Scavenger-induced scattering of wild boar carcasses over large distances and its implications for disease management.

Vertebrate scavengers provide essential ecosystem services such as accelerating carrion decomposition by consuming carcasses, exposing tissues to microbial and invertebrate decomposers, and recycling nutrients back into the ecosystem. Some scavengers do not consume carcasses on site but rather scatter their remains in the surroundings, which might have important implications for nutrient transport, forensic investigations and the spread of diseases such as African Swine Fever. However, only a few studies have investigated and measured the scatter distances. Using wild boar (Sus scrofa) carcasses and limbs, we monitored scavenging behavior and measured scatter distances of mammals. We placed 20 carcasses (up to 25 kg) and 21 separate limbs equipped with very high frequency (VHF) transmitters and monitored scavenger activity using camera traps in a mountainous region in southeast Germany. Except for one carcass, all other carcasses and limbs were scattered. We measured 72 scatter distances (of 89 scattering events; mean = 232 m, maximum = 1250 m), of which 75% were dispersed up to 407 m. Scavengers moved scattered pieces into denser vegetation compared to the half-open vegetation at provisioning sites. Red foxes (Vulpes vulpes) were the most common scavenger species, contributing to 72 scattering events (58 measured scatter distances). Our results provide evidence of scatter distances farther than previously assumed and have far-reaching implications for disease management or forensic investigations, as the broader surroundings of carcasses must be included in search efforts to remove infectious material or relevant body parts for forensic analysis.

Range-wide evolutionary relationships and historical demography of brown bears (Ursus arctos) revealed by whole-genome sequencing of isolated central Asian populations.

Phylogeographic studies uncover hidden pathways of divergence and inform conservation. Brown bears (Ursus arctos) have one of the broadest distributions of all land mammals, ranging from Eurasia to North America, and are an important model for evolutionary studies. Although several whole genomes were available for individuals from North America, Europe and Asia, limited whole-genome data were available from Central Asia, including the highly imperilled brown bears in the Gobi Desert. To fill this knowledge gap, we sequenced whole genomes from nine Asian brown bears from the Gobi Desert of Mongolia, Northern Mongolia and the Himalayas of Pakistan. We combined these data with published brown bear sequences from Europe, Asia and North America, as well as other bear species. Our goals were to determine the evolutionary relationships among brown bear populations worldwide, their genetic diversity and their historical demography. Our analyses revealed five major lineages of brown bears based on a filtered set of 684,081 single nucleotide polymorphisms. We found distinct evolutionary lineages of brown bears in the Gobi, Himalayas, northern Mongolia, Europe and North America. The lowest level of genetic diversity and the highest level of inbreeding were found in Pakistan, the Gobi Desert and Central Italy. Furthermore, the effective population size (Ne ) for all brown bears decreased over the last 70,000 years. Our results confirm the genetic distinctiveness and ancient lineage of brown bear subspecies in the Gobi Desert of Mongolia and the Himalayas of Pakistan and highlight their importance for conservation.

Landscape of fear or landscape of food? Moose hunting triggers an antipredator response in brown bears.

Hunters can affect the behavior of wildlife by inducing a landscape of fear, selecting individuals with specific traits, or altering resource availability across the landscape. Most research investigating the influence of hunting on wildlife resource selection has focused on target species and less attention has been devoted to nontarget species, such as scavengers that can be both attracted or repelled by hunting activities. We used resource selection functions to identify areas where hunters were most likely to kill moose (Alces alces) in south-central Sweden during the fall. Then, we used step-selection functions to determine whether female brown bears (Ursus arctos) selected or avoided these areas and specific resources during the moose hunting season. We found that, during both day and nighttime, female brown bears avoided areas where hunters were more likely to kill moose. We found evidence that resource selection by brown bears varied substantially during the fall and that some behavioral changes were consistent with disturbance associated with moose hunters. Brown bears were more likely to select concealed locations in young (i.e., regenerating) and coniferous forests and areas further away from roads during the moose hunting season. Our results suggest that brown bears react to both spatial and temporal variations in apparent risk during the fall: moose hunters create a landscape of fear and trigger an antipredator response in a large carnivore even if bears are not specifically targeted during the moose hunting season. Such antipredator responses might lead to indirect habitat loss and lower foraging efficiency and the resulting consequences should be considered when planning hunting seasons.

