Body mass is associated with hibernation length, body temperature, and heart rate in free-ranging brown bears.

BACKGROUND: Despite centuries of research, debate remains on the scaling of metabolic rate to mass especially for intraspecific cases. The high variation of body mass within brown bears presents a unique opportunity to study the intraspecific effects of body mass on physiological variables. The amplitude of metabolic rate reduction in hibernators is dependent on body mass of the species. Small hibernators have high metabolic rates when euthermic but experience a drastic decrease in body temperature during torpor, which is necessary to reach a very low metabolic rate. Conversely, large hibernators, such as the brown bear (Ursus arctos), show a moderate decrease in temperature during hibernation, thought to be related to the bear's large size. We studied body mass, abdominal body temperature, heart rate, and accelerometer-derived activity from 63 free-ranging brown bears (1-15 years old, 15-233 kg). We tested for relationships between body mass and body temperature, heart rate, and hibernation duration. RESULTS: The smallest individuals maintained lower body temperatures during hibernation, hibernated longer, and ended hibernation later than large bears. Unlike body temperature, winter heart rates were not associated with body mass. In summer, the opposite pattern was found, with smaller individuals having higher body temperature and daytime heart rates. Body mass was associated with body temperature in the winter hypometabolic state, even in a large hibernating mammal. Smaller bears, which are known to have higher thermal conductance, reached lower body temperatures during hibernation. During summer, smaller bears had higher body temperatures and daytime heart rates, a phenomenon not previously documented within a single mammalian species. CONCLUSION: We conclude that the smallest bears hibernated more deeply and longer than large bears, likely from a combined effect of basic thermodynamics, the higher need for energy savings, and a lower cost of warming up a smaller body.

Disentangling direct and indirect determinants of the duration of maternal care in brown bears: Environmental context matters.

The duration of maternal care, an important life-history trait affecting population dynamics, varies greatly within species. Yet, our understanding of its predictors is limited, mostly correlative and subject to misinterpretations, due to difficulties to disentangle the role of maternal- and offspring-related characteristics. We conducted path analysis on a dataset including 217 brown bear litters captured over a 29-year period in two populations in Sweden ('North' and 'South') facing contrasting environmental conditions to identify and quantify the causes of variation in the duration of maternal care (1.5 or 2.5 years). We showed that the causal determinants of the duration of maternal care were context-dependent. Contrary to their expected central role in the determination of the duration of maternal care, yearling mass and its direct determinants (i.e. litter size and maternal mass) were only important in the North population, where environmental conditions are harsher and the cost of extended maternal care presumably higher. In the South, the duration of maternal care was not caused by yearling mass nor any maternal or litter characteristics. Extension of maternal care may thus result from factors independent from maternal and offspring condition in the South, such as an artificial hunting-induced selection for longer maternal care through the legal protection of family groups. Our results provide an important contribution to our very limited knowledge of the direct and indirect determinants of the duration of maternal care and highlight the importance of accounting for the environmental context when assessing maternal reproductive tactics.

MicroRNAs facilitate skeletal muscle maintenance and metabolic suppression in hibernating brown bears.

Hibernating brown bears, Ursus arctos, undergo extended periods of inactivity and yet these large hibernators are resilient to muscle disuse atrophy. Physiological characteristics associated with atrophy resistance in bear muscle have been examined (e.g., muscle mechanics, neural activity) but roles for molecular signaling/regulatory mechanisms in the resistance to muscle wasting in bears still require investigation. Using quantitative reverse transcription PCR (RT-qPCR), the present study characterized the responses of 36 microRNAs linked with development, metabolism, and regeneration of skeletal muscle, in the vastus lateralis of brown bears comparing winter hibernating and summer active animals. Relative levels of mRNA of selected genes (mef2a, pax7, id2, prkaa1, and mstn) implicated upstream and downstream of the microRNAs were examined. Results indicated that hibernation elicited a myogenic microRNA, or "myomiR", response via MEF2A-mediated signaling. Upregulation of MEF2A-controlled miR-1 and miR-206 and respective downregulation of pax7 and id2 mRNA are suggestive of responses that promote skeletal muscle maintenance. Increased levels of metabolic microRNAs, such as miR-27, miR-29, and miR-33, may facilitate metabolic suppression during hibernation via mechanisms that decrease glucose uptake and fatty acid oxidation. This study identified myomiR-mediated mechanisms for the promotion of muscle regeneration, suppression of ubiquitin ligases, and resistance to muscle atrophy during hibernation mediated by observed increases in miR-206, miR-221, miR-31, miR-23a, and miR-29b. This was further supported by the downregulation of myomiRs associated with a muscle injury and inflammation (miR-199a and miR-223) during hibernation. The present study provides evidence of myomiR-mediated signaling pathways that are activated during hibernation to maintain skeletal muscle functionality in brown bears.

