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As an important extrinsic source of mortality, harvest should select for fast reproduction and accelerated life histories. However, if vulnerability to harvest depends upon female reproductive status, patterns of selectivity could diverge and favor alternative reproductive behaviors. Here, using more than 20 years of detailed data on survival and reproduction in a hunted large carnivore population, we show that protecting females with dependent young, a widespread hunting regulation, provides a survival benefit to females providing longer maternal care. This survival gain compensates for the females’ reduced reproductive output, especially at high hunting pressure, where the fitness benefit of prolonged periods of maternal care outweighs that of shorter maternal care. Our study shows that hunting regulation can indirectly promote slower life histories by modulating the fitness benefit of maternal care tactics. We provide empirical evidence that harvest regulation can induce artificial selection on female life history traits and affect demographic processes.

Knowledge of factors influencing the timing of reproduction is important for animal conservation and management. Brown bears (Ursus arctos) are able to vary the birth date of their cubs in response to their fat stores, but little information is available about the timing of implantation and parturition in free-ranging brown bears. Body temperature and activity of pregnant brown bears is higher during the gestation period than during the rest of hibernation and drops at parturition. We compared mean daily body temperature and activity levels of pregnant and nonpregnant females during preimplantation, gestation, and lactation. Additionally we tested whether age, litter size, primiparity, environmental conditions, and the start of hibernation influence the timing of parturition. The mean date of implantation was 1 December (SD = 12), the mean date of parturition was 26 January (SD = 12), and the mean duration of the gestation period was 56 days (SD = 2). The body temperature of pregnant females was higher during the gestation and lactation periods than that of nonpregnant bears. The body temperature of pregnant females decreased during the gestation period. Activity recordings were also used to determine the date of parturition. The parturition dates calculated with activity and body temperature data did not differ significantly and were the same in 50% of the females. Older females started hibernation earlier. The start of hibernation was earlier during years with favorable environmental conditions. Dates of parturition were later during years with good environmental conditions which was unexpected. We suggest that free-ranging pregnant brown bears in areas with high levels of human activities at the beginning of the denning period, as in our study area, might prioritize investing energy in early denning than in early parturition during years with favorable environmental conditions, as a strategy to prevent disturbances caused by human.

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 (l ¼ 0.975; 95% CI ¼ 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. population dynamics, harvesting, brown bear, sexually selected infanticide, behaviour, carnivore, ecology, behaviour

The ecological requirements of brown bears are poorly known in the Himalaya region, which complicates conservation efforts. We documented the diet of the Himalayan brown bear (Ursus arctos isabellinus) by combining classical scat analysis and a newly developed molecular genetic technique (the trnL approach), in Deosai National Park, Pakistan. Brown bears consumed over 50 plant species, invertebrates, ungulates, and several rodents. Eight plant families; Poaceae, Polygonaceae, Cyperaceae, Apiaceae, Asteraceae, Caryophyllaceae, Lamiaceae, and Rubiaceae were commonly eaten with graminoids comprising the bulk of the diet. Golden marmots comprised the major mammalian biomass in the park, and were also the main meat source for bears. Animal matter, making 36% of dietary content, contributed half of the digestible energy, due to its higher nutritious value. We did not find a significant temporal pattern in diet, perhaps because the availability of the major diet (graminoids) did not change over the foraging period. Male brown bears were more carnivorous than females, probably because of their larger size, which requires higher energy and also makes them more efficient in capturing marmots. Frequencies of three plant species were also significantly higher in male brown bears; Bistorta affinis, Carex diluta, and Carex sp. Diet of the brown bear differed significantly between the park and surrounding valleys. In valleys, diet consisted predominantly of graminoids and crops, whereas the park provided more nutritious and diverse foodThe estimated digestible energy available to brown bears in Deosai was the lowest documented among brown bear populations, due to the lack of fruits and a relatively lower meat content. The low nutritious diet and high cost of metabolism in a high-altitude environment, probably explains the very low reproductive potential of this population.

Human persecution and habitat loss have endangered large carnivore populations worldwide, but some are recovering, exacerbating old conflicts. Carnivores can injure and kill people; the most dramatic form of wildlife-human conflict. In Scandinavia, the brown bear (Ursus arctos) population increased from ~500 bears in 1977 to ~3300 in 2008, with an increase in injuries, fatalities, and public fear of bear attacks. We reviewed media coverage and interviewed victims to explore how bear population trends, hunter education, and other factors may have influenced the number of injuries and fatalities in Scandinavia from 1977 to 2016. We found 42 incidents with 42 injuries and 2 fatalities; 42 were adult men, one was an adult woman conducting forestry work, and one was a boy skiing off-piste. Thirty-three adult men were hunting bears, moose, or small game, often with a hunting dog, and 26 had shot at the bear at 8±11 m before injury. Eleven nonhunters were conducting forestry work, inspecting a hunting area, picking berries, tending livestock, hiking, harassing a denned bear, and one person was killed outside his house at night. Eight of the 11 incidents of nonhunters involved female bears with cubs; three of these family groups were in dens and two were on carcasses. The annual number of hunters injured/killed was mostly influenced by the increase in the bear population size. The pattern was similar regarding injuries/fatalities to other outdoor users, but the relation with the bear population size was weaker than for hunters, and the null model was equally supported. Bear physiology at denning may make encounters with bears more risky in the fall, when bears show prehibernation behavior. Awareness and education efforts, especially among hunters, seem important to ensure human safety. Recreationists and forestry workers should avoid dense vegetation or make noise to warn bears of their presence.