Toxic elements in arctic and sub-arctic brown bears: Blood concentrations of As, Cd, Hg and Pb in relation to diet, age, and human footprint.

Contamination with arsenic (As), cadmium (Cd), mercury (Hg) and lead (Pb) is a global concern impairing resilience of organisms and ecosystems. Proximity to emission sources increases exposure risk but remoteness does not alleviate it. These toxic elements are transported in atmospheric and oceanic pathways and accumulate in organisms. Mercury accumulates in higher trophic levels. Brown bears (Ursus arctos), which often live in remote areas, are long-lived omnivores, feeding on salmon (Oncorhynchus spp.) and berries (Vaccinium spp.), resources also consumed by humans. We measured blood concentrations of As, Cd, Hg and Pb in bears (n = 72) four years and older in Scandinavia and three national parks in Alaska, USA (Lake Clark, Katmai and Gates of the Arctic) using high-resolution, inductively-coupled plasma sector field mass spectrometry. Age and sex of the bears, as well as the typical population level diet was associated with blood element concentrations using generalized linear regression models. Alaskan bears consuming salmon had higher Hg blood concentrations compared to Scandinavian bears feeding on berries, ants (Formica spp.) and moose (Alces). Cadmium and Pb blood concentrations were higher in Scandinavian bears than in Alaskan bears. Bears using marine food sources, in addition to salmon in Katmai, had higher As blood concentrations than bears in Scandinavia. Blood concentrations of Cd and Pb, as well as for As in female bears increased with age. Arsenic in males and Hg concentrations decreased with age. We detected elevated levels of toxic elements in bears from landscapes that are among the most pristine on the planet. Sources are unknown but anthropogenic emissions are most likely involved. All study areas face upcoming change: Increasing tourism and mining in Alaska and more intensive forestry in Scandinavia, combined with global climate change in both regions. Baseline contaminant concentrations as presented here are important knowledge in our changing world.

Forests buffer the climate-induced decline of body mass in a mountain herbivore.

Climate change is known to affect key life-history traits, such as body mass, reproduction, and survival in many species. Animal populations inhabiting mountain habitats are adapted to extreme seasonal environmental conditions but are also expected to be especially vulnerable to climate change. Studies on mountain ungulates typically focus on populations or sections of populations living above the tree line, whereas populations inhabiting forested habitats are largely understudied. Here, we investigate whether forested areas can mitigate the impact of climatic change on life-history traits by evaluating the interactive effects of temperature and habitat characteristics on body mass variation in the Alpine chamois Rupicapra rupicapra rupicapra. We examined data of 20,573 yearling chamois collected from 1993 to 2019 in 28 mountain ranges in the Austrian Eastern Alps, characterized by different proportion of forest cover. Our results show that the temporal decline of chamois body mass is less pronounced in areas with greater proportion of forest cover. For chamois living in forest habitats only, no significant temporal change in body mass was detected. Variation in body mass was affected by the interaction between density and snow cover, as well as by the interaction between spring temperatures and forest cover, supporting the role of forests as thermal buffer against the effects of increasing temperatures on life-history traits in a mountain ungulate. In turn, this study suggests a buffering effect of forests against climate change impacts. Assessments of the consequences of climate change on the life-history traits and population dynamics of mountain-dwelling species should thus consider the plasticity of the species with respect to the use and availability of different habitat types.

Disentangling direct and indirect determinants of the duration of maternal care in brown bears: Environmental context matters.

