Contrasting Activity Patterns of Sympatric and Allopatric Black and Grizzly Bears

ABSTRACT The distribution of grizzly (Ursus arctos) and American black bears (U. americanus) overlaps in western North America. Few studies have detailed activity patterns where the species are sympatric and no studies contrasted patterns where populations are both sympatric and allopatric. We contrasted activity patterns for sympatric black and grizzly bears and for black bears allopatric to grizzly bears, how human influences altered patterns, and rates of grizzly-black bear predation. Activity patterns differed between black bear populations, with those sympatric to grizzly bears more day-active. Activity patterns of black bears allopatric with grizzly bears were similar to those of female grizzly bears; both were crepuscular and day-active. Male grizzly bears were crepuscular and night-active. Both species were more night-active and less day-active when ≤1 km from roads or developments. In our sympatric study area, 2 of 4 black bear mortalities were due to grizzly bear predation. Our results suggested patterns of activity that allowed for intra- and inter-species avoidance. National park management often results in convergence of locally high human densities in quality bear habitat. Our data provide additional understanding into how bears alter their activity patterns in response to other bears and humans and should help park managers minimize undesirable bear-human encounters when considering needs for temporal and spatial management of humans and human developments in bear habitats.     

Using spatially‐explicit capture–recapture models to explain variation in seasonal density patterns of sympatric ursids

Understanding how environmental factors interact to determine the abundance and distribution of animals is a primary goal of ecology, and fundamental to the conservation of wildlife populations. Studies of these relationships, however, often assume static environmental conditions, and rarely consider effects of competition with ecologically similar species. In many parts of their shared ranges, grizzly bears Ursus arctos and American black bears U. americanus have nearly complete dietary overlap and share similar life history traits. We therefore tested the hypothesis that density patterns of both bear species would reflect seasonal variation in available resources, with areas of higher primary productivity supporting higher densities of both species. We also hypothesized that interspecific competition would influence seasonal density patterns. Specifically, we predicted that grizzly bear density would be locally reduced due to the ability of black bears to more efficiently exploit patchy food resources such as seasonally abundant fruits. To test our hypotheses, we used detections of 309 grizzly and 597 black bears from two independent genetic sampling methods in spatially‐explicit capture–recapture (SECR) models. Our results suggest grizzly bear density was lower in areas of high black bear density during spring and summer, although intraspecific densities were also important, particularly during the breeding season. Black bears had lower densities in areas of high grizzly bear density in spring; however, density of black bears in early and late summer was best explained by primary productivity. Our results are consistent with the hypothesis that smaller‐bodied, more abundant black bears may influence the density patterns of behaviorally‐dominant grizzly bears through exploitative competition. We also suggest that seasonal variation in resource availability be considered in efforts to relate environmental conditions to animal density.     

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.     

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.     

Patch occupancy and abundance of local populations in landscapes differing in degree of habitat fragmentation: a case study of the colonial black‐headed gull,Chroicocephalus ridibundus

Aim This study investigated whether habitat fragmentation at the landscape level influences patch occupancy and abundance of the black‐headed gull, Chroicocephalus ridibundus, and whether the response of the species to environmental factors is consistent across replicated landscape plots. Location Water bodies (habitat patches) in southern Poland. Methods Surveys were conducted in two landscape types (four plots in each): (1) more‐fragmented landscape, in which habitat patches were small (mean size 2.2–6.2 ha) and far apart (mean distance 2.5–3.1 km); and (2) less‐fragmented landscape, in which habitat patches were large (mean size 9.2–16.5 ha) and separated by short distances (mean 0.9–1.4 km). Observations were performed twice in 284 potential habitat patches during the 2007 breeding season. Results Colonies were significantly more frequent and larger in the less‐fragmented landscapes than in the more‐fragmented ones. Probability of patch occupancy and number of breeding birds were positively related with patch size and these relationships were especially strong in the more‐fragmented landscapes. In the less‐fragmented landscapes, the occurrence of black‐headed gulls was negatively related to the distance to the nearest local population, but in the more‐fragmented landscapes such a relationship was not detected. As distance to the nearest habitat patch increased, the probability of the patch occupancy decreased in the more‐fragmented landscapes. Moreover, abundance was negatively influenced by distance to the nearest habitat patch, especially strongly in more‐fragmented landscapes. Proximity of corridors (rivers) positively influenced the occupation of patches regardless of landscape type. The number of islets positively influenced occupancy and abundance of local populations, and this relationship was stronger in the more‐fragmented landscapes. Main conclusions Our results are in agreement with predictions from metapopulation theory and are the first evidence that populations of black‐headed gulls may have a metapopulation structure. However, patch occupancy and abundance were differentially affected by explanatory variables in the more‐fragmented landscapes than in the less‐fragmented ones. This implies that it is impossible to derive, a priori, predictions about presence/abundance patterns based on only a single landscape.     

