Negative density-dependent dispersal in the American black bear (Ursus americanus) revealed by noninvasive sampling and genotyping

Although the dispersal of animals is influenced by a variety of factors, few studies have used a condition-dependent approach to assess it. The mechanisms underlying dispersal are thus poorly known in many species, especially in large mammals. We used 10 microsatellite loci to examine population density effects on sex-specific dispersal behavior in the American black bear, Ursus americanus. We tested whether dispersal increases with population density in both sexes. Fine-scale genetic structure was investigated in each of four sampling areas using Mantel tests and spatial autocorrelation analyses. Our results revealed male-biased dispersal pattern in low-density areas. As population density increased, females appeared to exhibit philopatry at smaller scales. Fine-scale genetic structure for males at higher densities may indicate reduced dispersal distances and delayed dispersal by subadults.     

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.     

Density and genetic structure of black bears in coastal South Carolina

The frequency of black bear (Ursus americanus) sightings, vehicle collisions, and nuisance incidents in the coastal region of South Carolina has increased over the past 4 decades. To develop the statewide Black Bear Management and Conservation Strategy, the South Carolina Department of Natural Resources needed reliable information for the coastal population. Because no such data were available, we initiated a study to determine population density and genetic structure of black bears. We selected 2 study areas that were representative of the major habitat types in the study region: Lewis Ocean Bay consisted primarily of Carolina Bays and pocosin habitats, whereas Carvers Bay was representative of extensive pine plantations commonly found in the region. We established hair snares on both study areas to obtain DNA from hair samples during 8 weekly sampling periods in 2008 and again in 2009. We used genotypes to obtain capture histories of sampled bears. We estimated density using spatially explicit capture–recapture (SECR) models and used information-theoretic procedures to fit parameters for capture heterogeneity and behavioral responses and to test if density and model parameters varied by year. Model-averaged density was 0.046 bears/km² (SE = 0.011) for Carvers Bay and 0.339 bears/km² (SE = 0.056) for Lewis Ocean Bay. Next, we sampled habitat covariates for all locations in the SECR sampling grid to derive spatially explicit estimates of density based on habitat characteristics. Addition of habitat covariates had substantial support, and accounted for differences in density between Carvers Bay and Lewis Ocean Bay; black bear density showed a negative association with the area of pine forests (4.5-km² scale) and a marginal, positive association with the area of pocosin habitat (0.3-km² scale). Bear density was not associated with pine forest at a smaller scale (0.3-km²), nor with major road density or an index of largest patch size. Predicted bear densities were low throughout the coastal region and only a few larger areas had high predicted densities, most of which were centered on public lands (e.g., Francis Marion National Forest, Lewis Ocean Bay). We sampled a third bear population in the Green Swamp area of North Carolina for genetic structure analyses and found no evidence of historic fragmentation among the 3 sampled populations. Neither did we find evidence of more recent barriers to gene exchange; with the exception of 1 recent migrant, Bayesian population assignment techniques identified only a single population cluster that incorporated all 3 sampled areas. Bears in the region may best be managed as 1 population. If the goal is to maintain or increase bear densities, demographic connectivity of high-density areas within the low-density landscape matrix is a key consideration and managers would need to mitigate potential impacts of planned highway expansions and anticipated development. Because the distribution of black bears in coastal South Carolina is not fully known, the regional map of potential black bear density can be used to identify focal areas for management and sites that should be surveyed for occupancy or where more intensive studies are needed. © 2012 The Wildlife Society.     

Spatial genetic structure and landscape connectivity in black bears: Investigating the significance of using different land cover datasets and classifications in landscape genetics analyses

