The shared preference niche of sympatric Asiatic black bears and sun bears in a tropical forest mosaic

Background

Ecologically similar species often coexist by partitioning use of habitats or resources. Such partitioning can occur through divergent or shared niches. We investigated overlap in habitat use and spatial co-occurrence by sympatric Asiatic black bears and sun bears in three habitats in Thailand, and thereby assessed which niche model best accounts for their coexistence.

Methods/Principal Findings

We used density of species-specific signs to assess habitat use. Signs of both bear species occurred in all three habitats, and on >60% of sampling transects. Both species fed mostly on fruit; insect feeding signs were uncommon, and were mostly from sun bears. Significant differences in habitat use occurred only in montane forest, the habitat in which fruit was most abundant; incidence of black bear sign there was six times higher than that of sun bears. Habitat use was similar between the two species in the other habitats, which comprised 85% of the area. Of 10 habitat attributes examined, fruiting tree density was the best predictor of occurrence for both species. Models that included interspecific competition (fresh foraging activity of the other species) were less supported than the top models without competition.

Conclusions/Significance

Bear species co-occurrence at both coarse and fine spatial scales and use of the same resources (fruit trees) indicated common niche preferences. However, their habitat use differed in ways expected from their physical differences: larger black bears dominated in the most fruit-rich habitat, and smaller sun bears used less-preferred insects. These results indicate broadly overlapping fundamental niches combined with asymmetric competition—features consistent with the concept of shared preference niches. This model of the niche has received little attention in ecology, but appears to be relatively common in nature.     

Dietary adjustability of grizzly bears and American black bears in Yellowstone National Park

Grizzly bears (Ursus arctos) and American black bears (U. americanus) are sympatric in much of Yellowstone National Park. Three primary bear foods, cutthroat trout (Oncorhynchus clarki), whitebark pine (Pinus albicaulis) nuts, and elk (Cervus elaphus), have declined in recent years. Because park managers and the public are concerned about the impact created by reductions in these foods, we quantified bear diets to determine how bears living near Yellowstone Lake are adjusting. We estimated diets using: 1) stable isotope and mercury analyses of hair samples collected from captured bears and from hair collection sites established along cutthroat trout spawning streams and 2) visits to recent locations occupied by bears wearing Global Positioning System collars to identify signs of feeding behavior and to collect scats for macroscopic identification of residues. Approximately 45 ± 22% ( ± SD) of the assimilated nitrogen consumed by male grizzly bears, 38 ± 20% by female grizzly bears, and 23 ± 7% by male and female black bears came from animal matter. These assimilated dietary proportions for female grizzly bears were the same as 10 years earlier in the Lake area and 30 years earlier in the Greater Yellowstone Ecosystem. However, the proportion of meat in the assimilated diet of male grizzly bears decreased over both time frames. The estimated biomass of cutthroat trout consumed by grizzly bears and black bears declined 70% and 95%, respectively, in the decade between 1997–2000 and 2007–2009. Grizzly bears killed an elk calf every 4.3 ± 2.7 days and black bears every 8.0 ± 4.0 days during June. Elk accounted for 84% of all ungulates consumed by both bear species. Whitebark pine nuts continue to be a primary food source for both grizzly bears and black bears when abundant, but are replaced by false-truffles (Rhizopogon spp.) in the diets of female grizzly bears and black bears when nut crops are minimal. Thus, both grizzly bears and black bears continue to adjust to changing resources, with larger grizzly bears continuing to occupy a more carnivorous niche than the smaller, more herbivorous black bear. © 2012 The Wildlife Society.     

Gene flow between insular, coastal and interior populations of brown bears in Alaska

The brown bears of coastal Alaska have been recently regarded as comprising from one to three distinct genetic groups. We sampled brown bears from each of the regions for which hypotheses of genetic uniqueness have been made, including the bears of the Kodiak Archipelago and the bears of Admiralty, Baranof and Chichagof (ABC) Islands in southeast Alaska. These samples were analysed with a suite of nuclear microsatellite markers. The 'big brown bears' of coastal Alaska were found to be part of the continuous continental distribution of brown bears, and not genetically isolated from the physically smaller 'grizzly bears' of the interior. By contrast, Kodiak brown bears appear to have experienced little or no genetic exchange with continental populations in recent generations. The bears of the ABC Islands, which have previously been shown to undergo little or no female-mediated gene flow with mainland populations, were found not to be genetically isolated from mainland bears. The data from the four insular populations indicate that female and male dispersal can be reduced or eliminated by water barriers of 2–4 km and 7km in width, respectively.     