Quantifying carnivory by grizzly bears in a multi-ungulate system

Grizzly bears (Ursus arctos) cohabit landscapes with a diversity of ungulates, but the degree to which grizzly bears are carnivorous is unclear and likely varies across landscapes. We used stable isotopes of carbon and nitrogen to infer seasonal diets of grizzly bears in northern British Columbia while studying predator–prey dynamics in a largely undisturbed system. We found high seasonal variation in diets among individuals; males consumed more ungulate prey than did females throughout the year. In fall, both sexes increased their consumption of prey; large ungulates constituted 51% and 32% of the fall diets of males and females, respectively. This increase in carnivory appeared to be primarily associated with consumption of elk (Cervus elaphus). Estimates of prey proportions in the diets of grizzly bears were highly sensitive to the range of tissue to diet discrimination values that were incorporated in isotope models. Small changes in discrimination values resulted in estimated prey intakes that varied more than threefold as a percentage of the diet depending on sex and season. We caution against using standard discrimination values, and we recommend that diet reconstructions using stable isotopes be based on tissue-specific values that would be appropriate for the species of interest. © 2012 The Wildlife Society.     

Components of Grizzly Bear Habitat Selection: Density,Habitats, Roads,and Mortality Risk

Abstract: We used resource selection functions (RSF) to estimate the relative probability of use for grizzly bears (Ursus arctos) adjacent to the Parsnip River, British Columbia, Canada, 1998-2003. We collected data from 30 radiocollared bears on a rolling plateau where a large portion of the landscape had been modified by human activities, primarily forestry. We also monitored 24 radiocollared bears in mountain areas largely inaccessible to humans. Bears that lived on the plateau existed at less than one-quarter the density of bears in the mountains. Plateau bears ate more high-quality food items, such as meat and berries, leading us to conclude that food limitation was not responsible for the differences in densities. We hypothesized that plateau bears were limited by human-caused mortality associated with roads constructed for forestry activities. Independent estimates of bear population size from DNA-based mark-recapture techniques allowed us to link populations to habitats using RSF models to scale habitat use patterns to population density. To evaluate whether differences in land-cover type, roads, or mortality risk could account for the disparity in density we used the mountain RSF model to predict habitat use and number of bears on the plateau and vice versa. We predicted increases ranging from 34 bears to 96 bears on the plateau when switching model coefficients, excluding land-cover types; when exchanging land-cover coefficients, the model predicted that the plateau population would be 9 bears lower than was observed. Large reductions in the numbers of mountain bears were predicted by habitat-selection models of bears using the plateau landscape. Although RSF models estimated in mountain and plateau landscapes could not predict bear use and abundance in the other areas, contrasts in models between areas provided a useful tool for examining the effects of human activities on grizzly bears.     

Bird community responses to habitat fragmentation: how consistent are they across landscapes?

