Andreas Zedrosser

Inversely density-dependent natal dispersal in brown bears Ursus arctos

There is considerable controversy in the literature about the presence of density dependence in dispersal. In this study, we exploit a data series from a long-term study (>18xa0years) on radio-marked brown bears (Ursus arctos L.) in two study areas in Scandinavia to investigate how individual-based densities influence the probability of natal dispersal and natal dispersal distances. Cumulatively, 32% and 46% of the females and 81% and 92% of the males dispersed before reaching 5xa0years of age in the northern and southern study area, respectively. Density had a negative effect on both the probability of dispersal and dispersal distances for the dispersing animals, when controlling for study area, sex and age, making this the first study to show that natal dispersal probability and distances are inversely density dependent in a large carnivore. We suggest that female–female competition for space caused females in higher density areas to settle closer to their natal area. For males, however, merging of demes, resulting in decreased relatedness and increased heterozygosity in an expanding population, might be the reason for shorter dispersal distances in males living at higher densities. This has been hypothesised for small mammals. The high proportion of dispersing female brown bears in Scandinavian compared with North American studies might be due to lower densities in Scandinavia and recent population expansion, with unoccupied areas available at the edges of the population. The longer dispersal distances in female Scandinavian brown bears suggest less social constraints on movements than for North American females. The longer dispersal distances by Scandinavian males may be due to increased searching for potential mates in peripheral areas with lower densities of females. These results, in addition to results of other brown bear studies, suggest that brown bears might be more territorial than previously thought, and that density is regulated by social interactions.

Should I stay or should I go? Natal dispersal in the brown bear

We studied the causes of natal dispersal of male and female brown bears, Ursus arctos, in two study areas in Sweden. Males had a higher dispersal probability (94%) than females (41%). For males, we found no difference in dispersal probability or mean age of dispersal between the study areas, in spite of differences in population density and sex ratio. Male–male competition did not seem to influence subadult male dispersal probability significantly. These results support the inbreeding avoidance hypothesis as the cause of male natal dispersal. For females, dispersal probability decreased with increasing maternal age and decreased with increasing body size, and an interaction between maternal age and body size suggested that the importance of body size decreased with increasing maternal age. Nondispersing females were closer to their mother than their dispersing sibling sisters were in the period between weaning and dispersal. Female littermates seemed to compete for philopatry, suggesting that a dominance hierarchy among female littermates based on body size may cause the subdominant sister to disperse. If juvenile females are born into matrilineal assemblages, surrounded mostly by related females, the competition for philopatry may not be as severe as when they are born into an area surrounded by mostly nonkin females. This hypothesis is supported by the decreasing importance of body size for dispersal with increasing maternal age. We suggest that natal dispersal in juvenile female brown bears can be explained by the resident fitness hypothesis.

POPULATION DENSITY AND FOOD CONDITIONS DETERMINE ADULT FEMALE BODY SIZE IN BROWN BEARS

Abstract We investigated growth and determinants of adult female body size in brown bears (Ursus arctos) in 2 study areas in Sweden. Scandinavian female brown bears reached 90% of their asymptotic size at 4.1–4.7 years. Four factors were considered in our analysis of the determinants of adult female size: annual food conditions, population density, multilocus heterozygosity, and yearling body size. Our results suggest that individual body size variation of female brown bears is negatively related to density-dependent factors and positively correlated to density-independent fluctuations in the environment. Density-dependent factors may operate by increasing competition for food, resulting in a decrease in body size. Food resources for brown bears in Sweden fluctuate annually in the boreal forest and influence individual body size. Multilocus heterozygosity and yearling body size were not important in explaining adult size, because initially smaller individuals show compensatory growth when experiencing good food conditions.