Denning in brown bears

1. Hibernation represents an adaptation for coping with unfavorable environmental conditions. For brown bears Ursus arctos, hibernation is a critical period as pronounced temporal reductions in several physiological functions occur.
2. Here, we review the three main aspects of brown bear denning: (1) den chronology, (2) den characteristics, and (3) hibernation physiology in order to identify (a) proximate and ultimate factors of hibernation as well as (b) research gaps and conservation priorities.
3. Den chronology, which varies by sex and reproductive status, depends on environmental factors, such as snow, temperature, food availability, and den altitude. Significant variation in hibernation across latitudes occurs for both den entry and exit.
4. The choice of a den and its surroundings may affect individual fitness, for example, loss of offspring and excessive energy consumption. Den selection is the result of broad‐ and fine‐scale habitat selection, mainly linked to den insulation, remoteness, and availability of food in the surroundings of the den location.
5. Hibernation is a metabolic challenge for the brown bears, in which a series of physiological adaptations in tissues and organs enable survival under nutritional deprivation, maintain high levels of lipids, preserve muscle, and bone and prevent cardiovascular pathologies such as atherosclerosis.

It is important to understand: (a) proximate and ultimate factors in denning behavior and the difference between actual drivers of hibernation (i.e., factors to which bears directly respond) and their correlates; (b) how changes in climatic factors might affect the ability of bears to face global climate change and the human‐mediated changes in food availability; (c) hyperphagia (period in which brown bears accumulate fat reserves), predenning and denning periods, including for those populations in which bears do not hibernate every year; and (d) how to approach the study of bear denning merging insights from different perspectives, that is, physiology, ecology, and behavior.     

Mercury in polar bears from Alaska

Alaskan polar bear (Ursus maritimus) muscle and liver samples collected in 1972 were analyzed for total mercury. Bears north of Alaska had more mercury than bears west of Alaska. The only difference between young and adult animals was in the northern area where adults had more mercury in liver tissue than young animals. Levels were probably not high enough to be a serious threat to bears.     

Early primiparity in brown bears

We documented 2 cases of unusually early primiparity in brown bearsUrsus arctos Linneaus, 1758 in an introduced population in central Austria. Two females gave birth at the age of 3 years.     

Shape complexity of space used by American black bears influenced by sex and intensity of use

Disentangling the complexities that influence animal space use poses substantial challenges based on decision trade-offs and constraints imposed on animals. Optimal decisions suggest that the spatial complexity of home-range shapes should be inversely related to energy conservation and fitness. Hence, the most beneficial shape should be the circle. We evaluated whether shape complexity (i.e., an index of circularity) of home ranges was influenced by two extrinsic (spatial heterogeneity, preferred habitat [i.e., deciduous forest]), and three intrinsic (sex, season [breeding, non-breeding], intensity of use) factors, with intensity of use indexed as contours containing core and peripheral areas. We estimated utilization distributions of 39 radiomarked adult American black bears (Ursus americanus), a habitat generalist, using fixed-kernel techniques and estimated 50% (core area) and 95% (peripheral area) contours. We fit a set of 47 models using linear modeling and ranked models using small-sample Akaike Information Criterion. Coefficients for the best model were the intrinsic factors intensity of use (reference category = core; 0.118; 95% CL = 0.064–0.173), sex (reference = female; 0.105; 95% CL = 0.043–0.167), and intercept (0.229; 95% CL = 0.186–0.272). Shape complexity was less for core areas than peripheral areas and less for females than males. Considering complex resource selection patterns within a fragmented landscape, both sexes appeared to use energy-maximizing strategies, although the increase in shape complexity for males may be an allometric relationship based on size dimorphism. Our approach supported the phenomenon of optimality as manifested through home-range shape complexity, but we suggest that assessment of this phenomenon for habitat specialists may yield different results, including the potential importance of intrinsic factors based on more restrictive limiting factors.     