Landscape genetic analyses allow detection of fine‐scale spatial genetic structure (SGS) and quantification of effects of landscape features on gene flow and connectivity. Typically, analyses require generation of resistance surfaces. These surfaces characteristically take the form of a grid with cells that are coded to represent the degree to which landscape or environmental features promote or inhibit animal movement. How accurately resistance surfaces predict association between the landscape and movement is determined in large part by (a) the landscape features used, (b) the resistance values assigned to features, and (c) how accurately resistance surfaces represent landscape permeability. Our objective was to evaluate the performance of resistance surfaces generated using two publicly available land cover datasets that varied in how accurately they represent the actual landscape. We genotyped 365 individuals from a large black bear population (Ursus americanus) in the Northern Lower Peninsula (NLP) of Michigan, USA at 12 microsatellite loci, and evaluated the relationship between gene flow and landscape features using two different land cover datasets. We investigated the relative importance of land cover classification and accuracy on landscape resistance model performance. We detected local spatial genetic structure in Michigan''s NLP black bears and found roads and land cover were significantly correlated with genetic distance. We observed similarities in model performance when different land cover datasets were used despite 21% dissimilarity in classification between the two land cover datasets. However, we did find the performance of land cover models to predict genetic distance was dependent on the way the land cover was defined. Models in which land cover was finely defined (i.e., eight land cover classes) outperformed models where land cover was defined more coarsely (i.e., habitat/non‐habitat or forest/non‐forest). Our results show that landscape genetic researchers should carefully consider how land cover classification changes inference in landscape genetic 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.     

No evidence for inbreeding avoidance in a great reed warbler population

Inbreeding depression may drive the evolution of inbreedingavoidance through dispersal and mate choice. In birds, manyspecies show female-biased dispersal, which is an effectiveinbreeding avoidance mechanism. In contrast, there is scarceevidence in birds for kin discriminative mate choice, whichmay, at least partly, reflect difficulties detecting it. First,kin discrimination may be realized as dispersal, and this isdifficult to distinguish from other causes of dispersal. Second,even within small, isolated populations, it is often difficultto determine the potential candidates available to a femalewhen choosing a mate. We sought evidence for inbreeding avoidancevia kin discrimination in a breeding population of great reedwarblers (Acrocephalus arundinaceus) studied over 17 years.Inbreeding depression is strong in the population, suggestingthat it would be adaptive to avoid relatives as mates. Detaileddata on timing of settlement and mate search movements madeit possible to identify candidate mates for each female, andlong-term pedigrees and resolved parentage enabled us to estimaterelatedness between females and their candidate mates. We foundno evidence for kin discrimination: mate choice was random withrespect to relatedness when all mate-choice events were considered,and, after correction for multiple tests, also in all breedingyears. We suggest that dispersal is a sufficient inbreedingavoidance mechanism in most situations, although the lack ofkin discriminative mate choice has negative consequences forsome females, because they end up mating with closely relatedmales that lowers their fitness.     

Variation in life history and demography of the American black bear

Variation in life history and demography across a species' range informs researchers about regional adaptations and affects whether managers can borrow information from other populations in decision-making. The American black bear (Ursus americanus) is a long-lived game species whose continued persistence depends on management of harvest and removal of habituated bears that come into conflict with humans. Understanding the demography of black bears guides efforts at management and conservation, yet detailed knowledge of many populations is typically lacking. I performed a hierarchical Bayesian meta-analysis of black bear demographic studies across the geographic range of the species to explore how vital rates vary across the range, what information they give us about population growth, and whether managers can justify borrowing information from other studies to inform management decisions. Cub, yearling, and adult survival and fecundity varied between eastern and western North America, whereas subadult survival did not show geographic structuring. Adult survival and fecundity appeared to trade off, with higher survival in the western portions of bears' range and higher fecundity in the east. Although adult survival had the highest elasticity, differences in reproduction drove differences in population growth rate. Mean population growth rate was higher in the east (0.99; 95% credible interval [CrI]: 0.96, 1.03) than the west (0.97; 95%CrI: 0.93, 1.01). Despite declining trends in the west, 34% of the distribution of population growth rate was >1, compared to 55% in the east. Further work needs to be done to address the cause of the apparent trade-off between adult survival and fecundity and explore how the estimated growth rates are likely to affect population status of black bears. Because population growth rates are close to 1 and small deviations could impact whether a population is considered increasing or decreasing, managers need to employ caution in borrowing vital rates from other populations. © 2011 The Wildlife Society.     