The duration of maternal care, an important life-history trait affecting population dynamics, varies greatly within species. Yet, our understanding of its predictors is limited, mostly correlative and subject to misinterpretations, due to difficulties to disentangle the role of maternal- and offspring-related characteristics. We conducted path analysis on a dataset including 217 brown bear litters captured over a 29-year period in two populations in Sweden ('North' and 'South') facing contrasting environmental conditions to identify and quantify the causes of variation in the duration of maternal care (1.5 or 2.5 years). We showed that the causal determinants of the duration of maternal care were context-dependent. Contrary to their expected central role in the determination of the duration of maternal care, yearling mass and its direct determinants (i.e. litter size and maternal mass) were only important in the North population, where environmental conditions are harsher and the cost of extended maternal care presumably higher. In the South, the duration of maternal care was not caused by yearling mass nor any maternal or litter characteristics. Extension of maternal care may thus result from factors independent from maternal and offspring condition in the South, such as an artificial hunting-induced selection for longer maternal care through the legal protection of family groups. Our results provide an important contribution to our very limited knowledge of the direct and indirect determinants of the duration of maternal care and highlight the importance of accounting for the environmental context when assessing maternal reproductive tactics.

Biologging reveals individual variation in behavioural predictability in the wild.

Recent research highlights the ecological importance of individual variation in behavioural predictability. Individuals may not only differ in their average expression of a behavioural trait (their behavioural type) and in their ability to adjust behaviour to changing environmental conditions (individual plasticity), but also in their variability around their average behaviour (predictability). However, quantifying behavioural predictability in the wild has been challenging due to limitations of acquiring sufficient repeated behavioural measures. We here demonstrate how common biologging data can be used to detect individual variation in behavioural predictability in the wild and reveal the coexistence of highly predictable individuals along with unpredictable individuals within the same population. We repeatedly quantified two behaviours-daily movement distance and diurnal activity-in 62 female brown bears Ursus arctos tracked across 187 monitoring years. We calculated behavioural predictability over the short term (50 consecutive monitoring days within 1 year) and long term (across monitoring years) as the residual intra-individual variability (rIIV) of behaviour around the behavioural reaction norm. We tested whether predictability varies systematically across average behavioural types and whether it is correlated across functionally distinct behaviours, that is, daily movement distance and amount of diurnal activity. Brown bears showed individual variation in behavioural predictability from predictable to unpredictable individuals. For example, the standard deviation around the average daily movement distance within one monitoring year varied up to fivefold from 1.1 to 5.5 km across individuals. Individual predictability for both daily movement distance and diurnality was conserved across monitoring years. Individual predictability was correlated with behavioural type where individuals which were on average more diurnal and mobile were also more unpredictable in their behaviour. In contrast, more nocturnal individuals moved less and were more predictable in their behaviour. Finally, individual predictability in daily movement distance and diurnality was positively correlated, suggesting that individual predictability may be a quantitative trait in its own regard that could evolve and is underpinned by genetic variation. Unpredictable individuals may cope better with stochastic events and unpredictability may hence be an adaptive behavioural response to increased predation risk. Coexistence of predictable and unpredictable individuals may therefore ensure adaptable and resilient populations.

Grading fecal consistency in an omnivorous carnivore, the brown bear: Abandoning the concept of uniform feces.

Grading the fecal consistency of carnivores is a frequently used tool for monitoring gut health and overall digestion. Several fecal consistency grading systems are available for mainly felids and canids. No such system exists for the brown bear (Ursus arctos Linnaeus, 1758). We aim at extending current fecal consistency grading systems with a scoring system for brown bears. The system was set up during a diet study with nine individuals fed a variety of diets including beef meat, rabbit, fruit, and grass-fruit-pellet mix in an incomplete crossover design. One additional individual was included opportunistically and was fed the typical zoo diet (vegetable-fruit-meat-pellet diet). All feces from the collection period were photographed, graded by "handling the feces" and visually inspected for dietary components. Based on a total of 446 feces, a six-point scale for uniform fecal consistencies was established. In 11% of all feces, two distinct consistencies could be distinguished, a feature that appears in other carnivore species as well. Hence, an additional grading system for dual consistencies was developed. The fecal consistency of brown bears is heavily dependent on the diet items processed before defecation with the general observation that the more vegetation or whole prey, the firmer the feces, and at certain proportions of the latter, the higher the chance for dual fecal consistencies to occur. The results indicate that in bears, diet may have a strong effect on fecal consistency, hampering animal health assessments without prior knowledge of the diet.

