Movement and habitat selection of a large carnivore in response to human infrastructure differs by life stage.

BACKGROUND: The movement extent of mammals is influenced by human-modified areas, which can affect population demographics. Understanding how human infrastructure influences movement at different life stages is important for wildlife management. This is true especially for large carnivores, due to their substantial space requirements and potential for conflict with humans. METHODS: We investigated human impact on movement and habitat selection by GPS-collared male brown bears (Ursus arctos) in two life stages (residents and dispersers) in central Sweden. We identified dispersers visually based on their GPS locations and used hidden Markov models to delineate dispersal events. We used integrated step selection analysis (iSSA) to infer movement and habitat selection at a local scale (availability defined by hourly relocations), and resource selection functions (RSFs) to infer habitat selection at a landscape scale (availability defined by the study area extent). RESULTS: Movement of residents on a local scale was facilitated by small forestry roads as they moved faster and selected areas closer to forestry roads, and they avoided areas closer to larger public roads and buildings on both scales. Dispersers were more ambivalent in their response to human infrastructure. Dispersers increased their speed closer to small forestry roads and larger public roads, did not exhibit selection for or against any road class, and avoided areas closer to buildings only at local scale. Dispersers did not select for any features on the landscape, which is likely explained by the novelty of the landscape or their naivety towards it. CONCLUSION: Our results show that movement in male brown bears is life stage-dependent and indicate that connectivity maps derived from movement data of dispersing animals may provide more numerous and more realistic pathways than those derived from resident animal data alone. This suggests that data from dispersing animals provide more realistic models for reconnecting populations and maintaining connectivity than if data were derived from resident animals alone.

Social environment shapes female settlement decisions in a solitary carnivore.

How and where a female selects an area to settle and breed is of central importance in dispersal and population ecology as it governs range expansion and gene flow. Social structure and organization have been shown to influence settlement decisions, but its importance in the settlement of large, solitary mammals is largely unknown. We investigate how the identity of overlapping conspecifics on the landscape, acquired during the maternal care period, influences the selection of settlement home ranges in a non-territorial, solitary mammal using location data of 56 female brown bears (Ursus arctos). We used a resource selection function to determine whether females' settlement behavior was influenced by the presence of their mother, related females, familiar females, and female population density. Hunting may remove mothers and result in socio-spatial changes before settlement. We compared overlap between settling females and their mother's concurrent or most recent home ranges to examine the settling female's response to the absence or presence of her mother on the landscape. We found that females selected settlement home ranges that overlapped their mother's home range, familiar females, that is, those they had previously overlapped with, and areas with higher density than their natal ranges. However, they did not select areas overlapping related females. We also found that when mothers were removed from the landscape, female offspring selected settlement home ranges with greater overlap of their mother's range, compared with mothers who were alive. Our results suggest that females are acquiring and using information about their social environment when making settlement decisions.

Fear the reaper: ungulate carcasses may generate an ephemeral landscape of fear for rodents.

Animal carcasses provide an ephemeral pulse of nutrients for scavengers that use them. Carcass sites can increase species interactions and/or ephemeral, localized landscapes of fear for prey within the vicinity. Few studies have applied the landscape of fear to carcasses. Here, we use a mass die-off of reindeer caused by lightning in Norway to test whether rodents avoided larger scavengers (e.g. corvids and fox). We used the presence and abundance of faeces as a proxy for carcass use over the course of 2 years and found that rodents showed the strongest avoidance towards changes in raven abundance (ß = -0.469, s.e. = 0.231, p-value = 0.0429), but not fox, presumably due to greater predation risk imposed by large droves of raven. Moreover, the emergence of rodent occurrence within the carcass area corresponded well with the disappearance of raven during the second year of the study. We suggest that carcasses have the potential to shape the landscape of fear for prey, but that the overall effects of carcasses on individual fitness and populations of species ultimately depend on the carcass regime, e.g. carcass size, count, and areal extent, frequency and the scavenger guild. We discuss conservation implications and how carcass provisioning and landscapes of fear could be potentially used to manage populations and ecosystems, but that there is a gap in understanding that must first be bridged.