Sex-biased natal dispersal and inbreeding avoidance in American black bears as revealed by spatial genetic analyses

We tested the hypothesis that sex-biased natal dispersal reduces close inbreeding in American black bears, a solitary species that exhibits nearly complete male dispersal and female philopatry. Using microsatellite DNA and spatial data from reproductively mature bears (>or= 4 years old), we examined the spatial genetic structure of two distinct populations in New Mexico from 1993 to 2000. As predicted, relatedness (r) and the frequency of close relationships (parent-offspring or full siblings) decreased with distance among female dyads, but little change was observed among male or opposite-sex dyads. Neighbouring females were more closely related than neighbouring males. The potential for inbreeding was low. Most opposite-sex pairs that lived sufficiently close to facilitate mating were unrelated, and few were close relatives. We found no evidence that bears actively avoided inbreeding in their selection of mates from this nearby pool, as mean r and relationship frequencies did not differ between potential and actual mating pairs (determined by parentage analysis). These basic patterns were apparent in both study areas despite a nearly two-fold difference in density. However, the sex bias in dispersal was less pronounced in the lower-density area, based on proportions of bears with male and female relatives residing nearby. This result suggests that male bears may respond to reduced competition by decreasing their rate or distance of dispersal. Evidence supports the hypothesis that inbreeding avoidance is achieved by means of male-biased dispersal but also indicates that competition (for mates or resources) modifies dispersal patterns.     

Phylogeography of mitochondrial DNA variation in brown bears and polar bears

We analyzed 286 nucleotides of the middle portion of the mitochondrial cytochrome b gene of 61 brown bears from three locations in Alaska and 55 polar bears from Arctic Canada and Arctic Siberia to test our earlier observations of paraphyly between polar bears and brown bears as well as to test the extreme uniqueness of mitochondrial DNA types of brown bears on Admiralty, Baranof, and Chichagof (ABC) islands of southeastern Alaska. We also investigated the phylogeography of brown bears of Alaska's Kenai Peninsula in relation to other Alaskan brown bears because the former are being threatened by increased human development. We predicted that: (1) mtDNA paraphyly between brown bears and polar bears would be upheld, (2) the mtDNA uniqueness of brown bears of the ABC islands would be upheld, and (3) brown bears of the Kenai Peninsula would belong to either clade II or clade III of brown bears of our earlier studies of mtDNA. All of our predictions were upheld through the analysis of these additional samples.     

Static habitat partitioning and dynamic selection by sympatric young Atlantic salmon and brown trout in south-west England streams

Direct underwater observation of micro‐habitat use by 1838 young Atlantic salmon Salmo salar [mean L_T 7·9 ± 3.1(s.d.) cm, range 3·19] and 1227 brown trout Salmo trutta (L_T 10·9 ± 5·0 cm, range 3·56) showed both species were selective in habitat use, with differences between species and fish size. Atlantic salmon and brown trout selected relatively narrow ranges for the two micro‐habitat variables snout water velocity and height above bottom, but with differences between size‐classes. The smaller fishes <7 cm held positions in slower water closer to the bottom. On a larger scale, the Atlantic salmon more often used shallower stream areas, compared with brown trout. The larger parr preferred the deeper stream areas. Atlantic salmon used higher and slightly more variable mean water velocities than brown trout. Substrata used by the two species were similar. Finer substrata, although variable, were selected at the snout position, and differences were pronounced between size‐classes. On a meso‐habitat scale, brown trout were more frequently observed in slow pool‐glide habitats, while young Atlantic salmon favoured the faster high‐gradient meso‐habitats. Small juveniles <7 cm of both species were observed most frequently in riffle‐chute habitats. Atlantic salmon and brown trout segregated with respect to use of habitat, but considerable niche overlap between species indicated competitive interactions. In particular, for small fishes <7 cm of the two species, there was almost complete niche overlap for use of water depth, while they segregated with respect to water velocity. Habitat suitability indices developed for both species for mean water velocity and water depth, tended to have their optimum at lower values compared with previous studies in larger streams, with Atlantic salmon parr in the small streams occupying the same habitat as favoured by brown trout in larger streams. The data indicate both species may be flexible in their habitat selection depending on habitat availability. Species‐specific habitat overlap between streams may be complete. However, between‐species habitat partitioning remains similar.     