Fears from the past? The innate ability of dogs to detect predator scents.

Throughout the animal kingdom, antipredator mechanisms are an evolutionary driving force to enable the survival of species classified as prey. Information regarding a predator's location can be determined through chemosensory cues from urine, faeces, visual and/or acoustic signals and anal gland secretions; and in several lab and field-based studies it has been seen that these cues mediate behavioural changes within prey species. These behaviours are often linked to fear and avoidance, which will in turn increase the prey's survival rate. In many studies dogs (Canis lupus familiaris) have been used as a predator species, however, no research has addressed a dog's innate ability to detect predator scents, hence the rationale behind this study. We assessed the innate ability of the untrained domestic dog to detect faecal scents of wild Eurasian brown bear (Ursus arctos arctos) and European lynx (Lynx lynx). The study monitored 82 domestic dogs across the UK and Norway. The dogs were exposed to the two predator faecal scents from Eurasian brown bear and European lynx, a herbivore faecal scent of Eurasian beaver (Castor fiber) and water control. Measurements were taken upon the time spent within a 40 cm radius of each scent and changes in the dog's heart rate when within this 40 cm radius. We found dogs spent a decreased length of time around the predator scents and had an increased heart rate in relation to their basal heart rate. We conclude that dogs can innately sense predator scents of brown bear and lynx and elicit fear towards these odours, as shown through behavioural and physiological changes.

Causes and consequences of inverse density-dependent territorial behaviour and aggression in a monogamous mammal.

Territoriality is an important process shaping population dynamics, and the defence of a territory is crucial for individuals to increase the duration of territory occupancy and, consequently, reproductive success. However, little is known about how the frequency of territory intrusions and subsequent territorial behaviours and aggression by territory owners are affected by external factors, such as population density. This is important because it can affect mate change (the replacement of one pair member) and dispersal, a key ecological process. The aim of this study was to investigate the behavioural and spatial response of territory owners to intruder pressure as a function of population density in a territorial, monogamous mammal, the Eurasian beaver (Castor fiber). Using a combination of GPS technology, scent experiments, camera trap data and tail scar observations from an individual-based long-term study, we investigated the factors influencing spatial movement patterns by territory owners in response to a simulated intruder and the factors affecting territory intrusions. We found consistent inverse density-dependent patterns in territorial behaviours and evidence of conspecific aggression. At lower densities, territory owners detected more simulated intrusions, showed more territorial reactions and experienced increased conspecific aggression as indicated by tail scars, suggesting increased intruder pressure. Inverse density-dependent territorial behaviour and aggression suggest a potential mechanistic link between inverse density-dependent natal dispersal and mate change. At low population densities, increased dispersal amplifies intruder pressure, leading to the observed increases in territorial behaviours, conspecific aggression and previously observed mate turnover, which in turn might increase natal dispersal. Our study demonstrates how population density can affect the behaviour and space use of individuals, which is important for territory occupancy and fitness.

Metabolic reprogramming involving glycolysis in the hibernating brown bear skeletal muscle.