Female‐biased dispersal alone can reduce the occurrence of inbreeding in black grouse (Tetrao tetrix)

Although inbreeding depression and mechanisms for kin recognition have been described in natural bird populations, inbreeding avoidance through mate choice has rarely been reported suggesting that sex‐biased dispersal is the main mechanism reducing the risks of inbreeding. However, a full understanding of the effect of dispersal on the occurrence of inbred matings requires estimating the inbreeding risks prior to dispersal. Combining pairwise relatedness measures and kinship assignments, we investigated in black grouse whether the observed occurrence of inbred matings was explained by active kin discrimination or by female‐biased dispersal. In this large continuous population, copulations between close relatives were rare. As female mate choice was random for relatedness, females with more relatives in the local flock tended to mate with genetically more similar males. To quantify the initial risks of inbreeding, we measured the relatedness to the males of females captured in their parental flock and virtually translocated female hatchlings in their parental and to more distant flocks. These tests indicated that dispersal decreased the likelihood of mating with relatives and that philopatric females had higher inbreeding risks than the actual breeding females. As females do not discriminate against relatives, the few inbred matings were probably due to the variance in female dispersal propensity and dispersal distance. Our results support the view that kin discrimination mate choice is of little value if dispersal effectively reduces the risks of inbreeding.     

Integrating video and genetic data to estimate annual age-structured apparent survival of American black bears

Camera or genetic data are increasingly used to estimate wildlife abundance and density. We integrated video camera data with genetic data over 7 years to estimate annual age-structured apparent survival of American black bears (Ursus americanus). We identified 70 individuals through meticulous scrutiny of 7531 video captures, cross-referenced with 721 genetic captures from hair samples concurrently collected from stations in view of cameras. We used the Cormack–Jolly–Seber model in Program Mark to estimate annual age-structured apparent survival for yearling males, yearling females, 2+ year-old males, and 2+ year-old females. We manually calculated cub survival. We compared parameter estimates based on combined video and genetic data with those based on only genetic data. Combining video and genetic data provided a means to test video-based identification accuracy, which was highest for females (97%–100%). Annual apparent survival was highest for yearling females (φ = 0.92, SE = 0.07), followed by 2+ year-old females (φ = 0.88, SE = 0.05), 2+ year-old males (φ = 0.84, SE = 0.06), and yearling males (φ = 0.80, SE = 0.14). Annual cub survival (φ = 0.86, SE = 0.07) was likely biased because we could not account for mortality that occurred in-den through early spring. Annual apparent survival and recapture probabilities derived from only genetic data were lower than those derived from combined video and genetic data. Our finding that noninvasive data can be used to estimate annual age-structured apparent survival of a species with relatively indistinct traits is broadly relevant to wildlife research and conservation.     

Role of male‐biased dispersal in inbreeding avoidance in the grey‐sided vole (Myodes rufocanus)

Effects of male‐biased dispersal on inbreeding avoidance were investigated in a semi‐natural population of Myodes (formerly Clethrionomys) rufocanus using a large outdoor enclosure (3 ha). Parentage of 918 voles weaned from 215 litters and relatedness of mates were analysed using microsatellite loci, and dispersal distances were obtained from mark–recapture live‐trapping data. Natal and breeding male‐biased dispersal was observed. There remained, however, chances that incestuous mating could occur, because not all males dispersed from their natal site, and 51 matings occurred between relatives (relatedness r > 0). The number of weaned juveniles from inbred litters was significantly smaller than that from non‐inbred litters. Fourteen incestuous matings occurred between close relatives (r ≥ 0.25), most of which were those between non‐littermate maternal half siblings (four cases) and those between paternal half siblings (seven cases). When comparing the observed frequencies to the expected ones generated by combining every oestrous female with a male randomly chosen from her surroundings, the observed values for inbreeding of r ≥ 0.25 were significantly smaller than the expectations, while no difference was observed for inbreeding of 0 < r < 0.25. These results suggest that male‐biased dispersal is partly effective to avoid incestuous mating, but it does not provide complete separation of male and female close relatives. Additional mechanisms such as kin discrimination based on familiarity may work in inbreeding avoidance of the vole.     

Kin encounter rate and inbreeding avoidance in canids

Mating with close kin can lead to inbreeding depression through the expression of recessive deleterious alleles and loss of heterozygosity. Mate selection may be affected by kin encounter rate, and inbreeding avoidance may not be uniform but associated with age and social system. Specifically, selection for kin recognition and inbreeding avoidance may be more developed in species that live in family groups or breed cooperatively. To test this hypothesis, we compared kin encounter rate and the proportion of related breeding pairs in noninbred and highly inbred canid populations. The chance of randomly encountering a full sib ranged between 1-8% and 20-22% in noninbred and inbred canid populations, respectively. We show that regardless of encounter rate, outside natal groups mates were selected independent of relatedness. Within natal groups, there was a significant avoidance of mating with a relative. Lack of discrimination against mating with close relatives outside packs suggests that the rate of inbreeding in canids is related to the proximity of close relatives, which could explain the high degree of inbreeding depression observed in some populations. The idea that kin encounter rate and social organization can explain the lack of inbreeding avoidance in some species is intriguing and may have implications for the management of populations at risk.     

