fear, individuals need to decide where and when to move, and what resources to choose. Foraging theory predicts the outcome of these decisions based on energetic trade-offs, but complex interactions between perceived predation risk and preferences of foragers for certain functional traits of their resources are rarely considered. Here, we studied the interactive effects of perceived predation risk on food trait preferences and foraging behavior in bank voles (Myodes glareolus) in experimental landscapes. Individuals (n = 19) were subjected for periods of 24 h to two extreme, risk-uniform landscapes (either risky or safe), containing 25 discrete food patches, filled with seeds of four plant species in even amounts. Seeds varied in functional traits: size, nutrients, and shape. We evaluated whether and how risk modifies forager preference for functional traits. We also investigated whether perceived risk and distance from shelter affected giving-up density (GUD), time in patches, and number of patch visits. In safe landscapes, individuals increased time spent in patches, lowered GUD and visited distant patches more often compared to risky landscapes. Individuals preferred bigger seeds independent of risk, but in the safe treatment they preferred fat-rich over carb-rich seeds. Thus, higher densities of resource levels remained in risky landscapes, while in safe landscapes resource density was lower and less diverse due to selective foraging. Our results suggest that the interaction of perceived risk and dietary preference adds an additional layer to the cascading effects of a landscape of fear which affects biodiversity at resource level.

Forager-mediated cascading effects on food resource species diversity.

Perceived predation risk varies in space and time. Foraging in this landscape of fear alters forager-resource interactions via cascading nonconsumptive effects. Estimating these indirect effects is difficult in natural systems. Here, we applied a novel measure to quantify the diversity at giving-up density that allows to test how spatial variation in perceived predation risk modifies the diversity of multispecies resources at local and regional spatial levels. Furthermore, we evaluated whether the nonconsumptive effects on resource species diversity can be explained by the preferences of foragers for specific functional traits and by the forager species richness. We exposed rodents of a natural community to artificial food patches, each containing an initial multispecies resource community of eight species (10 items each) mixed in sand. We sampled 35 landscapes, each containing seven patches in a spatial array, to disentangle effects at local (patch) and landscape levels. We used vegetation height as a proxy for perceived predation risk. After a period of three nights, we counted how many and which resource species were left in each patch to measure giving-up density and resource diversity at the local level (alpha diversity) and the regional level (gamma diversity and beta diversity). Furthermore, we used wildlife cameras to identify foragers and assess their species richness. With increasing vegetation height, i.e., decreasing perceived predation risk, giving-up density, and local alpha and regional gamma diversity decreased, and patches became less similar within a landscape (beta diversity increased). Foragers consumed more of the bigger and most caloric resources. The higher the forager species richness, the lower the giving-up density, and alpha and gamma diversity. Overall, spatial variation of perceived predation risk of foragers had measurable cascading effects on local and regional resource species biodiversity, independent of the forager species. Thus, nonconsumptive predation effects modify forager-resource interactions and might act as an equalizing mechanism for species coexistence.

Fitness, risk taking, and spatial behavior covary with boldness in experimental vole populations.

Individuals of a population may vary along a pace-of-life syndrome from highly fecund, short-lived, bold, dispersive "fast" types at one end of the spectrum to less fecund, long-lived, shy, plastic "slow" types at the other end. Risk-taking behavior might mediate the underlying life history trade-off, but empirical evidence supporting this hypothesis is still ambiguous. Using experimentally created populations of common voles (Microtus arvalis)-a species with distinct seasonal life history trajectories-we aimed to test whether individual differences in boldness behavior covary with risk taking, space use, and fitness. We quantified risk taking, space use (via automated tracking), survival, and reproductive success (via genetic parentage analysis) in 8 to 14 experimental, mixed-sex populations of 113 common voles of known boldness type in large grassland enclosures over a significant part of their adult life span and two reproductive events. Populations were assorted to contain extreme boldness types (bold or shy) of both sexes. Bolder individuals took more risks than shyer ones, which did not affect survival. Bolder males but not females produced more offspring than shy conspecifics. Daily home range and core area sizes, based on 95% and 50% Kernel density estimates (20 ± 10 per individual, n = 54 individuals), were highly repeatable over time. Individual space use unfolded differently for sex-boldness type combinations over the course of the experiment. While day ranges decreased for shy females, they increased for bold females and all males. Space use trajectories may, hence, indicate differences in coping styles when confronted with a novel social and physical environment. Thus, interindividual differences in boldness predict risk taking under near-natural conditions and have consequences for fitness in males, which have a higher reproductive potential than females. Given extreme inter- and intra-annual fluctuations in population density in the study species and its short life span, density-dependent fluctuating selection operating differently on the sexes might maintain (co)variation in boldness, risk taking, and pace-of-life.

Breeding state and season affect interspecific interaction types: indirect resource competition and direct interference.

Indirect resource competition and interference are widely occurring mechanisms of interspecific interactions. We have studied the seasonal expression of these two interaction types within a two-species, boreal small mammal system. Seasons differ by resource availability, individual breeding state and intraspecific social system. Live-trapping methods were used to monitor space use and reproduction in 14 experimental populations of bank voles Myodes glareolus in large outdoor enclosures with and without a dominant competitor, the field vole Microtus agrestis. We further compared vole behaviour using staged dyadic encounters in neutral arenas in both seasons. Survival of the non-breeding overwintering bank voles was not affected by competition. In the spring, the numbers of male bank voles, but not of females, were reduced significantly in the competition populations. Bank vole home ranges expanded with vole density in the presence of competitors, indicating food limitation. A comparison of behaviour between seasons based on an analysis of similarity revealed an avoidance of costly aggression against opponents, independent of species. Interactions were more aggressive during the summer than during the winter, and heterospecific encounters were more aggressive than conspecific encounters. Based on these results, we suggest that interaction types and their respective mechanisms are not either-or categories and may change over the seasons. During the winter, energy constraints and thermoregulatory needs decrease direct aggression, but food constraints increase indirect resource competition. Direct interference appears in the summer, probably triggered by each individual's reproductive and hormonal state and the defence of offspring against conspecific and heterospecific intruders. Both interaction forms overlap in the spring, possibly contributing to spring declines in the numbers of subordinate species.