The dilemma of female mate selection in the brown bear, a species with sexually selected infanticide

Because of differential investment in gametes between sexes, females tend to be the more selective sex. Based on this concept, we investigate mate selection in a large carnivore: the brown bear (Ursus arctos). We hypothesize that, in this species with sexually selected infanticide (SSI), females may be faced with a dilemma: either select a high-quality partner based on phenotypic criteria, as suggested by theories of mate choice, or rather mate with future potentially infanticidal males as a counter-strategy to SSI. We evaluated which male characteristics were important in paternity assignment. Among males available in the vicinity of the females, the largest, most heterozygous and less inbred and also the geographically closest males were more often the fathers of the females next litter. We suggest that female brown bears may select the closest males as a counter-strategy to infanticide and exercise a post-copulatory cryptic choice, based on physical attributes, such as a large body size, reflecting male genetic quality. However, male–male competition either in the form of fighting before copulation or during the post-copulatory phase, in the form of sperm competition, cannot entirely be ruled out.

Archiving primary data: solutions for long-term studies

The recent trend for journals to require open access to primary data included in publications has been embraced by many biologists, but has caused apprehension amongst researchers engaged in long-term ecological and evolutionary studies. A worldwide survey of 73 principal investigators (Pls) with long-term studies revealed positive attitudes towards sharing data with the agreement or involvement of the PI, and 93% of PIs have historically shared data. Only 8% were in favor of uncontrolled, open access to primary data while 63% expressed serious concern. We present here their viewpoint on an issue that can have non-trivial scientific consequences. We discuss potential costs of public data archiving and provide possible solutions to meet the needs of journals and researchers.

Male reproductive strategy explains spatiotemporal segregation in brown bears

1. Spatiotemporal segregation is often explained by the risk for offspring predation or by differences in physiology, predation risk vulnerability or competitive abilities related to size dimorphism. 2. Most large carnivores are size dimorphic and offspring predation is often intraspecific and related to nonparental infanticide (NPI). NPI can be a foraging strategy, a strategy to reduce competition, or a male reproductive strategy. Spatiotemporal segregation is widespread among large carnivores, but its nature remains poorly understood. 3. We evaluated three hypotheses to explain spatiotemporal segregation in the brown bear, a size-dimorphic large carnivore in which NPI is common; the ‘NPI – foraging/competition hypothesis, i.e. NPI as a foraging strategy or a strategy to reduce competition, the ‘NPI – sexual selection hypothesis’, i.e. infanticide as a male reproductive strategy and the ‘body size hypothesis’, i.e. body-size-related differences in physiology, predation risk vulnerability or competitive ability causes spatiotemporal segregation. To test these hypotheses, we quantified spatiotemporal segregation among adult males, lone adult females and females with cubs-of-the-year, based on GPS-relocation data (2006–2010) and resource selection functions in a Scandinavian population. 4. We found that spatiotemporal segregation was strongest between females with cubs-of-the-year and adult males during the mating season. During the mating season, females with cubs-of-the-year selected their resources, in contrast to adult males, in less rugged landscapes in relative close proximity to certain human-related variables, and in more open habitat types. After the mating season, females with cubs-of-the-year markedly shifted their resource selection towards a pattern more similar to that of their conspecifics. No strong spatiotemporal segregation was apparent between females with cubs-of-the-year and conspecifics during the mating and the postmating season. 5. The ‘NPI – sexual selection hypothesis’ best explained spatiotemporal segregation in our study system. We suggest that females with cubs-of-the-year alter their resource selection to avoid infanticidal males. In species exhibiting NPI as a male reproductive strategy, female avoidance of infanticidal males is probably more common than observed or reported, and may come with a fitness cost if females trade safety for optimal resources.

The effects of primiparity on reproductive performance in the brown bear

We studied the effects of primiparity on litter size, offspring size, and cub loss in brown bears (Ursus arctos) in two study areas (north, south) in Sweden from 1987 to 2006. Sexually selected infanticide (SSI) has been suggested previously as a mortality factor in our study populations. Females in the south became primiparous earlier than females in the north. Primiparous females had significantly smaller litters of cubs than multiparous females. We found no evidence that primiparity was costly in terms of the interlitter interval. Primiparous mothers had a higher probability of cub loss than multiparous mothers. The probability of cub loss was analyzed separately for the pre-mating and the mating season. The probability of cub loss by primiparous females in the pre-mating season increased with both increasing population density and deteriorating food conditions, whereas the probability of cub loss during the mating season decreased with increasing age of primiparity and increased with male turnover (a variable predicting SSI). The temporal patterns of cub loss by primiparous females suggested that the critical times for reproductive success by primiparous females were the pre-mating season (from birth to shortly after leaving the den) and the mating season. Cub loss in these periods was independent and caused by different factors. Cub loss before the mating season seemed to be most influenced by food conditions, whereas that during the mating season appeared to be caused by SSI.