BACKGROUND: In mammals, the hibernating state is characterized by biochemical adjustments, which include metabolic rate depression and a shift in the primary fuel oxidized from carbohydrates to lipids. A number of studies of hibernating species report an upregulation of the levels and/or activity of lipid oxidizing enzymes in muscles during torpor, with a concomitant downregulation for glycolytic enzymes. However, other studies provide contrasting data about the regulation of fuel utilization in skeletal muscles during hibernation. Bears hibernate with only moderate hypothermia but with a drop in metabolic rate down to ~ 25% of basal metabolism. To gain insights into how fuel metabolism is regulated in hibernating bear skeletal muscles, we examined the vastus lateralis proteome and other changes elicited in brown bears during hibernation. RESULTS: We show that bear muscle metabolic reorganization is in line with a suppression of ATP turnover. Regulation of muscle enzyme expression and activity, as well as of circulating metabolite profiles, highlighted a preference for lipid substrates during hibernation, although the data suggested that muscular lipid oxidation levels decreased due to metabolic rate depression. Our data also supported maintenance of muscle glycolysis that could be fuelled from liver gluconeogenesis and mobilization of muscle glycogen stores. During hibernation, our data also suggest that carbohydrate metabolism in bear muscle, as well as protein sparing, could be controlled, in part, by actions of n-3 polyunsaturated fatty acids like docosahexaenoic acid. CONCLUSIONS: Our work shows that molecular mechanisms in hibernating bear skeletal muscle, which appear consistent with a hypometabolic state, likely contribute to energy and protein savings. Maintenance of glycolysis could help to sustain muscle functionality for situations such as an unexpected exit from hibernation that would require a rapid increase in ATP production for muscle contraction. The molecular data we report here for skeletal muscles of bears hibernating at near normal body temperature represent a signature of muscle preservation despite atrophying conditions.

Sociodemographic factors modulate the spatial response of brown bears to vacancies created by hunting.

There is a growing recognition of the importance of indirect effects from hunting on wildlife populations, e.g. social and behavioural changes due to harvest, which occur after the initial offtake. Nonetheless, little is known about how the removal of members of a population influences the spatial configuration of the survivors. We studied how surviving brown bears (Ursus arctos) used former home ranges that had belonged to casualties of the annual bear hunting season in southcentral Sweden (2007-2015). We used resource selection functions to explore the effects of the casualty's and survivor's sex, age and their pairwise genetic relatedness, population density and hunting intensity on survivors' spatial responses to vacated home ranges. We tested the competitive release hypothesis, whereby survivors that increase their use of a killed bear's home range are presumed to have been released from intraspecific competition. We found strong support for this hypothesis, as survivors of the same sex as the casualty consistently increased their use of its vacant home range. Patterns were less pronounced or absent when the survivor and casualty were of opposite sex. Genetic relatedness between the survivor and the casualty emerged as the most important factor explaining increased use of vacated male home ranges by males, with a stronger response from survivors of lower relatedness. Relatedness was also important for females, but it did not influence use following removal; female survivors used home ranges of higher related female casualties more, both before and after death. Spatial responses by survivors were further influenced by bear age, population density and hunting intensity. We have shown that survivors exhibit a spatial response to vacated home ranges caused by hunting casualties, even in nonterritorial species such as the brown bear. This spatial reorganization can have unintended consequences for population dynamics and interfere with management goals. Altogether, our results underscore the need to better understand the short- and long-term indirect effects of hunting on animal social structure and their resulting distribution in space.

Hunting promotes sexual conflict in brown bears.

The removal of individuals through hunting can destabilize social structure, potentially affecting population dynamics. Although previous studies have shown that hunting can indirectly reduce juvenile survival through increased sexually selected infanticide (SSI), very little is known about the spatiotemporal effects of male hunting on juvenile survival. Using detailed individual monitoring of a hunted population of brown bears (Ursus arctos) in Sweden (1991-2011), we assessed the spatiotemporal effect of male removal on cub survival. We modelled cub survival before, during and after the mating season. We used three proxies to evaluate spatial and temporal variation in male turnover; distance and timing of the closest male killed and number of males that died around a female's home range centre. Male removal decreased cub survival only during the mating season, as expected in seasonal breeders with SSI. Cub survival increased with distance to the closest male killed within the previous 1·5 years, and it was lower when the closest male killed was removed 1·5 instead of 0·5 year earlier. We did not detect an effect of the number of males killed. Our results support the hypothesis that social restructuring due to hunting can reduce recruitment and suggest that the distribution of the male deaths might be more important than the overall number of males that die. As the removal of individuals through hunting is typically not homogenously distributed across the landscape, spatial heterogeneity in hunting pressure may cause source-sink dynamics, with lower recruitment in areas of high human-induced mortality.