Specific shifts in the endocannabinoid system in hibernating brown bears.

In small hibernators, global downregulation of the endocannabinoid system (ECS), which is involved in modulating neuronal signaling, feeding behavior, energy metabolism, and circannual rhythms, has been reported to possibly drive physiological adaptation to the hibernating state. In hibernating brown bears (Ursus arctos), we hypothesized that beyond an overall suppression of the ECS, seasonal shift in endocannabinoids compounds could be linked to bear's peculiar features that include hibernation without arousal episodes and capacity to react to external disturbance. We explored circulating lipids in serum and the ECS in plasma and metabolically active tissues in free-ranging subadult Scandinavian brown bears when both active and hibernating. In winter bear serum, in addition to a 2-fold increase in total fatty acid concentration, we found significant changes in relative proportions of circulating fatty acids, such as a 2-fold increase in docosahexaenoic acid C22:6 n-3 and a decrease in arachidonic acid C20:4 n-6. In adipose and muscle tissues of hibernating bears, we found significant lower concentrations of 2-arachidonoylglycerol (2-AG), a major ligand of cannabinoid receptors 1 (CB1) and 2 (CB2). Lower mRNA level for genes encoding CB1 and CB2 were also found in winter muscle and adipose tissue, respectively. The observed reduction in ECS tone may promote fatty acid mobilization from body fat stores, and favor carbohydrate metabolism in skeletal muscle of hibernating bears. Additionally, high circulating level of the endocannabinoid-like compound N-oleoylethanolamide (OEA) in winter could favor lipolysis and fatty acid oxidation in peripheral tissues. We also speculated on a role of OEA in the conservation of an anorexigenic signal and in the maintenance of torpor during hibernation, while sustaining the capacity of bears to sense stimuli from the environment.

Metabolic reprogramming involving glycolysis in the hibernating brown bear skeletal muscle.

BACKGROUND: In mammals, the hibernating state is characterized by biochemical adjustments, which include metabolic rate depression and a shift in the primary fuel oxidized from carbohydrates to lipids. A number of studies of hibernating species report an upregulation of the levels and/or activity of lipid oxidizing enzymes in muscles during torpor, with a concomitant downregulation for glycolytic enzymes. However, other studies provide contrasting data about the regulation of fuel utilization in skeletal muscles during hibernation. Bears hibernate with only moderate hypothermia but with a drop in metabolic rate down to ~ 25% of basal metabolism. To gain insights into how fuel metabolism is regulated in hibernating bear skeletal muscles, we examined the vastus lateralis proteome and other changes elicited in brown bears during hibernation. RESULTS: We show that bear muscle metabolic reorganization is in line with a suppression of ATP turnover. Regulation of muscle enzyme expression and activity, as well as of circulating metabolite profiles, highlighted a preference for lipid substrates during hibernation, although the data suggested that muscular lipid oxidation levels decreased due to metabolic rate depression. Our data also supported maintenance of muscle glycolysis that could be fuelled from liver gluconeogenesis and mobilization of muscle glycogen stores. During hibernation, our data also suggest that carbohydrate metabolism in bear muscle, as well as protein sparing, could be controlled, in part, by actions of n-3 polyunsaturated fatty acids like docosahexaenoic acid. CONCLUSIONS: Our work shows that molecular mechanisms in hibernating bear skeletal muscle, which appear consistent with a hypometabolic state, likely contribute to energy and protein savings. Maintenance of glycolysis could help to sustain muscle functionality for situations such as an unexpected exit from hibernation that would require a rapid increase in ATP production for muscle contraction. The molecular data we report here for skeletal muscles of bears hibernating at near normal body temperature represent a signature of muscle preservation despite atrophying conditions.

