Aim We aimed to describe the large-scale patterns in population density of roe deer Caprelous capreolus in Europe and to determine the factors shaping variation in their abundance. Location Europe.Methods We collated data on roe deer population density from 72 localities spanning 25°latitude and 48°longitude and analysed them in relation to a range of environmental factors: vegetation productivity (approximated by the fraction of photosynthetically active radiation) and forest cover as proxies for food supply, winter severity, summer drought and presence or absence of large predators (wolf, Canis lupus, and Eurasian lynx, Lynx lynx), hunter harvest and a competitor (red deer, Cervus elaphus). ResultsRoe deer abundance increased with the overall productivity of vegetation cover and with lower forest cover (sparser forest cover means that a higher proportion of overall plant productivity is allocated to ground vegetation and thus is available to roe deer). The effect of large predators was relatively weak in highly productive environments and in regions with mild climate, but increased markedly in regions with low vegetation productivity and harsh winters. Other potentially limiting factors (hunting, summer drought and competition with red deer) had no significant impact on roe deer abundance. Main conclusionsThe analyses revealed the combined effect of bottom-up and top-down control on roe deer: on a biogeographical scale, population abundance of roe deer has been shaped by food-related factors and large predators, with additive effects of the two species of predators. The results have implications for management of roe deer populations in Europe. First, an increase in roe deer abundance can be expected as environmental productivity increases due to climate change. Secondly, recovery plans for large carnivores should take environmental productivity and winter severity into account when predicting their impact on prey.

Distribution, quantity and quality of food resources aff ect the diet and several other life-history traits of large mammals. Supplemental feeding of wildlife has high potential for infl uencing the behaviour and diet of opportunistic omnivores, such as bears. Supplemental feeding of brown bears Ursus arctos is a common practice in several European countries, but the eff ects of this controversial and expensive management measure on bear diet and behaviour are poorly understood. We analysed 714 brown bear scats collected throughout the year in three regions of Slovenia with diff erent densities of supplemental feeding sites. Supplemental food was the most important food category in the bear diet and represented 34% of the annual estimated dietary energy content (maize: 22%, livestock carrion: 12%). Th e proportion of supplemental food in the diet varied with season and region, being highest in spring and in the region with the highest density of feeding sites. However, considerable seasonal changes in bear diet, despite year-round access to supplemental food, suggest that bears prefer high-energy natural food sources, particularly insects, fruits, and hard mast, when available. Despite high availability and use of supplemental food, human -bear confl icts are frequent in Slovenia. In addition, evidence from earlier studies suggests that changes in diet and foraging behaviour due to supplemental feeding may aff ect several aspects of bear biology and in some cases increase the probability of human -bear confl icts. Th us, we caution against promoting unconditional supplemental feeding as a measure to prevent or reduce human -bear confl icts.

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We tested six hypotheses to explain expected geographical differences in body masses of 1,771 brown bears (Ursus arctos) from northern and southern Europe (Sweden and Norway compared with Slovenia and Croatia): Bergmann's rule, the fasting endurance hypothesis, and the dietary meat hypothesis, which predicted larger bears in the north; and hypotheses stressing the role of high primary productivity, high population density, low seasonality, and length of the growing season, which predicted larger bears in the south. Although brown bear populations in North America vary greatly in body mass, we found no significant difference in body mass between the two European populations using a new analytical approach incorporating modeled age-standardized body masses in linear models, when correcting for sex and season. The greater variation in North America may be due primarily to the presence of large bears that feed on salmon (Oncorhynchus spp.), which does not occur in Europe. Asymptotic body masses were 115 +/- 9 (SE) kg in spring and 141 +/- 9 kg in autumn for southern females, 248 +/- 25 and 243 +/- 24 kg for southern males, 96 +/- 2 and 158 +/- 4 kg for northern females, and 201 +/- 4 and 273 +/- 6 kg for northern males, respectively. Northern bears gained more body mass before hibernation and lost more during hibernation than southern bears, probably because hibernation was twice as long in the north. Northern bears gained and southern bears lost mass during the spring, perhaps due to the greater availability and use of protein-rich food in spring in the north. As reproductive success in bears is correlated with adult female body mass in interpopulation comparisons, brown bears may have relatively similar reproductive rates throughout Europe, although minimum age at primiparity and litter interval are lower in the south.

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