Hunters select for behavioral traits in a large carnivore.

Human harvest can induce selection on life history and morphological traits, leading to ecological and evolutionary responses. Our understanding of harvest-induced selection on behavioral traits is, however, very limited. Here, we assessed whether hunters harvest, consciously or not, individuals with specific behavioral traits. We used long-term, detailed behavioral and survival data of a heavily harvested brown bear (Ursus arctos) population in Sweden. We found that hunters harvested male bears that were less active during legal hunting hours and had lower movement rates. Also, hunters harvested male and female bears that used habitats closer to roads. We provide an empirical example that individual behavior can modulate vulnerability to hunting and that hunters could exert a selective pressure on wildlife behaviors. This study increases our understanding of the complex interactions between harvest method, human behavior, and animal behavior that are at play in harvest-induced selection and provides better insight into the full effects of human harvest on wild populations.

Hunting promotes spatial reorganization and sexually selected infanticide.

Harvest can affect the ecology and evolution of wild species. The removal of key individuals, such as matriarchs or dominant males, can disrupt social structure and exacerbate the impact of hunting on population growth. We do not know, however, how and when the spatiotemporal reorganization takes place after removal and if such changes can be the mechanism that explain a decrease in population growth. Detailed behavioral information from individually monitored brown bears, in a population where hunting increases sexually selected infanticide, revealed that adult males increased their use of home ranges of hunter-killed neighbors in the second year after their death. Use of a hunter-killed male's home range was influenced by the survivor's as well as the hunter-killed male's age, population density, and hunting intensity. Our results emphasize that hunting can have long-term indirect effects which can affect population viability.

Human shields mediate sexual conflict in a top predator.

Selecting the right habitat in a risky landscape is crucial for an individual's survival and reproduction. In predator-prey systems, prey often can anticipate the habitat use of their main predator and may use protective associates (i.e. typically an apex predator) as shields against predation. Although never tested, such mechanisms should also evolve in systems in which sexual conflict affects offspring survival. Here, we assessed the relationship between offspring survival and habitat selection, as well as the use of protective associates, in a system in which sexually selected infanticide (SSI), rather than interspecific predation, affects offspring survival. We used the Scandinavian brown bear (Ursus arctos) population with SSI in a human-dominated landscape as our model system. Bears, especially adult males, generally avoid humans in our study system. We used resource selection functions to contrast habitat selection of GPS-collared mothers that were successful (i.e. surviving litters, n = 19) and unsuccessful (i.e. complete litter loss, n = 11) in keeping their young during the mating season (2005-2012). Habitat selection was indeed a predictor of litter survival. Successful mothers were more likely to use humans as protective associates, whereas unsuccessful mothers avoided humans. Our results suggest that principles of predator-prey and fear ecology theory (e.g. non-consumptive and cascading effects) can also be applied to the context of sexual conflict.

Infanticide as a male reproductive strategy has a nutritive risk effect in brown bears.

Behavioural strategies to reduce predation risk can incur costs, which are often referred to as risk effects. A common strategy to avoid predation is spatio-temporal avoidance of predators, in which prey typically trade optimal resources for safety. Analogous with predator-prey theory, risk effects should also arise in species with sexually selected infanticide (SSI), in which females with dependent offspring avoid infanticidal males. SSI can be common in brown bear (Ursus arctos) populations and explains spatio-temporal segregation among reproductive classes. Here, we show that in a population with SSI, females with cubs-of-the-year had lower quality diets than conspecifics during the SSI high-risk period, the mating season. After the mating season, their diets were of similar quality to diets of their conspecifics. Our results suggest a nutritive risk effect of SSI, in which females with cubs-of-the-year alter their resource selection and trade optimal resources for offspring safety. Such risk effects can add to female costs of reproduction and may be widespread among species with SSI.