Can hunting data be used to estimate unbiased population parameters? A case study on brown bears.

Quantifying temporal changes in harvested populations is critical for applied and fundamental research. Unbiased data are required to detect true changes in phenotypic distribution or population size. Because of the difficulty of collecting detailed individual data from wild populations, data from hunting records are often used. Hunting records, however, may not represent a random sample of a population. We aimed to detect and quantify potential bias in hunting records. We compared data from a long-term monitoring project with hunting records of brown bears (Ursus arctos) in Sweden and investigated temporal trends (1996-2013) in the ratio of yearlings to adult females, yearling mass and adult female mass. Data from hunting records underestimated the decline in yearling and adult female mass over time, most likely owing to the legal protection of family groups from hunting, but reflected changes in the ratio of yearlings to adult females more reliably. Although hunting data can be reliable to approximate population abundance in some circumstances, hunting data can represent a biased sample of a population and should be used with caution in management and conservation decisions.

Quantifying consistent individual differences in habitat selection.

Habitat selection is a fundamental behaviour that links individuals to the resources required for survival and reproduction. Although natural selection acts on an individual's phenotype, research on habitat selection often pools inter-individual patterns to provide inferences on the population scale. Here, we expanded a traditional approach of quantifying habitat selection at the individual level to explore the potential for consistent individual differences of habitat selection. We used random coefficients in resource selection functions (RSFs) and repeatability estimates to test for variability in habitat selection. We applied our method to a detailed dataset of GPS relocations of brown bears (Ursus arctos) taken over a period of 6 years, and assessed whether they displayed repeatable individual differences in habitat selection toward two habitat types: bogs and recent timber-harvest cut blocks. In our analyses, we controlled for the availability of habitat, i.e. the functional response in habitat selection. Repeatability estimates of habitat selection toward bogs and cut blocks were 0.304 and 0.420, respectively. Therefore, 30.4 and 42.0 % of the population-scale habitat selection variability for bogs and cut blocks, respectively, was due to differences among individuals, suggesting that consistent individual variation in habitat selection exists in brown bears. Using simulations, we posit that repeatability values of habitat selection are not related to the value and significance of ß estimates in RSFs. Although individual differences in habitat selection could be the results of non-exclusive factors, our results illustrate the evolutionary potential of habitat selection.

Temporal effects of hunting on foraging behavior of an apex predator: Do bears forego foraging when risk is high?

Avoiding predators most often entails a food cost. For the Scandinavian brown bear (Ursus arctos), the hunting season coincides with the period of hyperphagia. Hunting mortality risk is not uniformly distributed throughout the day, but peaks in the early morning hours. As bears must increase mass for winter survival, they should be sensitive to temporal allocation of antipredator responses to periods of highest risk. We expected bears to reduce foraging activity at the expense of food intake in the morning hours when risk was high, but not in the afternoon, when risk was low. We used fine-scale GPS-derived activity patterns during the 2 weeks before and after the onset of the annual bear hunting season. At locations of probable foraging, we assessed abundance and sugar content, of bilberry (Vaccinium myrtillus), the most important autumn food resource for bears in this area. Bears decreased their foraging activity in the morning hours of the hunting season. Likewise, they foraged less efficiently and on poorer quality berries in the morning. Neither of our foraging measures were affected by hunting in the afternoon foraging bout, indicating that bears did not allocate antipredator behavior to times of comparably lower risk. Bears effectively responded to variation in risk on the scale of hours. This entailed a measurable foraging cost. The additive effect of reduced foraging activity, reduced forage intake, and lower quality food may result in poorer body condition upon den entry and may ultimately reduce reproductive success.