Aim The woodland ecosystems of south‐eastern Australia have been extensively disturbed by agriculture and urbanization. Herein, the occurrence of birds in woodland remnants in three distinct landscapes was analysed to examine the effects of different types of landscape matrices on species richness vs. area and species richness vs. isolation relationships and individual species responses to woodland fragmentation. Location The study system comprised three distinct woodland landscapes of the northern Australian Capital Territory and bordering areas of New South Wales. These landscapes (termed agricultural, peri‐urban and urban) are located within 50 km of each other, have remnant fragments of similar age, size, isolation, woodland cover, elevation and climates. The major distinguishing feature of the three landscapes was the properties of the habitats surrounding the numerous woodland remnants. Methods Bird surveys, using an area‐search methodology, were conducted in 1999 and 2000 in 127 remnants in the three landscapes to determine bird species presence/absence. Each remnant was characterized by measures of remnant area, isolation and habitat complexity. To characterize differences between each landscape, we conducted an analysis of the amount of tree cover and human disturbance in each landscape using SPOT imagery and aerial photographs. Linear regressions of woodland‐dependent species richness vs. remnant area and remnant isolation for the three different landscapes were calculated to see if there were any apparent differences. Binomial logistic regressions were used to determine the relationships between the occurrence of each species and the size and isolation of woodland habitat, in each landscape. Results All the landscapes displayed a significant (P < 0.01) species vs. area relationship, but the slope of the urban relationship was significantly greater than those of the other landscapes. In contrast, only the agricultural landscape displayed a significant (P < 0.01) species richness vs. isolation relationship. When individual species were investigated, we found species that were: (1) apparently insensitive to reduction in remnant area and increase in isolation across all landscapes, (2) absent in small remnants in all landscapes, (3) absent in small remnants in all landscapes and also absent in isolated remnants in the agricultural landscape, (4) absent in isolated remnants in the agricultural landscape, and (5) absent in small remnants in the urban landscape. Threshold values (50% probability of occurrence) for area and isolation for individual species were highly variable across the three landscapes. Main conclusions These results indicate that woodland bird communities have a varying response to habitat fragmentation in different landscapes. Whilst we cannot be sure how representative our chosen landscapes are of other similarly composed landscapes, these results suggest that the type of landscape matrix may have a considerable influence on how bird species are affected by woodland fragmentation in the region. For instance, the properties of a matrix may influence both the resources available in the landscape as a whole for different bird species, and the connectivity (dispersal of birds), between woodland remnants. We encourage further research that examines these hypotheses and argue that the management of the matrix should be included in conservation strategies for fragmented landscapes.     

Contrasting habitat use of morphologically similar bat species with differing conservation status in south‐eastern Australia

The east‐coast free‐tailed bat Mormopterus norfolkensis Gray, 1839 is a threatened insectivorous bat that is poorly known and as such conservation management strategies are only broadly prescribed. Insectivorous bats that use human‐modified landscapes are often adapted to foraging in open microhabitats. However, few studies have explored whether open‐adapted bats select landscapes with more of these microhabitat features. We compared three morphologically similar and sympatric, molossid bats (genus Mormopterus) with different conservation status in terms of their association with vegetation, climate, landform and land‐use attributes at landscape and local habitat element scales. We predicted that these species would use similar landscape types, with semi‐cleared and low density urban landscapes used more than forested and heavily cleared landscapes. Additionally, we explored which environmental variables best explained the occurrence of each species by constructing post‐hoc models and habitat suitability maps. Contrary to predictions, we found that the three species varied in their habitat use with no one landscape type used more extensively than other types. Overall, M. norfolkensis was more likely to occur in low‐lying, non‐urban, riparian habitats with little vegetation cover. Mormopterus species 2 occupied similar habitats, but was more tolerant of urban landscapes. In contrast, Mormopterus species 4 occurred more often in cleared than forested landscapes, particularly dry landscapes with little vegetation cover. The extensive use of coastal floodplains by the threatened M. norfolkensis is significant because these habitats are under increasing pressure from human land‐uses and the predicted increase in urbanization is likely to further reduce the amount of suitable habitat.     

Insights into wolf presence in human‐dominated landscapes: the relative role of food availability,humans and landscape attributes

Aim Understanding which human or environmental factors interact to enable or to limit the occurrence and persistence of large carnivores in human‐dominated landscapes is an important issue for their effective conservation, especially under the current scenario of global change where most of their former habitat is being transformed by humans. Location NW Iberian Peninsula. Methods We combine data on the distribution of Iberian wolves (Canis lupus signatus) living in a human‐dominated landscape in NW Spain and variation and partitioning methods to investigate the relative importance of three groups of predictors: food availability, humans and landscape attributes – each group expected to have unequal effects on wolf reproduction and survival – and their interactions on the occurrence of this species. Results We found that the group of predictors related with landscape attributes (altitude, roughness and refuge) strongly determined wolf occurrence, followed by humans and food availability. Variance partitioning analysis revealed that the three most important components determining wolf occurrence were related with landscape attributes: (1) the joint effects of the three predictor groups, (2) the joint effect of humans and landscape attributes and (3) the pure effect of landscape attributes. Altitude had the main independent contribution to explain the probability of wolf occurrence. Main conclusions In human‐dominated landscapes, the occurrence of wolves is the result of a complex interaction among several environmental and human factors. Our results suggest that the characteristics of the landscape (spatial context) – factors associated with the security of wolves facilitating that animals go unnoticed by humans, wolf movements, dispersal events and short‐time colonization – become more important in human‐dominated landscapes and may have played a key role in the occurrence and persistence of this species throughout decades modulating the relationship between humans and wolf distribution.     