Sociodemographic factors modulate the spatial response of brown bears to vacancies created by hunting.

There is a growing recognition of the importance of indirect effects from hunting on wildlife populations, e.g. social and behavioural changes due to harvest, which occur after the initial offtake. Nonetheless, little is known about how the removal of members of a population influences the spatial configuration of the survivors. We studied how surviving brown bears (Ursus arctos) used former home ranges that had belonged to casualties of the annual bear hunting season in southcentral Sweden (2007-2015). We used resource selection functions to explore the effects of the casualty's and survivor's sex, age and their pairwise genetic relatedness, population density and hunting intensity on survivors' spatial responses to vacated home ranges. We tested the competitive release hypothesis, whereby survivors that increase their use of a killed bear's home range are presumed to have been released from intraspecific competition. We found strong support for this hypothesis, as survivors of the same sex as the casualty consistently increased their use of its vacant home range. Patterns were less pronounced or absent when the survivor and casualty were of opposite sex. Genetic relatedness between the survivor and the casualty emerged as the most important factor explaining increased use of vacated male home ranges by males, with a stronger response from survivors of lower relatedness. Relatedness was also important for females, but it did not influence use following removal; female survivors used home ranges of higher related female casualties more, both before and after death. Spatial responses by survivors were further influenced by bear age, population density and hunting intensity. We have shown that survivors exhibit a spatial response to vacated home ranges caused by hunting casualties, even in nonterritorial species such as the brown bear. This spatial reorganization can have unintended consequences for population dynamics and interfere with management goals. Altogether, our results underscore the need to better understand the short- and long-term indirect effects of hunting on animal social structure and their resulting distribution in space.

Seasonal changes in eicosanoid metabolism in the brown bear.

Polyunsaturated fatty acids (PUFAs) exert several important functions across organ systems. During winter, hibernators divert PUFAs from oxidation, retaining them in their tissues and membranes, to ensure proper body functions at low body temperature. PUFAs are also precursors of eicosanoids with pro- and anti-inflammatory properties. This study investigated seasonal changes in eicosanoid metabolism of free-ranging brown bears (Ursus arctos). By using a lipidomic approach, we assessed (1) levels of specific omega-3 and omega-6 fatty acids involved in the eicosanoid cascade and (2) concentrations of eicosanoids in skeletal muscle and blood plasma of winter hibernating and summer active bears. We observed significant seasonal changes in the specific omega-3 and omega-6 precursors. We also found significant seasonal alterations of eicosanoid levels in both tissues. Concentrations of pro-inflammatory eicosanoids, such as thromboxane B2, 5-hydroxyeicosatetraenoic acid (HETE), and 15-HETE and 18-HETE, were significantly lower in muscle and/or plasma of hibernating bears compared to summer-active animals. Further, plasma and muscle levels of 5,6-epoxyeicosatrienoic acid (EET), as well as muscle concentration of 8,9-EET, tended to be lower in bears during winter hibernation vs. summer. We also found lower plasma levels of anti-inflammatory eicosanoids, such as 15dPGJ2 and PGE3, in bears during winter hibernation. Despite of the limited changes in omega-3 and omega-6 precursors, plasma and muscle concentrations of the products of all pathways decreased significantly, or remained unchanged, independent of their pro- or anti-inflammatory properties. These findings suggest that hibernation in bears is associated with a depressed state of the eicosanoid cascade.

Biochemical Foundations of Health and Energy Conservation in Hibernating Free-ranging Subadult Brown Bear Ursus arctos.

