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.

High-throughput microsatellite genotyping in ecology: improved accuracy, efficiency, standardization and success with low-quantity and degraded DNA.

Microsatellite markers have played a major role in ecological, evolutionary and conservation research during the past 20 years. However, technical constrains related to the use of capillary electrophoresis and a recent technological revolution that has impacted other marker types have brought to question the continued use of microsatellites for certain applications. We present a study for improving microsatellite genotyping in ecology using high-throughput sequencing (HTS). This approach entails selection of short markers suitable for HTS, sequencing PCR-amplified microsatellites on an Illumina platform and bioinformatic treatment of the sequence data to obtain multilocus genotypes. It takes advantage of the fact that HTS gives direct access to microsatellite sequences, allowing unambiguous allele identification and enabling automation of the genotyping process through bioinformatics. In addition, the massive parallel sequencing abilities expand the information content of single experimental runs far beyond capillary electrophoresis. We illustrated the method by genotyping brown bear samples amplified with a multiplex PCR of 13 new microsatellite markers and a sex marker. HTS of microsatellites provided accurate individual identification and parentage assignment and resulted in a significant improvement of genotyping success (84%) of faecal degraded DNA and costs reduction compared to capillary electrophoresis. The HTS approach holds vast potential for improving success, accuracy, efficiency and standardization of microsatellite genotyping in ecological and conservation applications, especially those that rely on profiling of low-quantity/quality DNA and on the construction of genetic databases. We discuss and give perspectives for the implementation of the method in the light of the challenges encountered in wildlife studies.

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.

Drivers of hibernation in the brown bear.

BACKGROUND: Hibernation has been a key area of research for several decades, essentially in small mammals in the laboratory, yet we know very little about what triggers or ends it in the wild. Do climatic factors, an internal biological clock, or physiological processes dominate? Using state-of-the-art tracking and monitoring technology on fourteen free-ranging brown bears over three winters, we recorded movement, heart rate (HR), heart rate variability (HRV), body temperature (Tb), physical activity, ambient temperature (TA), and snow depth to identify the drivers of the start and end of hibernation. We used behavioral change point analyses to estimate the start and end of hibernation and convergent cross mapping to identify the causal interactions between the ecological and physiological variables over time. RESULTS: To our knowledge, we have built the first chronology of both ecological and physiological events from before the start to the end of hibernation in the field. Activity, HR, and Tb started to drop slowly several weeks before den entry. Bears entered the den when snow arrived and when ambient temperature reached 0 °C. HRV, taken as a proxy of sympathetic nervous system activity, dropped dramatically once the bear entered the den. This indirectly suggests that denning is tightly coupled to metabolic suppression. During arousal, the unexpected early rise in Tb (two months before den exit) was driven by TA, but was independent of HRV. The difference between Tb and TA decreased gradually suggesting that bears were not thermoconforming. HRV increased only three weeks before exit, indicating that late activation of the sympathetic nervous system likely finalized restoration of euthermic metabolism. Interestingly, it was not until TA reached the presumed lower critical temperature, likely indicating that the bears were seeking thermoneutrality, that they exited the den. CONCLUSIONS: We conclude that brown bear hibernation was initiated primarily by environmental cues, but terminated by physiological cues.

Consequences of a demographic bottleneck on genetic structure and variation in the Scandinavian brown bear.

The Scandinavian brown bear went through a major decline in population size approximately 100 years ago, due to intense hunting. After being protected, the population subsequently recovered and today numbers in the thousands. The genetic diversity in the contemporary population has been investigated in considerable detail, and it has been shown that the population consists of several subpopulations that display relatively high levels of genetic variation. However, previous studies have been unable to resolve the degree to which the demographic bottleneck impacted the contemporary genetic structure and diversity. In this study, we used mitochondrial and microsatellite DNA markers from pre- and postbottleneck Scandinavian brown bear samples to investigate the effect of the bottleneck. Simulation and multivariate analysis suggested the same genetic structure for the historical and modern samples, which are clustered into three subpopulations in southern, central and northern Scandinavia. However, the southern subpopulation appears to have gone through a marked change in allele frequencies. When comparing the mitochondrial DNA diversity in the whole population, we found a major decline in haplotype numbers across the bottleneck. However, the loss of autosomal genetic diversity was less pronounced, although a significant decline in allelic richness was observed in the southern subpopulation. Approximate Bayesian computations provided clear support for a decline in effective population size during the bottleneck, in both the southern and northern subpopulations. These results have implications for the future management of the Scandinavian brown bear because they indicate a recent loss in genetic diversity and also that the current genetic structure may have been caused by historical ecological processes rather than recent anthropogenic persecution.

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.

A new individual-based spatial approach for identifying genetic discontinuities in natural populations.