Spatial Analysis of Factors Influencing Long-Term Stress in the Grizzly Bear (Ursus arctos) Population of Alberta,Canada

Non-invasive measures for assessing long-term stress in free ranging mammals are an increasingly important approach for understanding physiological responses to landscape conditions. Using a spatially and temporally expansive dataset of hair cortisol concentrations (HCC) generated from a threatened grizzly bear (Ursus arctos) population in Alberta, Canada, we quantified how variables representing habitat conditions and anthropogenic disturbance impact long-term stress in grizzly bears. We characterized spatial variability in male and female HCC point data using kernel density estimation and quantified variable influence on spatial patterns of male and female HCC stress surfaces using random forests. Separate models were developed for regions inside and outside of parks and protected areas to account for substantial differences in anthropogenic activity and disturbance within the study area. Variance explained in the random forest models ranged from 55.34% to 74.96% for males and 58.15% to 68.46% for females. Predicted HCC levels were higher for females compared to males. Generally, high spatially continuous female HCC levels were associated with parks and protected areas while low-to-moderate levels were associated with increased anthropogenic disturbance. In contrast, male HCC levels were low in parks and protected areas and low-to-moderate in areas with increased anthropogenic disturbance. Spatial variability in gender-specific HCC levels reveal that the type and intensity of external stressors are not uniform across the landscape and that male and female grizzly bears may be exposed to, or perceive, potential stressors differently. We suggest observed spatial patterns of long-term stress may be the result of the availability and distribution of foods related to disturbance features, potential sexual segregation in available habitat selection, and may not be influenced by sources of mortality which represent acute traumas. In this wildlife system and others, conservation and management efforts can benefit by understanding spatial- and gender-based stress responses to landscape conditions.     

Estimating distemper virus dynamics among wolves and grizzly bears using serology and Bayesian state‐space models

Many parasites infect multiple hosts, but estimating the transmission across host species remains a key challenge in disease ecology. We investigated the within and across host species dynamics of canine distemper virus (CDV) in grizzly bears (Ursus arctos) and wolves (Canis lupus) of the Greater Yellowstone Ecosystem (GYE). We hypothesized that grizzly bears may be more likely to be exposed to CDV during outbreaks in the wolf population because grizzly bears often displace wolves while scavenging carcasses. We used serological data collected from 1984 to 2014 in conjunction with Bayesian state‐space models to infer the temporal dynamics of CDV. These models accounted for the unknown timing of pathogen exposure, and we assessed how different testing thresholds and the potential for testing errors affected our conclusions. We identified three main CDV outbreaks (1999, 2005, and 2008) in wolves, which were more obvious when we used higher diagnostic thresholds to qualify as seropositive. There was some evidence for increased exposure rates in grizzly bears in 2005, but the magnitude of the wolf effect on bear exposures was poorly estimated and depended upon our prior distributions. Grizzly bears were exposed to CDV prior to wolf reintroduction and during time periods outside of known wolf outbreaks, thus wolves are only one of several potential routes for grizzly bear exposures. Our modeling approach accounts for several of the shortcomings of serological data and is applicable to many wildlife disease systems, but is most informative when testing intervals are short. CDV circulates in a wide range of carnivore species, but it remains unclear whether the disease persists locally within the GYE carnivore community or is periodically reintroduced from distant regions with larger host populations.     