Brown bears (Ursus arctos) hibernate for 5-7 months without eating, drinking, urinating, and defecating at a metabolic rate of only 25% of the summer activity rate. Nonetheless, they emerge healthy and alert in spring. We quantified the biochemical adaptations for hibernation by comparing the proteome, metabolome, and hematological features of blood from hibernating and active free-ranging subadult brown bears with a focus on conservation of health and energy. We found that total plasma protein concentration increased during hibernation, even though the concentrations of most individual plasma proteins decreased, as did the white blood cell types. Strikingly, antimicrobial defense proteins increased in concentration. Central functions in hibernation involving the coagulation response and protease inhibition, as well as lipid transport and metabolism, were upheld by increased levels of very few key or broad specificity proteins. The changes in coagulation factor levels matched the changes in activity measurements. A dramatic 45-fold increase in sex hormone-binding globulin levels during hibernation draws, for the first time, attention to its significant but unknown role in maintaining hibernation physiology. We propose that energy for the costly protein synthesis is reduced by three mechanisms as follows: (i) dehydration, which increases protein concentration without de novo synthesis; (ii) reduced protein degradation rates due to a 6 °C reduction in body temperature and decreased protease activity; and (iii) a marked redistribution of energy resources only increasing de novo synthesis of a few key proteins. The comprehensive global data identified novel biochemical strategies for bear adaptations to the extreme condition of hibernation and have implications for our understanding of physiology in general.

Competition between apex predators? Brown bears decrease wolf kill rate on two continents.

Trophic interactions are a fundamental topic in ecology, but we know little about how competition between apex predators affects predation, the mechanism driving top-down forcing in ecosystems. We used long-term datasets from Scandinavia (Europe) and Yellowstone National Park (North America) to evaluate how grey wolf (Canis lupus) kill rate was affected by a sympatric apex predator, the brown bear (Ursus arctos). We used kill interval (i.e. the number of days between consecutive ungulate kills) as a proxy of kill rate. Although brown bears can monopolize wolf kills, we found no support in either study system for the common assumption that they cause wolves to kill more often. On the contrary, our results showed the opposite effect. In Scandinavia, wolf packs sympatric with brown bears killed less often than allopatric packs during both spring (after bear den emergence) and summer. Similarly, the presence of bears at wolf-killed ungulates was associated with wolves killing less often during summer in Yellowstone. The consistency in results between the two systems suggests that brown bear presence actually reduces wolf kill rate. Our results suggest that the influence of predation on lower trophic levels may depend on the composition of predator communities.

Hunting promotes sexual conflict in brown bears.

The removal of individuals through hunting can destabilize social structure, potentially affecting population dynamics. Although previous studies have shown that hunting can indirectly reduce juvenile survival through increased sexually selected infanticide (SSI), very little is known about the spatiotemporal effects of male hunting on juvenile survival. Using detailed individual monitoring of a hunted population of brown bears (Ursus arctos) in Sweden (1991-2011), we assessed the spatiotemporal effect of male removal on cub survival. We modelled cub survival before, during and after the mating season. We used three proxies to evaluate spatial and temporal variation in male turnover; distance and timing of the closest male killed and number of males that died around a female's home range centre. Male removal decreased cub survival only during the mating season, as expected in seasonal breeders with SSI. Cub survival increased with distance to the closest male killed within the previous 1·5 years, and it was lower when the closest male killed was removed 1·5 instead of 0·5 year earlier. We did not detect an effect of the number of males killed. Our results support the hypothesis that social restructuring due to hunting can reduce recruitment and suggest that the distribution of the male deaths might be more important than the overall number of males that die. As the removal of individuals through hunting is typically not homogenously distributed across the landscape, spatial heterogeneity in hunting pressure may cause source-sink dynamics, with lower recruitment in areas of high human-induced mortality.

Staying cool or staying safe in a human-dominated landscape: which is more relevant for brown bears?