We conducted thirteen immobilizations of previously collared hibernating two- to four-year-old brown bears (Ursus arctos) weighing 21-66 kg in central Sweden in winter 2010 and 2011 for comparative physiology research. Here we report, for the first time, an effective protocol for the capture and anesthesia of free-ranging brown bears during hibernation and an assessment of the disturbance the captures caused. Bears were darted in anthill, soil, or uprooted tree dens on eleven occasions, but two bears in rock dens fled and were darted outside the den. We used medetomidine at 0.02-0.06 mg/kg and zolazepam-tiletamine at 0.9-2.8 mg/kg for anesthesia. In addition, ketamine at 1.5 mg/kg was hand-injected intramuscularly in four bears and in six it was included in the dart at 1.1-3.0 mg/kg. Once anesthetized, bears were removed from the dens. In nine bears, arterial blood samples were analyzed immediately with a portable blood gas analyzer. We corrected hypoxemia in seven bears (PaO(2) 57-74 mmHg) with supplemental oxygen. We placed the bears back into the dens and antagonized the effect of medetomidine with atipamezole. Capturing bears in the den significantly increased the risk of den abandonment. One of twelve collared bears that were captured remained at the original den until spring, and eleven, left their dens (mean ± standard deviation) 3.2±3.6 (range 0.5-10.5) days after capture. They used 1.9±0.9 intermediate resting sites, during 6.2±7.8 days before entering a new permanent den. The eleven new permanent dens were located 730±589 m from the original dens. We documented that it was feasible and safe to capture hibernating brown bears, although they behaved differently than black bears. When doing so, researchers should use 25% of the doses used for helicopter darting during the active period and should consider increased energetic costs associated with den abandonment.

One of the classic approaches in environmental economics is the environmental Kuznets curve, which predicts that when a national economy grows from low to medium levels, threats to biodiversity conservation increase, but they decrease when the economy moves from medium to high. We evaluated this approach by examining how population densities of the brown bear (Ursus arctos), gray wolf (Canis lupus), and Eurasian lynx (Lynx lynx) were related to the national economy in 24 European countries. We used forest proportions, the existence of a compensation system, and country group (former socialist countries, Nordic countries, other countries) as covariates in a linear model with the first- and the second-order polynomial terms of per capita gross domestic product (GDP). Country group was treated as a random factor, but remained insignificant and was ignored. All models concerning brown bear and wolf provided evidence that population densities decreased with increasing GDP, but densities of lynx were virtually independent of GDP. Models for the wolf explained >80% of the variation in densities, without a difference between the models with all independent variables and the model with only GDP. For the bear, the model with GDP alone accounted for 10%, and all three variables 33%, of the variation in densities. Wolves exhibit a higher capacity for dispersal and reproduction than bear or lynx, but still exists at the lowest densities in wealthy European countries. We are aware that several other factors, not available for our models, influenced large carnivore densities. Based on the pronounced differences among large carnivore species in their countrywide relationships between densities and GDP, and a strikingly high relationship for the gray wolf, we suggest that our results reflected differences in political history and public acceptance of these species among countries. The compensation paid for the damages caused by the carnivores is not a key to higher carnivore densities, but might be necessity for the presence of large carnivores to be accepted in countries with high GDP.

Recreation is a crucial contribution of nature to people, relevant for forest ecosystems. Large carnivores (LCs) are important components of forests, however, their contribution to forest recreational value has not yet been evaluated. Given the current expansion of LC populations, the ongoing forest conservation debate, and the increasing use of nature for recreational purposes, this is a timely study. We used discrete choice experiments and willingness-to-travel to determine people’ preferences for both forest structural characteristics and presence of four LC species in Poland (N = 1097 respondents) and Norway (N = 1005). In both countries, two-thirds of the respondents (termed ‘wildness-positive’) perceived LCs as contributing positively to forest recreational value and preferred to visit old forests with trees of different species and ages and presence of dead wood (i.e. natural forests). Respondents with negative preferences towards LCs preferred more intensively managed forest (‘wildness-negative’); their preferences were stronger than in wildness-positive respondents and in Norway. Preferences towards wild nature were highly polarized and there were hardly neutral people. Our results showed a strong link between preferences for LC presence and forest structure, and reflected the dualism of human-nature relationships. This study highlights the need to consider the contribution of forests and LCs to human recreation services in ecosystem management policies.

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 β 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.

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