Seasonal habitat use and selection by grizzly bears in Northern British Columbia

We defined patterns of habitat use and selection by female grizzly bears (Ursus arctos) in the Besa-Prophet watershed of northern British Columbia. We fitted 13 adult females with Geographic Positioning System (GPS) radio-collars and monitored them between 2001 and 2004. We examined patterns of habitat selection by grizzly bears relative to topographical attributes and 3 potential surrogates of food availability: land-cover class, vegetation biomass or quality (as measured by the Normalized Difference Vegetation Index), and selection value for prey species themselves (moose [Alces alces], elk [Cervus elaphus], woodland caribou [Rangifer tarandus], Stone's sheep [Ovis dalli stonei]). Although vegetation biomass and quality, and selection values for prey were important in seasonal selection by some individual bears, land-cover class, elevation, aspect, and vegetation diversity most influenced patterns of habitat selection across grizzly bears, which rely on availability of plant foods and encounters with ungulate prey. Grizzly bears as a group avoided conifer stands and areas of low vegetation diversity, and selected for burned land-cover classes and high vegetation diversity across seasons. They also selected mid elevations from what was available within seasonal ranges. Quantifying relative use of different attributes helped place selection patterns within the context of the landscape. Grizzly bears used higher elevations (1,595 ± 31 m SE) in spring and lower elevations (1,436 ± 27 m) in fall; the range of average elevations used among individuals was highest (500 m) during the summer. During all seasons, grizzly bears most frequented aspects with high solar gain. Use was distributed across 10 land-cover classes and depended on season. Management and conservation actions must maintain a diverse habitat matrix distributed across a large elevational gradient to ensure persistence of grizzly bears as levels of human access increase in the northern Rocky Mountains. © 2011 The Wildlife Society.     

Spatial genetic patterns indicate mechanism and consequences of large carnivore cohabitation within development

Patterns of human development are shifting from concentrated housing toward sprawled housing intermixed with natural land cover, and wildlife species increasingly persist in close proximity to housing, roads, and other anthropogenic features. These associations can alter population dynamics and evolutionary trajectories. Large carnivores increasingly occupy urban peripheries, yet the ecological consequences for populations established entirely within urban and exurban landscapes are largely unknown. We applied a spatial and landscape genetics approach, using noninvasively collected genetic data, to identify differences in black bear spatial genetic patterns across a rural‐to‐urban gradient and quantify how development affects spatial genetic processes. We quantified differences in black bear dispersal, spatial genetic structure, and migration between differing levels of development within a population primarily occupying areas with >6 houses/km² in western Connecticut. Increased development disrupted spatial genetic structure, and we found an association between increased housing densities and longer dispersal. We also found evidence that roads limited gene flow among bears in more rural areas, yet had no effect among bears in more developed ones. These results suggest dispersal behavior is condition‐dependent and indicate the potential for landscapes intermixing development and natural land cover to facilitate shifts toward increased dispersal. These changes can affect patterns of range expansion and the phenotypic and genetic composition of surrounding populations. We found evidence that subpopulations occupying more developed landscapes may be sustained by male‐biased immigration, creating potentially detrimental demographic shifts.     

Prevalence of Trichinella spp. in black bears, grizzly bears, and wolves in the Dehcho Region, Northwest Territories, Canada, including the first report of T. nativa in a grizzly bear from Canada

Samples of muscle from 120 black bears (Ursus americanus), 11 grizzly bears (Ursus arctos), and 27 wolves (Canis lupus) collected in the Dehcho Region of the Northwest Territories from 2001 to 2010 were examined for the presence of Trichinella spp. larvae using a pepsin-HCl digestion assay. Trichinella spp. larvae were found in eight of 11 (73%) grizzly bears, 14 of 27 (52%) wolves, and seven of 120 (5.8%) black bears. The average age of positive grizzly bears, black bears, and wolves was 13.5, 9.9, and approximately 4 yr, respectively. Larvae from 11 wolves, six black bears, and seven grizzly bears were genotyped. Six wolves were infected with T. nativa and five with Trichinella T6, four black bears were infected with T. nativa and two with Trichinella T6, and all seven grizzly bears were infected with Trichinella T6 and one of them had a coinfection with T. nativa. This is the first report of T. nativa in a grizzly bear from Canada. Bears have been linked to trichinellosis outbreaks in humans in Canada, and black bears are a subsistence food source for residents of the Dehcho region. In order to assess food safety risk it is important to monitor the prevalence of Trichinella spp. in both species of bear and their cohabiting mammalian food sources.     