The scale-dependent impact of wolf predation risk on resource selection by three sympatric ungulates

Resource selection is a fundamental ecological process impacting population dynamics and ecosystem structure. Understanding which factors drive selection is vital for effective species- and landscape-level management. We used resource selection probability functions (RSPFs) to study the influence of two forms of wolf (Canis lupus) predation risk, snow conditions and habitat variables on white-tailed deer (Odocoileus virginianus), elk (Cervus elaphus) and moose (Alces alces) resource selection in central Ontario's mixed forest French River-Burwash ecosystem. Direct predation risk was defined as the frequency of a predator's occurrence across the landscape and indirect predation risk as landscape features associated with a higher risk of predation. Models were developed for two winters, each at two spatial scales, using a combination of GIS-derived and ground-measured data. Ungulate presence was determined from snow track transects in 64 16- and 128 1-km(2) resource units, and direct predation risk from GPS radio collar locations of four adjacent wolf packs. Ungulates did not select resources based on the avoidance of areas of direct predation risk at any scale, and instead exhibited selection patterns that tradeoff predation risk minimization with forage and/or mobility requirements. Elk did not avoid indirect predation risk, while both deer and moose exhibited inconsistent responses to this risk. Direct predation risk was more important to models than indirect predation risk but overall, abiotic topographical factors were most influential. These results indicate that wolf predation risk does not limit ungulate habitat use at the scales investigated and that responses to spatial sources of predation risk are complex, incorporating a variety of anti-predator behaviours. Moose resource selection was influenced less by snow conditions than cover type, particularly selection for dense forest, whereas deer showed the opposite pattern. Temporal and spatial scale influenced resource selection by all ungulate species, underlining the importance of incorporating scale into resource selection studies.     

Spatial variation in density of American black bears in northern Yellowstone National Park

The quality and availability of resources are known to influence spatial patterns of animal density. In Yellowstone National Park, relationships between the availability of resources and the distribution of grizzly bears (Ursus arctos) have been explored but have yet to be examined in American black bears (Ursus americanus). We conducted non-invasive genetic sampling during 2017–2018 (mid-May to mid-July) and applied spatially explicit capture-recapture models to estimate density of black bears and examine associations with landscape features. In both years, density estimates were higher in forested vegetation communities, which provide food resources and thermal and security cover preferred by black bears, compared with non-forested areas. In 2017, density also varied by sex, with female densities being higher than males. Based on our estimates, the northern range of Yellowstone National Park supports one of the highest densities of black bears (20 black bears/100 km²) in the northern Rocky Mountains (6–12 black bears/100 km² in other regions). Given these high densities, black bears could influence other wildlife populations more than previously thought, such as through displacement of sympatric predators from kills. Our study provides the first spatially explicit estimates of density for black bears within an ecosystem that contains the majority of North America's large mammal species. Our density estimates provide a baseline that can be used for future research and management decisions of black bears, including efforts to reduce human–bear conflicts.     

Sea ice resource selection models for polar bears in the Barents Sea subpopulation

The extent, thickness and age of Arctic sea ice has dramatically declined since the late 1990s, and these trends are predicted to continue. Exploring the habitat use of sea‐ice‐dependent species can help us understand which resources they use and how their distribution responds to a changing environment. The goal of this study was to develop predictive models of the habitat use of an Arctic apex predator. Polar bear Ursus maritimus habitat use in the Barents Sea subpopulation was modelled with seasonal resource selection functions (RSFs) using satellite‐linked telemetry data from 294 collars deployed on female polar bears between 1991 and 2015. Polar bears selected habitat in the Marginal Ice Zone, with a preference for intermediate sea ice concentrations (40–80%). They spent most time in areas with relatively short travel distances to 15 or 75% ice concentration, and during spring and autumn they exhibited a preference for sea ice areas over the continental shelf or over shallower bathymetry). Predictions of the distribution of polar bears in the Barents Sea area can be made for specific sea ice scenarios using these models. Two such predictive distribution maps based on the autumn seasonal model were made and validated against two independent polar bear survey datasets collected in August 2004 and August 2015. The distribution of optimal polar bear habitat has shifted strongly northwards in all seasons of the year during the 25 yr study period.     