Evaluation of Connectivity Among American Black Bear Populations in Georgia

Habitat fragmentation and loss contribute to isolation of wildlife populations and increased extinction risks for various species, including many large carnivores. We studied a small and isolated population of American black bears (Ursus americanus) that is of conservation concern in central Georgia, USA (i.e., central Georgia bear population [CGBP]). Our goal was to evaluate the potential for demographic and genetic interchange from neighboring bear populations to the CGBP. To evaluate resource selection and movement potential, we used 35,487 global positioning system locations collected every 20 minutes from 2012 to 2014 from 33 male bears in the CGBP. We then developed a step selection function model based on conditional logistic regression. Male bears chose steps that avoided crops, roads, and human developments and were closer to forests and woody wetlands than expected based on availability. We used a geographic information system to simulate 300 bear movement paths from nearby bear populations in northern Florida, northern Georgia, and southern Georgia to estimate the potential for immigration to the CGBP. Only 4 simulated movement paths from the nearby populations intersected the CGBP. The creation of a hypothetical 1-km-wide corridor between the southern Georgia population and the CGBP produced only minor improvements in interchange. Our findings suggest that demographic connectivity between the CGBP and surrounding bear populations may be limited, and coupled with previous works showing genetic isolation in the CGBP, that creation of corridors may have only marginal effects on restoring gene flow, at least in the near term. Management actions such as translocation and the establishment of stepping stone populations may be needed to increase the genetic diversity and demographic stability of bears in the CGBP. © 2021 The Wildlife Society.     

Sex-biased dispersal and natal philopatry in the diamondback terrapin, Malaclemys terrapin

Nesting ecology and population studies indicate that diamondback terrapins (Malaclemys terrapin) exhibit nest site fidelity and high habitat fidelity. However, genetic studies indicate high levels of gene flow. Because dispersal affects the genetics and population dynamics of a species, we used six highly polymorphic microsatellite markers to investigate sex-biased dispersal and natal philopatry of M. terrapin in Barnegat Bay, NJ. We compared results of spatial autocorrelation analysis, assignment methods and Wright's F(ST) estimators to a mark-recapture analysis. Mark-recapture analysis over a 4-year period indicated that most individuals have relatively small home ranges (<2 km), with mature females displaying greater home ranges than males. Goodness of fit analysis of our mark-recapture study indicated that some juvenile males were likely transient individuals moving through our study location. Mean assignment indices and first-generation migrant tests indicated that mature males were more prone to disperse than mature females, but first-generation migrant tests indicated that per capita there are more female than male dispersers. Thus, the relative importance of males and females on gene flow in terrapin populations may change in relation to population sex ratios. Spatial autocorrelation analysis indicated that mature females exhibited natal philopatry to nesting beaches, but first-generation migrant tests indicated that a small number of females failed to nest on natal beaches. Finally, we discuss the important conservation implications of male-biased dispersal and natal philopatry in the diamondback terrapin.     