Influence of whitebark pine decline on fall habitat use and movements of grizzly bears in the Greater Yellowstone Ecosystem

When abundant, seeds of the high‐elevation whitebark pine (WBP; Pinus albicaulis) are an important fall food for grizzly bears (Ursus arctos) in the Greater Yellowstone Ecosystem. Rates of bear mortality and bear/human conflicts have been inversely associated with WBP productivity. Recently, mountain pine beetles (Dendroctonus ponderosae) have killed many cone‐producing WBP trees. We used fall (15 August–30 September) Global Positioning System locations from 89 bear years to investigate temporal changes in habitat use and movements during 2000–2011. We calculated Manly–Chesson (MC) indices for selectivity of WBP habitat and secure habitat (≥500 m from roads and human developments), determined dates of WBP use, and documented net daily movement distances and activity radii. To evaluate temporal trends, we used regression, model selection, and candidate model sets consisting of annual WBP production, sex, and year. One‐third of sampled grizzly bears had fall ranges with little or no mapped WBP habitat. Most other bears (72%) had a MC index above 0.5, indicating selection for WBP habitats. From 2000 to 2011, mean MC index decreased and median date of WBP use shifted about 1 week later. We detected no trends in movement indices over time. Outside of national parks, there was no correlation between the MC indices for WBP habitat and secure habitat, and most bears (78%) selected for secure habitat. Nonetheless, mean MC index for secure habitat decreased over the study period during years of good WBP productivity. The wide diet breadth and foraging plasticity of grizzly bears likely allowed them to adjust to declining WBP. Bears reduced use of WBP stands without increasing movement rates, suggesting they obtained alternative fall foods within their local surroundings. However, the reduction in mortality risk historically associated with use of secure, high‐elevation WBP habitat may be diminishing for bears residing in multiple‐use areas.     

Complementary food resources of carnivory and frugivory affect local abundance of an omnivorous carnivore

A major unresolved question for omnivorous carnivores, like most species of bears, is to what degree are populations influenced by bottom–up (food supply) or top–down (human‐caused mortality) processes. Most previous work on bear populations has focused on factors that limit survival (top–down) assuming little effect of food resource supply. When food resources are considered, most often they consider only the availability/supply of a single resource, particularly marine‐subsidized or terrestrial sources of protein (carnivory) or alternately hard or soft mast (frugivory). Little has been done to compare the importance of each of these factors for omnivorous bears or test whether complementary resources better explain individual animal and population measures such as density, vital rates, and body size. We compared landscape patterns of digestible energy (kcal) for buffaloberry (a key source of carbohydrate) and ungulate matter (a key source of protein and lipid) to local measures in grizzly bear Ursus arctos abundance at DNA hair snag sites in west‐central Alberta, Canada. We tested support for bottom–up hypotheses in either single (carnivory [meat] versus frugivory [fruit]) or complementary (additive or multiplicative) food resources, while accounting for a well‐known top–down limiting factor affecting bear survival (road density). We found support for both top–down and bottom–up factors with complementary resources (co‐limitation) supported over single resource supplies of either meat or fruit. Our study suggests that the availability of food resources that provide complementary nutrients is more important in predicting local bear abundance than single foods or nutrients (e.g. protein) or simply energy per se. This suggests a nutritionally multidimensional bottom–up limitation for a low density interior population of grizzly bears.     

Grizzly bear responses to restrictions of recreation in Yellowstone National Park

Avoiding humans will be more difficult and energetically costly for animals as outdoor recreation increases and people venture farther into wildland areas that provide high-quality habitat for wildlife. Restricting human access can be an attractive management tool to mitigate effects of human recreation activities on wildlife; however, the efficacy of such measures is rarely assessed. In 1982, Yellowstone National Park identified areas important to grizzly bears (Ursus arctos) to help protect critical grizzly bear habitat and reduce the likelihood of human injuries by bears. Referred to as bear management areas (BMAs), human access is restricted in these areas for 2–8 months each year, with timing and type of restrictions varying by area. We examined 2 datasets to evaluate grizzly bear selection of BMAs and differences of bear density in BMAs and non-BMAs. First, we used 17 years of recent global positioning system telemetry data for grizzly bears to assess their selection of BMAs during periods when human access was allowed, and when access was restricted. We used step-selection functions to test the hypothesis that bears spend time in places that allow them to avoid people and select quality food sources. There was support that grizzly bears differentially select for BMAs regardless of whether human access was restricted at the time, compared with areas outside BMAs, and that selection changed with sex and season. Only males during the summer and hyperphagic seasons changed their selection of BMAs based on whether access restrictions were in place, and overall, male bears preferred unrestricted BMAs (BMAs without restrictions in place). Females preferentially selected BMAs regardless of whether the area had access restrictions in place only during the mating season. Individuals varied widely in their preference for BMAs and access restrictions. Bears likely choose to spend time in BMAs based on available food resources rather than restrictions to human access. Supporting this interpretation, our analyses indicated that a greater proportion of BMA in an area was associated with higher densities of grizzly bear. Thus, restrictions to human access likely help reduce the potential for human–bear interactions, accomplishing one of the original objectives for establishing the BMAs.     