Pigeon et al. (2016) Staying cool in a changing landscape: the influence of maximum daily ambient temperature on grizzly bear habitat selection. Oecologia 181:1101. doi: 10.1007/s00442-016-3630-5 analyzed the effect of ambient temperature on the habitat selection of grizzly bears (Ursus arctos) in Alberta, Canada. They concluded that temperature played a significant role in bear habitat selection and that it was unlikely that human activity introduced biases to the habitat selection of bears. However, Pigeon et al. did not consider variables related to human activities in their analyses. They also misinterpreted previous research that has accounted for temperature in the habitat selection of brown bears. There is much literature published on the negative effects of human disturbance on wildlife in general and on bears in particular. Downplaying the role of human disturbance could have important negative consequences if, in fact, human disturbance were a more important factor than thermoregulation. Indeed, dismissing the importance of human influence, in the face of contradictory evidence, could tempt managers to disregard an important factor that is difficult and often unpopular to deal with in their conservation plans.

Can hunting data be used to estimate unbiased population parameters? A case study on brown bears.

Quantifying temporal changes in harvested populations is critical for applied and fundamental research. Unbiased data are required to detect true changes in phenotypic distribution or population size. Because of the difficulty of collecting detailed individual data from wild populations, data from hunting records are often used. Hunting records, however, may not represent a random sample of a population. We aimed to detect and quantify potential bias in hunting records. We compared data from a long-term monitoring project with hunting records of brown bears (Ursus arctos) in Sweden and investigated temporal trends (1996-2013) in the ratio of yearlings to adult females, yearling mass and adult female mass. Data from hunting records underestimated the decline in yearling and adult female mass over time, most likely owing to the legal protection of family groups from hunting, but reflected changes in the ratio of yearlings to adult females more reliably. Although hunting data can be reliable to approximate population abundance in some circumstances, hunting data can represent a biased sample of a population and should be used with caution in management and conservation decisions.

Quantifying consistent individual differences in habitat selection.

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 ß 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.

Temporal effects of hunting on foraging behavior of an apex predator: Do bears forego foraging when risk is high?

Avoiding predators most often entails a food cost. For the Scandinavian brown bear (Ursus arctos), the hunting season coincides with the period of hyperphagia. Hunting mortality risk is not uniformly distributed throughout the day, but peaks in the early morning hours. As bears must increase mass for winter survival, they should be sensitive to temporal allocation of antipredator responses to periods of highest risk. We expected bears to reduce foraging activity at the expense of food intake in the morning hours when risk was high, but not in the afternoon, when risk was low. We used fine-scale GPS-derived activity patterns during the 2 weeks before and after the onset of the annual bear hunting season. At locations of probable foraging, we assessed abundance and sugar content, of bilberry (Vaccinium myrtillus), the most important autumn food resource for bears in this area. Bears decreased their foraging activity in the morning hours of the hunting season. Likewise, they foraged less efficiently and on poorer quality berries in the morning. Neither of our foraging measures were affected by hunting in the afternoon foraging bout, indicating that bears did not allocate antipredator behavior to times of comparably lower risk. Bears effectively responded to variation in risk on the scale of hours. This entailed a measurable foraging cost. The additive effect of reduced foraging activity, reduced forage intake, and lower quality food may result in poorer body condition upon den entry and may ultimately reduce reproductive success.

The relative importance of direct and indirect effects of hunting mortality on the population dynamics of brown bears.

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 (λ = 0.975; 95% CI = 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.

Y chromosome haplotype distribution of brown bears (Ursus arctos) in Northern Europe provides insight into population history and recovery.

High-resolution, male-inherited Y-chromosomal markers are a useful tool for population genetic analyses of wildlife species, but to date have only been applied in this context to relatively few species besides humans. Using nine Y-chromosomal STRs and three Y-chromosomal single nucleotide polymorphism markers (Y-SNPs), we studied whether male gene flow was important for the recent recovery of the brown bear (Ursus arctos) in Northern Europe, where the species declined dramatically in numbers and geographical distribution during the last centuries but is expanding now. We found 36 haplotypes in 443 male extant brown bears from Sweden, Norway, Finland and northwestern Russia. In 14 individuals from southern Norway from 1780 to 1920, we found two Y chromosome haplotypes present in the extant population as well as four Y chromosome haplotypes not present among the modern samples. Our results suggested major differences in genetic connectivity, diversity and structure between the eastern and the western populations in Northern Europe. In the west, our results indicated that the recovered population originated from only four male lineages, displaying pronounced spatial structuring suggestive of large-scale population size increase under limited male gene flow within the western subpopulation. In the east, we found a contrasting pattern, with high haplotype diversity and admixture. This first population genetic analysis of male brown bears shows conclusively that male gene flow was not the main force of population recovery.