The population concept is central in evolutionary and conservation biology, but identifying the boundaries of natural populations is often challenging. Here, we present a new approach for assessing spatial genetic structure without the a priori assumptions on the locations of populations made by adopting an individual-centred approach. Our method is based on assignment tests applied in a moving window over an extensively sampled study area. For each individual, a spatially explicit probability surface is constructed, showing the estimated probability of finding its multilocus genotype across the landscape, and identifying putative migrants. Population boundaries are localized by estimating the mean slope of these probability surfaces over all individuals to identify areas with genetic discontinuities. The significance of the genetic discontinuities is assessed by permutation tests. This new approach has the potential to reveal cryptic population structure and to improve our ability to understand gene flow dynamics across landscapes. We illustrate our approach by simulations and by analysing two empirical datasets: microsatellite data of Ursus arctos in Scandinavia, and amplified fragment length polymorphism (AFLP) data of Rhododendron ferrugineum in the Alps.

Assumed and inferred spatial structure of populations: the Scandinavian brown bears revisited.

We reanalysed the spatial structure of the Scandinavian brown bear (Ursus arctos) population based on multilocus genotypes. We used data from a former study that had presumed a priori a specific population subdivision based on four subpopulations. Using two independent methods (neighbour-joining trees and Bayesian assignment tests), we analysed the data without any prior presumption about the spatial structure. A subdivision of the population into three subpopulations emerged from our study. The genetic pattern of these subpopulations matched the three geographical clusters of individuals present in the population. We recommend considering the Scandinavian brown bear population as consisting of three (instead of four) subpopulations. Our results underline the importance of determining genetic structure from the data, without presupposing a structure, even when there seems to be good reason to do so.

Recolonizing carnivores and naïve prey: conservation lessons from Pleistocene extinctions.

The current extinction of many of Earth's large terrestrial carnivores has left some extant prey species lacking knowledge about contemporary predators, a situation roughly parallel to that 10,000 to 50,000 years ago, when naive animals first encountered colonizing human hunters. Along present-day carnivore recolonization fronts, brown (also called grizzly) bears killed predator-naive adult moose at disproportionately high rates in Scandinavia, and moose mothers who lost juveniles to recolonizing wolves in North America's Yellowstone region developed hypersensitivity to wolf howls. Although prey that had been unfamiliar with dangerous predators for as few as 50 to 130 years were highly vulnerable to initial encounters, behavioral adjustments to reduce predation transpired within a single generation. The fact that at least one prey species quickly learns to be wary of restored carnivores should negate fears about localized prey extinction.

Nuclear DNA microsatellite analysis of genetic diversity and gene flow in the Scandinavian brown bear (Ursus arctos).

In the 1930s, the Scandinavian brown bear was close to extinction due to vigorous extermination programmes in Norway and Sweden. Increased protection of the brown bear in Scandinavia has resulted in the recovery of four subpopulations, which currently contain close to 1000 individuals. Effective conservation and management of the Scandinavian brown bear requires knowledge of the current levels of genetic diversity and gene flow among the four subpopulations. Earlier studies of mitochondrial DNA (mtDNA) diversity revealed extremely low levels of genetic variation, and population structure that grouped the three northern subpopulations in one genetic clade and the southernmost subpopulation in a second highly divergent clade. In this study, we extended the analysis of genetic diversity and gene flow in the Scandinavian brown bear using data from 19 nuclear DNA microsatellite loci. Results from the nuclear loci were strikingly different than the mtDNA results. Genetic diversity levels in the four subpopulations were equivalent to diversity levels in nonbottlenecked populations from North America, and significantly higher than levels in other bottlenecked and isolated brown bear populations. Gene flow levels between subpopulations ranged from low to moderate and were correlated with geographical distance. The substantial difference in results obtained using mtDNA and nuclear DNA markers stresses the importance of collecting data from both types of genetic markers before interpreting data and making recommendations for the conservation and management of natural populations. Based on the results from the mtDNA and nuclear DNA data sets, we propose one evolutionarily significant unit and four management units for the brown bear in Scandinavia.

The effect of matrix on the occurrence of hazel grouse (Bonasa bonasia) in isolated habitat fragments.

The aim of this study was to determine the effect of matrix on the occurrence of hazel grouse (Bonasa bonasia) in habitat fragments. The study was conducted in two kinds of landscape: (1) an agricultural landscape, where the censused forest habitat fragments were surrounded by farmland, and (2) in an intensively managed forested landscape, where the censused habitat fragments were surrounded by nonhabitat coniferous forest. Occupied and unoccupied habitat fragments in the agricultural landscape differed significantly in distance to the nearest suitable continuous habitat, with hazel grouse occurring only in habitat fragments closer than 100 m from continuous forest. In the intensively managed forest landscape, the effect of isolation was less evident, but there might be a threshold around 2 km. Effects of isolation occurred over much shorter distances when the surrounding habitats consisted of farmland than when it was forested habitats. The size of the habitat fragments was important in both landscapes, with larger habitat fragments more often containing hazel grouse.