Factors associated with black bear density and implications for management

Wildlife density estimates are important to accurately formulate population management objectives and understand the relationship between habitat characteristics and a species’ abundance. Despite advances in density and abundance estimation methods, management of common game species continues to be challenged by a lack of reliable population estimates. In Washington, USA, statewide American black bear (Ursus americanus) abundance estimates are predicated on density estimates derived from research in the 1970s and are hypothesized to be a function of precipitation and vegetation, with higher densities in western Washington. To evaluate current black bear density and landscape relationships in Washington, we conducted a 4-year capture-recapture study in 2 areas of the North Cascade Mountains using 2 detection methods, non-invasive DNA collection and physical capture and deployment of global positioning system (GPS) collars. We integrated GPS telemetry from collared bears with spatial capture-recapture (SCR) data and created a SCR-resource selection model to estimate density as a function of spatial covariates and test the hypothesis that density is higher in areas with greater vegetative food resources. We captured and collared 118 bears 132 times and collected 7,863 hair samples at hair traps where we identified 537 bears from 1,237 detections via DNA. The most-supported model in the western North Cascades depicted a negative relationship between black bear density and an index of human development. We estimated bear density at 20.1 bears/100 km², but density varied from 13.5/100 km² to 27.8 bears/100 km² depending on degree of human development. The model best supported by the data in the eastern North Cascades estimated an average density of 19.2 bears/100 km², which was positively correlated with primary productivity, with resulting density estimates ranging from 7.1/100 km² to 33.6 bears/100 km². The hypothesis that greater precipitation and associated vegetative production in western Washington supports greater bear density compared to eastern Washington was not supported by our data. In western Washington, empirically derived average density estimates (including cubs) were nearly 50% lower than managers expected prior to our research. In eastern Washington average black bear density was predominantly as expected, but localized areas of high primary productivity supported greater than anticipated bear densities. Our findings underscore the importance that black bear density is not likely uniform and management risk may be increased if an average density is applied at too large a scale. Disparities between expected and empirically derived bear density illustrate the need for more rigorous monitoring to understand processes that affect population numbers throughout the jurisdiction, and suggest that management plans may need to be reevaluated to determine if current harvest strategies are achieving population objectives. © 2019 The Wildlife Society.     

Cryptic behaviours, inverse genetic landscapes, and spatial avoidance of inbreeding in the Pacific jumping mouse

Although the behaviour of individuals is known to impact the genetic make-up of a population, observed behavioural patterns do not always correspond to patterns of genetic structure. In particular, philopatric or dispersal-limited species often display lower-than-expected values of relatedness or inbreeding suggestive of the presence of cryptic migration, dispersal, or mating behaviours. I used a combination of microsatellite and mark-recapture data to test for the influence of such behaviours in a dispersal-limited species, the Pacific jumping mouse, within a semi-isolated population over three seasons. Despite short dispersal distances and a low rate of first generation migrants, heterozygosities were high and inbreeding values were low. Dispersal was male-biased; interestingly however, this pattern was only present when dispersal was considered to include movement away from paternal home range. Not unexpectedly, males were polygynous; notably, some females were also found to be polyandrous, selecting multiple neighbouring mates for their single annual litter. Patterns of genetic structure were consistent with these more inconspicuous behavioural patterns. Females were more closely related than males and isolation by distance was present only in females. Furthermore, detailed genetic landscapes revealed the existence of strong, significant negative correlations, with areas of low genetic distance among females overlapping spatially with areas of high genetic distance among males. These results support the hypothesis that the detected cryptic components of dispersal and mating behaviour are reducing the likelihood of inbreeding in this population through paternally driven spatial mixing of male genotypes and polyandry of females.     

Longevity and reuse of black bear dens in managed forests of coastal British Columbia

We evaluated longevity and reuse of denning structures by American black bears (Ursus americanus) in coastal temperate rainforests of British Columbia from 1992 to 2010 to assess potential impacts of forest management on these critical habitat features over time. We identified 67 dens during a 4-yr intensive radio-telemetry study (1992–1995): 40 dens of 21 radio-collared black bears and 27 dens found incidentally. Dens occurred in or beneath large diameter trees or wooden structures derived from trees (i.e., logs, root boles, and stumps; = 143 cm diameter, SD = 49 cm). Longevity of dens varied by type, tree species that formed the den, and forest management that occurred at or near the den. Twenty-four of 28 dens of radio-collared bears that were monitored were still usable in 2010, whereas only 5 of 14 dens found incidentally were still usable in 2010. We assessed reuse of bear dens 3 times following initial identification: during the radio-telemetry study, again in 2000, and finally in 2010. Radio-collared bears reused dens from previous years on 7 of 25 potential occasions during the course of the radio-telemetry study. Upon assessment in 2000 and 2010, 17 of 24 (71%) available dens first used by radio-collared bears were reused at least once between 1993 and 2010. The high rate of reuse may indicate low availability of den structures in our study area. Because black bears in coastal British Columbia only used trees or structures derived from trees for winter dens and forest harvesting reduces the supply of these necessary structures, conservation and recruitment of suitable den trees is necessary if maintaining black bear populations is a management goal in these areas. © 2011 The Wildlife Society.