Den Entry Behavior in Scandinavian Brown Bears: Implications for Preventing Human Injuries.

Encounters between Scandinavian brown bears (Ursus arctos) and humans that result in human injuries and fatalities typically coincide with den entry in October and November, and commonly occur near a den. Our aim was to determine when bears arrive at their dens, identify potential predictors of this event, document behavior and activity associated with this period, and attempt to explain the increased risk of bear-caused human injuries in this period. We analyzed global positioning system (GPS) location and activity data from brown bears in south-central Sweden, using generalized linear mixed models, statistical process control, and activity analyses. Bears arrived at their den sites between 6 October and 1 December. Timing varied by reproductive category, bear age, and year. Half of all bears significantly reduced their activity before arriving at the den area: on average 2,169 m away from the den and 1.8 days before arrival. The other half reduced their activity after arriving at the den area. The latter bears took longer time to reach hibernation activity levels, but we did not find a difference in the start date of hibernation between the 2 groups. Bears also appeared to be sensitive to disturbance in this period, with higher den abandonment rates than later in winter, particularly for males and for bears that had not visited their den sites previously. Den entry occurred from October to December, with high variability and poor predictability of its timing. Therefore, restricting hunting or other recreation activities to reduce risk of injury by bears and disturbing bears probably would be both impractical and ineffective. Our findings can be used to educate hunters about bear behavior at this time of year. Many people associate dens with an increased risk of a bear responding aggressively to disturbance to defend its den, but our results indicate that other behavioral, and possibly physiological, changes in this period also may be involved. © 2014 The Authors. The Journal of Wildlife Management published by The Wildlife Society.

A "clearcut" case? Brown bear selection of coarse woody debris and carpenter ants on clearcuts.

Forest management alters habitat characteristics, resulting in various effects among and within species. It is crucial to understand how habitat alteration through forest management (e.g. clearcutting) affects animal populations, particularly with unknown future conditions (e.g. climate change). In Sweden, brown bears (Ursus arctos) forage on carpenter ants (Camponotus herculeanus) during summer, and may select for this food source within clearcuts. To assess carpenter ant occurrence and brown bear selection of carpenter ants, we sampled 6999 coarse woody debris (CWD) items within 1019 plots, of which 902 were within clearcuts (forests ⩽30 years of age) and 117 plots outside clearcuts (forests >30 years of age). We related various CWD and site characteristics to the presence or absence of carpenter ant galleries (nests) and bear foraging sign at three spatial scales: the CWD, plot, and clearcut scale. We tested whether both absolute and relative counts (the latter controlling for the number of CWD items) of galleries and bear sign in plots were higher inside or outside clearcuts. Absolute counts were higher inside than outside clearcuts for galleries (mean counts; inside: 1.8, outside: 0.8). CWD was also higher inside (mean: 6.8) than outside clearcuts (mean: 4.0). However, even after controlling for more CWD inside clearcuts, relative counts were higher inside than outside clearcuts for both galleries (mean counts; inside: 0.3, outside: 0.2) and bear sign (mean counts; inside: 0.03, outside: 0.01). Variables at the CWD scale best explained gallery and bear sign presence than variables at the plot or clearcut level, but bear selection was influenced by clearcut age. CWD circumference was important for both carpenter ant and bear sign presence. CWD hardness was most important for carpenter ant selection. However, the most important predictor for bear sign was the presence or absence of carpenter ant galleries. Bears had a high foraging "success" rate (⩾88%) in foraging CWD where galleries also occurred, which was assessed by summing CWD items with the concurrence of bear sign and galleries, divided by the sum of all CWD with bear sign. Clearcuts appeared to increase the occurrence of a relatively important summer food item, the carpenter ant, on Swedish managed forests for the brown bear. However, the potential benefit of this increase can only be determined from a better understanding of the seasonal and interannual variation of the availability and use of other important brown bear food items, berries (e.g. Vaccinium myrtillus and Empetrum spp.), as well as other primary needs for bears (e.g. secure habitat and denning habitat), within the landscape mosaic of managed forests.