Hazards Affecting Grizzly Bear Survival in the Greater Yellowstone Ecosystem

Abstract: During the past 2 decades, the grizzly bear (Ursus arctos) population in the Greater Yellowstone Ecosystem (GYE) has increased in numbers and expanded its range. Early efforts to model grizzly bear mortality were principally focused within the United States Fish and Wildlife Service Grizzly Bear Recovery Zone, which currently represents only about 61% of known bear distribution in the GYE. A more recent analysis that explored one spatial covariate that encompassed the entire GYE suggested that grizzly bear survival was highest in Yellowstone National Park, followed by areas in the grizzly bear Recovery Zone outside the park, and lowest outside the Recovery Zone. Although management differences within these areas partially explained differences in grizzly bear survival, these simple spatial covariates did not capture site-specific reasons why bears die at higher rates outside the Recovery Zone. Here, we model annual survival of grizzly bears in the GYE to 1) identify landscape features (i.e., foods, land management policies, or human disturbances factors) that best describe spatial heterogeneity among bear mortalities, 2) spatially depict the differences in grizzly bear survival across the GYE, and 3) demonstrate how our spatially explicit model of survival can be linked with demographic parameters to identify source and sink habitats. We used recent data from radiomarked bears to estimate survival (1983–2003) using the known-fate data type in Program MARK. Our top models suggested that survival of independent (age ≥ 2 yr) grizzly bears was best explained by the level of human development of the landscape within the home ranges of bears. Survival improved as secure habitat and elevation increased but declined as road density, number of homes, and site developments increased. Bears living in areas open to fall ungulate hunting suffered higher rates of mortality than bears living in areas closed to hunting. Our top model strongly supported previous research that identified roads and developed sites as hazards to grizzly bear survival. We also demonstrated that rural homes and ungulate hunting negatively affected survival, both new findings. We illustrate how our survival model, when linked with estimates of reproduction and survival of dependent young, can be used to identify demographically the source and sink habitats in the GYE. Finally, we discuss how this demographic model constitutes one component of a habitat-based framework for grizzly bear conservation. Such a framework can spatially depict the areas of risk in otherwise good habitat, providing a focus for resource management in the GYE.     

A study of predominant aerobic microflora of black bears (Ursus americanus) and grizzly bears (Ursus arctos) in northwestern Alberta

Swab specimens were obtained from nasal, rectal, and preputial or vaginal areas of 37 grizzly and 17 black bears, captured during May to June of 1981 to 1983, to determine the types and frequency of predominant aerobic microflora. Bacterial genera most frequently isolated from bears were Escherichia, Citrobacter, Hafnia, Proteus, Staphylococcus, and Streptococcus species, comprising about 65% of the isolates. Erwinia, Xanthomonas, Agrobacterium, Rhizobium, and Gluconobacter/Acetobacter were also isolated but at lower frequencies (less than 5%). Comparison of bacterial generic composition using similarity quotient values showed no appreciable differences between grizzly and black bear flora. Also, no outstanding differences in bacterial generic composition were observed among grizzly bear samples; however, differences were noted among black bear samples. Fungal genera most commonly encountered included Cryptococcus, Rhodotorula, Cladosporium, Penicillium, Sporobolomyces, and Candida. In general, the microflora of both bear types were marked by generic diversity and random distribution. The majority of microorganisms isolated from the plant samples in the study area were also found in bear samples. This observation and the presence of certain water and soil bacteria in samples from bears suggest that the predominant microflora of both grizzly and black bears were transient and probably influenced by their foraging habits and surrounding environments.