OpinionArchiving Primary Data: Solutions for Long-Term Studies

The recent trend for journals to require open access to primary data included in publications has been embraced by many biologists, but has caused apprehension amongst researchers engaged in long-term ecological and evolutionary studies. A worldwide survey of 73 principal investigators (Pls) with long-term studies revealed positive attitudes towards sharing data with the agreement or involvement of the PI, and 93% of PIs have historically shared data. Only 8% were in favor of uncontrolled, open access to primary data while 63% expressed serious concern. We present here their viewpoint on an issue that can have non-trivial scientific consequences. We discuss potential costs of public data archiving and provide possible solutions to meet the needs of journals and researchers.

Hunting Patterns, Ban on Baiting, and Harvest Demographics of Brown Bears in Sweden

Abstract We analyzed harvest data to describe hunting patterns and harvest demography of brown bears (Ursus arctos) killed in 3 geographic regions in Sweden during 1981–2004. In addition, we investigated the effects of a ban on baiting, instituted in 2001, and 2 major changes in the quota system: a switch to sex-specific quotas in 1992 and a return to total quotas in 1999. Brown bears (n = 887) were harvested specifically by bear hunters and incidentally by moose (Alces alces) hunters. Both hunter categories harvested bears 1) using dogs (37%), 2) by still hunting (30%), 3) with the use of bait (18%), and 4) by stalking (16%). The proportion of bears killed with different harvest methods varied among regions and between bear- and moose-oriented hunters. We found differences between male (52%) and female bears (48%) with respect to the variables that explained age. Moose-oriented hunters using still hunting harvested the youngest male bears. Bears harvested during the first management period (1981–1991) were older and had greater odds of being male than during the subsequent period. It appears that hunters harvesting bears in Sweden are less selective than their North American counterparts, possibly due to differences in the hunting system. When comparing the 4 years immediately prior to the ban on baiting with the 4 years following the ban, we found no differences in average age of harvested bears, sex ratio, or proportion of bears killed with stalking, still hunting, and hunting with dogs, suggesting that the ban on baiting in Sweden had no immediate effect on patterns of brown bear harvest demography and remaining hunting methods. As the demographic and evolutionary side effects of selective harvesting receive growing attention, wildlife managers should be aware that differences in harvest systems between jurisdictions may cause qualitative and quantitative differences in harvest biases.

Noninvasive genetic studies of brown bears using power poles

One difficulty in the conservation of endangered wildlife is the lack of reliable information on its status. This lack of knowledge can often be attributed to financial and logistic constraints as well as the lack of trained personnel to collect data. We test a simple method to study bears in the southern Balkans by inspecting power poles, which are used by bears for marking and rubbing purposes. We created a network of barbed-wire fitted poles for the collection of hair samples, evenly distributed throughout six study areas. During 87 sampling sessions in the main study area, we collected 191 samples and identified six microsatellite loci that were variable enough for individual bear identification. The most and best-quality hair samples were collected during the mating period, and DNA was most successfully extracted from samples remaining <4xa0weeks in the field. In the six study areas, we identified 47 bears. An advantage of using power poles for hair sampling is their availability and accessibility; no bait is required, and the network can be easily set up. A drawback may be an unequal capture probability of sex and age classes of bears. Despite this limitation, using power poles proved to be a simple and cheap method for the noninvasive genetic study of bears that did not require any prior knowledge on habitat use and activity patterns. The method is suitable for large-scale surveys to estimate distribution and relative densities of bears and could also be applied for studying other species.