The relative importance of direct and indirect effects of hunting mortality on the population dynamics of brown bears.

There is increasing evidence of indirect effects of hunting on populations. In species with sexually selected infanticide (SSI), hunting may decrease juvenile survival by increasing male turnover. We aimed to evaluate the relative importance of direct and indirect effects of hunting via SSI on the population dynamics of the Scandinavian brown bear (Ursus arctos). We performed prospective and retrospective demographic perturbation analyses for periods with low and high hunting pressures. All demographic rates, except yearling survival, were lower under high hunting pressure, which led to a decline in population growth under high hunting pressure (λ = 0.975; 95% CI = 0.914-1.011). Hunting had negative indirect effects on the population through an increase in SSI, which lowered cub survival and possibly also fecundity rates. Our study suggests that SSI could explain 13.6% of the variation in population growth. Hunting also affected the relative importance of survival and fecundity of adult females for population growth, with fecundity being more important under low hunting pressure and survival more important under high hunting pressure. Our study sheds light on the importance of direct and indirect effects of hunting on population dynamics, and supports the contention that hunting can have indirect negative effects on populations through SSI.

Socio-ecological features other than sex affect habitat selection in the socially obligate monogamous Eurasian beaver.

Habitat selection is a context-dependent mechanism, in which both the internal state as well as external factors affect the behavior and decisions of an individual. This is well known for polygamous mammals, which are typically sexually dimorphic, and often express great variability in behavior and habitat selection between individuals as well between the sexes. Among monogamous mammals, however, variability in habitat selection should be explained by group characteristics and the presence of offspring rather than by sex. We evaluated this hypothesis in a socially monogamous rodent, the Eurasian beaver (Castor fiber), in a saturated Norwegian population. For the first time in this species we applied GPS tracking devices (N = 22 adult beavers, in 15 territories, 2009-2013), and used resource selection functions (i) to document population-wide habitat selection and the importance of 'territory' therein, and (ii) to evaluate which socio-ecological factors explained potential individual differences in habitat selection. We found that variation in habitat selection was stronger between territories than between years or individuals nested by territory. We identified that family size and the presence of kits, but not sex, explained individual variation in habitat selection. Adults with kits and/or larger families tended to exhibit low risk-taking behavior (avoiding human-related variables such as roads, buildings, and agricultural land), and stayed close to their main lodge (parental care). Our results show that habitat selection is a context-dependent mechanism even in a species which expresses very little behavioral and morphological dimorphism.

A "clearcut" case? Brown bear selection of coarse woody debris and carpenter ants on clearcuts.