Bears are simply voles writ large: social structure determines the mechanisms of intrinsic population regulation in mammals.

The literature reveals opposing views regarding the importance of intrinsic population regulation in mammals. Different models have been proposed; adding importance to contrasting life histories, body sizes and social interactions. Here we evaluate current theory based on results from two Scandinavian projects studying two ecologically different mammal species with contrasting body sizes and life history traits: the root vole Microtus oeconomus and the brown bear Ursus arctos. We emphasize four inter-linked behavioral aspects-territoriality, dispersal, social inhibition of breeding, and infanticide-that together form a density-dependent syndrome with potentially regulatory effects on population growth. We show that the two species are similar in all four behaviors and thus the overall regulatory syndrome. Females form matrilineal assemblages, female natal dispersal is negatively density dependent and breeding is suppressed in philopatric young females. In both species, male turnover due to extrinsic mortality agents cause infanticide with negative effects on population growth. The sex-biased and density-dependent dispersal patterns promote the formation of matrilineal clusters which, in turn, leads to reproductive suppression with potentially regulatory effects. Hence, we show that intrinsic population regulation interacting with extrinsic mortality agents may occur irrespective of taxon, life history and body size. Our review stresses the significance of a mechanistic approach to understanding population ecology. We also show that experimental model populations are useful to elucidate natural populations of other species with similar social systems. In particular, such experiments should be combined with methodical innovations that may unravel the effects of cryptic intrinsic mechanisms such as infanticide.

Low cardiac output as physiological phenomenon in hibernating, free-ranging Scandinavian brown bears (Ursus arctos) - an observational study.

BACKGROUND: Despite 5-7 months of physical inactivity during hibernation, brown bears (Ursus arctos) are able to cope with physiological conditions that would be detrimental to humans. During hibernation, the tissue metabolic demands fall to 25% of the active state. Our objective was to assess cardiac function associated with metabolic depression in the hibernating vs. active states in free-ranging Scandinavian brown bears. METHODS: We performed echocardiography on seven free-ranging brown bears in Dalarna, Sweden, anesthetized with medetomidine-zolazepam-tiletamine-ketamine during winter hibernation in February 2013 and with medetomidine-zolazepam-tiletamine during active state in June 2013. We measured cardiac output noninvasively using estimates of hemodynamics obtained by pulsed wave Doppler echocardiography and 2D imaging. Comparisons were made using paired T-tests. RESULTS: During hibernation, all hemodynamic indices were significantly decreased (hibernating vs. active state): mean heart rate was 26.0 (standard deviation (SD): 5.6) beats per min vs. 75.0 (SD: 17.1) per min (P=0.002), mean stroke volume 32.3 (SD: 5.2) ml vs. 47.1 (SD: 7.9) ml (P=0.008), mean cardiac output 0.86 (SD: 0.31) l/min vs. 3.54 (SD: 1.04) l/min (P=0.003), and mean cardiac index 0.63 (SD: 0.21) l/min/kg vs. 2.45 (SD: 0.52) l/min/ m2 (P<0.001). Spontaneous echo contrast was present in all cardiac chambers in all seven bears during hibernation, despite the absence of atrial arrhythmias and valvular disease. CONCLUSION: Free-ranging brown bears demonstrate hemodynamics comparable to humans during active state, whereas during hibernation, we documented extremely low-flow hemodynamics. Understanding these physiological changes in bears may help to gain insight into the mechanisms of cardiogenic shock and heart failure in humans.