Size-selective and sex-selective predation by brown bears on sockeye salmon

Breeding activity increases the vulnerability of many animals to predation, and such predation can affect the subset of animals successfully reproducing. To study the ways in which predation might affect the evolution of Pacific salmon, we measured the intensity and selectivity of predation by bears (primarily brown bears, Ursus arctos) on mature sockeye salmon (Oncorhynchus nerka) breeding in a series of small, spring-fed ponds and creeks near Pedro Bay, Alaska, from 1994 to 1998. Bears killed male salmon more often than females; males constituted 60% of the kills but only 35% of the salmon that died of senescence. The bears also killed fish that were larger, on average, than those dying of senescence (males: 462 vs 452 mm; females: 453 vs 443 mm). The level of predation varied greatly, from 4% (females) and 10% (males) in 1994 to 100% of both sexes in 1996 and 1997. The rate of predation also varied among habitats, being lower in larger ponds than in smaller, shallower ponds and the very small interconnecting creeks. Despite the intense and size-selective predation, the salmon in safer habitats (large ponds) were not larger than those in riskier habitats, and salmon densities were only slightly higher in the safer areas. Compared to a nearby population that experiences no bear predation (Woody Island), the male sockeye salmon from the Pedro Pond system had shallower bodies (i.e., less exposure in shallow water) for a given length, consistent with the hypothesis that selective predation can affect the extent of sexual dimorphism among populations. However, the average length at age for both males and females was greater in the Pedro Pond fish, indicating that selective factors besides predation affect length. Overall, the results indicate that bears can be an agent of natural selection within (and perhaps between) sockeye salmon populations, and predation can greatly affect reproductive success among individuals and years for the population as a whole. Received: 6 April 1999 / Accepted: 1 June 1999     

Habitat use by black bears in relation to conspecifics and competitors

Sympatric black bears (Ursus americanus) and brown bears (Ursus arctos) are common in many boreal systems; however, few predator assemblages are known to coexist on a single seasonally abundant large prey item. In lowland southwestern interior Alaska, black bears and brown bears are considered the primary cause of moose (Alces alces) calf mortality during the first 6 weeks of life. The objective of this study was to document habitat use of global-positioning system (GPS)-collared black bears during peak and non-peak seasons of black bear-induced and brown bear-induced moose calf mortality within southwestern interior Alaska, in spring 2002. We compared habitats of GPS-collared black bears to those of presumably uncollared black bears and brown bears at their moose calf mortality sites. Results from this study suggest that GPS-collared black bears use similar habitat as conspecifics more than expected during the peak period of black bear predation on moose calves, whereas they use habitat in proportion to home range availability during the peak in brown bear predation on moose calves. Sex-specific Ivlev's electivity indices describe greater than expected use of mixed-deciduous forest and needleleaf forest by male GPS-collared black bears during the peak of moose calf predation, whereas females have a tendency to use these habitats less than expected. Juvenile GPS-collared black bears largely use the same habitat as other sympatric predators during the peak of moose calf predation, whereas during the non-peak period juveniles use opposite habitats as adult GPS-collared black bears. The outcome of this study offers possible explanations (e.g., sex, age) for spatial overlap or segregation in one member of a complex predator guild in relation to a seasonal pulse of preferred prey.     

Social network analysis of mating patterns in American black bears (Ursus americanus)

Nonrandom mating can structure populations and has important implications for population‐level processes. Investigating how and why mating deviates from random is important for understanding evolutionary processes as well as informing conservation and management. Prior to the implementation of parentage analyses, understanding mating patterns in solitary, elusive species like bears was virtually impossible. Here, we capitalize on a long‐term genetic data set collected from black bears (Ursus americanus) (N = 2422) in the Northern Lower Peninsula (NLP) of Michigan, USA. We identified mated pairs using parentage analysis and applied logistic regression (selection) models that controlled for features of the social network, to quantify the effects of individual characteristics, and spatial and population demographic factors on mating dynamics. Logistic regression models revealed that black bear mating was associated with spatial proximity of mates, male age, the time a pair had coexisted, local population density and relatedness. Mated pairs were more likely to contain older males. On average, bears tended to mate with nearby individuals to whom they were related, which does not support the existence of kin recognition in black bears. Pairwise relatedness was especially high for mated pairs containing young males. Restricted dispersal and high male turnover from intensive harvest mortality of NLP black bears are probably the underlying factors associated with younger male bears mating more often with female relatives. Our findings illustrate how harvest has the potential to disrupt the social structure of game species, which warrants further attention for conservation and management.