Forest management alters habitat characteristics, resulting in various effects among and within species. It is crucial to understand how habitat alteration through forest management (e.g. clearcutting) affects animal populations, particularly with unknown future conditions (e.g. climate change). In Sweden, brown bears (Ursus arctos) forage on carpenter ants (Camponotus herculeanus) during summer, and may select for this food source within clearcuts. To assess carpenter ant occurrence and brown bear selection of carpenter ants, we sampled 6999 coarse woody debris (CWD) items within 1019 plots, of which 902 were within clearcuts (forests ⩽30 years of age) and 117 plots outside clearcuts (forests >30 years of age). We related various CWD and site characteristics to the presence or absence of carpenter ant galleries (nests) and bear foraging sign at three spatial scales: the CWD, plot, and clearcut scale. We tested whether both absolute and relative counts (the latter controlling for the number of CWD items) of galleries and bear sign in plots were higher inside or outside clearcuts. Absolute counts were higher inside than outside clearcuts for galleries (mean counts; inside: 1.8, outside: 0.8). CWD was also higher inside (mean: 6.8) than outside clearcuts (mean: 4.0). However, even after controlling for more CWD inside clearcuts, relative counts were higher inside than outside clearcuts for both galleries (mean counts; inside: 0.3, outside: 0.2) and bear sign (mean counts; inside: 0.03, outside: 0.01). Variables at the CWD scale best explained gallery and bear sign presence than variables at the plot or clearcut level, but bear selection was influenced by clearcut age. CWD circumference was important for both carpenter ant and bear sign presence. CWD hardness was most important for carpenter ant selection. However, the most important predictor for bear sign was the presence or absence of carpenter ant galleries. Bears had a high foraging "success" rate (⩾88%) in foraging CWD where galleries also occurred, which was assessed by summing CWD items with the concurrence of bear sign and galleries, divided by the sum of all CWD with bear sign. Clearcuts appeared to increase the occurrence of a relatively important summer food item, the carpenter ant, on Swedish managed forests for the brown bear. However, the potential benefit of this increase can only be determined from a better understanding of the seasonal and interannual variation of the availability and use of other important brown bear food items, berries (e.g. Vaccinium myrtillus and Empetrum spp.), as well as other primary needs for bears (e.g. secure habitat and denning habitat), within the landscape mosaic of managed forests.

Does despotic behavior or food search explain the occurrence of problem brown bears in Europe?

Bears foraging near human developments are often presumed to be responding to food shortage, but this explanation ignores social factors, in particular despotism in bears. We analyzed the age distribution and body condition index (BCI) of shot brown bears in relation to densities of bears and people, and whether the shot bears were killed by managers (i.e., problem bears; n = 149), in self-defense (n = 51), or were hunter-killed nonproblem bears (n = 1,896) during 1990-2010. We compared patterns between areas with (Slovenia) and without supplemental feeding (Sweden) of bears relative to 2 hypotheses. The food-search/food-competition hypothesis predicts that problem bears should have a higher BCI (e.g., exploiting easily accessible and/or nutritious human-derived foods) or lower BCI (e.g., because of food shortage) than nonproblem bears, that BCI and human density should have a positive correlation, and problem bear occurrence and seasonal mean BCI of nonproblem bears should have a negative correlation (i.e., more problem bears during years of low food availability). Food competition among bears additionally predicts an inverse relationship between BCI and bear density. The safety-search/naivety hypothesis (i.e., avoiding other bears or lack of human experience) predicts no relationship between BCI and human density, provided no dietary differences due to spatiotemporal habitat use among bears, no relationship between problem bear occurrence and seasonal mean BCI of nonproblem bears, and does not necessarily predict a difference between BCI for problem/nonproblem bears. If food competition or predation avoidance explained bear occurrence near settlements, we predicted younger problem than nonproblem bears and a negative correlation between age and human density. However, if only food search explained bear occurrence near settlements, we predicted no relation between age and problem or nonproblem bear status, or between age and human density. We found no difference in BCI or its variability between problem and nonproblem bears, no relation between BCI and human density, and no correlation between numbers of problem bears shot and seasonal mean BCI for either country. The peak of shot problem bears occurred from April to June in Slovenia and in June in Sweden (i.e., during the mating period when most intraspecific predation occurs and before fall hyperphagia). Problem bears were younger than nonproblem bears, and both problem and nonproblem bears were younger in areas of higher human density. These age differences, in combination with similarities in BCI between problem and nonproblem bears and lack of correlation between BCI and human density, suggested safety-search and naïve dispersal to be the primary mechanisms responsible for bear occurrence near settlements. Younger bears are less competitive, more vulnerable to intraspecific predation, and lack human experience, compared to adults. Body condition was inversely related to the bear density index in Sweden, whereas we found no correlation in Slovenia, suggesting that supplemental feeding may have reduced food competition, in combination with high bear harvest rates. Bears shot in self-defense were older and their BCI did not differ from that of nonproblem bears. Reasons other than food shortage apparently explained why most bears were involved in encounters with people or viewed as problematic near settlements in our study.