The potential use of stable-isotope analyses (6I3C and 6I5N) to estimate bear diets was assessed in 40-day feeding trials using American black bears (Ursus americanus). Bear plasma and red blood cells have half-lives of -4 days and -28 days, respectively. The isotopic signature of bear plasma is linearly related to that of the diet, and with the exception of adipose tissue, there is no isotopic fractionation across bear tissues. Isotopic analyses were used to estimate the diets of three bear populations: Pleistocene cave bears (U. speleaus) in Europe, grizzly bears (Ursus arctos horribilis) inhabiting the Columbia River drainage prior to 1931, and brown bears (U. arctos) of Chichagof and Admiralty islands, Alaska. Cave bears were omnivores with terrestrially produced meat contributing from 4.1 to 78% (58 f 14%) of their metabolized carbon and nitrogen. Salmon contributed from 33 to 90% (58 f 23%) of the metabolized carbon and nitrogen in grizzly bears from the Columbia River drainage. Finally, most brown bears on Chichagof and Admiralty islands feed upon salmon during the late summer and fall; however, a subpopulation of bears exists that does not utilize salmon.

During the past 2 decades, the grizzly bear (Ursus arctos) population in the Greater Yellowstone Ecosystem (GYE) has increased in numbers and expanded in range. Understanding temporal, environmental, and spatial variables responsible for this change is useful in evaluating what likely influenced grizzly bear demographics in the GYE and where future management efforts might benefit conservation and management. We used recent data from radio-marked bears to estimate reproduction (1983)(1984)(1985)(1986)(1987)(1988)(1989)(1990)(1991)(1992)(1993)(1994)(1995)(1996)(1997)(1998)(1999)(2000)(2001)(2002) and survival (1983)(1984)(1985)(1986)(1987)(1988)(1989)(1990)(1991)(1992)(1993)(1994)(1995)(1996)(1997)(1998)(1999)(2000)(2001); these we combined into models to evaluate demographic vigor (lambda [k]). We explored the influence of an array of individual, temporal, and spatial covariates on demographic vigor.We identified an important relationship between k and where a bear resides within the GYE. This potential for a source-sink dynamic in the GYE, coupled with concerns for managing sustainable mortality, reshaped our thinking about how management agencies might approach longterm conservation of the species. Consequently, we assessed the current spatial dynamic of the GYE grizzly bear population. Throughout, we followed the information-theoretic approach. We developed suites of a priori models that included individual, temporal, and spatial covariates that potentially affected reproduction and survival. We selected our best approximating models using Akaike's information criterion (AIC) adjusted for small sample sizes and overdispersion (AIC c or QAIC c , respectively).We provide recent estimates for reproductive parameters of grizzly bears based on 108 adult ( .3 years old ) females observed for 329 bearyears. We documented production of 104 litters with cub counts for 102 litters. Mean age of females producing their first litter was 5.81 years and ranged from 4 to 7 years. Proportion of nulliparous females that produced cubs at age 4-7 years was 9.8, 29.4, 56.4, and 100%, respectively. Mean (6SE) litter size (n ¼ 102) was 2.0 6 0.1. The proportion of litters of 1, 2, and 3 cubs was 0.18, 0.61, and 0.22, respectively. Mean yearling litter size (n ¼ 57 ) was 2.0 6 0.1. The proportion of litters containing 1, 2, 3, and 4 yearlings was 0.26, 0.51, 0.21, and 0.02, respectively. The proportion of radio-marked females accompanied by cubs varied among years from 0.05 to 0.60; the mean was 0.316 6 0.03. Reproductive rate was estimated as 0.318 female cubs/female/year. We evaluated the probability of producing a litter of 0-3 cubs relative to a suite of individual and temporal covariates using multinomial logistic regression. Our best models indicated that reproductive output, measured as cubs per litter, was most strongly influenced by indices of population size and whitebark pine (Pinus albicaulis) cone production. Our data suggest a possible density-dependent response in reproductive output, although perinatal mortality could...

Population fragmentation compromises population viability, reduces a species ability to respond to climate change, and ultimately may reduce biodiversity. We studied the current state and potential causes of fragmentation in grizzly bears over approximately 1,000,000 km2 of western Canada, the northern United States (US), and southeast Alaska. We compiled much of our data from projects undertaken with a variety of research objectives including population estimation and trend, landscape fragmentation, habitat selection, vital rates, and response to human development. Our primary analytical techniques stemmed from genetic analysis of 3,134 bears, supplemented with radiotelemetry data from 792 bears. We used 15 locus microsatellite data coupled with measures of genetic distance, isolation‐by‐distance (IBD) analysis, analysis of covariance (ANCOVA), linear multiple regression, multi‐factorial correspondence analysis (to identify population divisions or fractures with no a priori assumption of group membership), and population‐assignment methods to detect individual migrants between immediately adjacent areas. These data corroborated observations of inter‐area movements from our telemetry database. In northern areas, we found a spatial genetic pattern of IBD, although there was evidence of natural fragmentation from the rugged heavily glaciated coast mountains of British Columbia (BC) and the Yukon. These results contrasted with the spatial pattern of fragmentation in more southern parts of their distribution. Near the Canada–US border area, we found extensive fragmentation that corresponded to settled mountain valleys and major highways. Genetic distances across developed valleys were elevated relative to those across undeveloped valleys in central and northern BC. In disturbed areas, most inter‐area movements detected were made by male bears, with few female migrants identified. North–south movements within mountain ranges (Mts) and across BC Highway 3 were more common than east–west movements across settled mountain valleys separating Mts. Our results suggest that relatively distinct subpopulations exist in this region, including the Cabinet, Selkirk South, and the decades‐isolated Yellowstone populations. Current movement rates do not appear sufficient to consider the subpopulations we identify along the Canada–US border as 1 inter‐breeding unit. Although we detected enough male movement to mediate gene flow, the current low rate of female movement detected among areas is insufficient to provide a demographic rescue effect between areas in the immediate future (0–15 yr). In Alberta, we found fragmentation corresponded to major east–west highways (Highways 3, 11, 16, and 43) and most inter‐area movements were made by males. Gene flow and movement rates between Alberta and BC were highest across the Continental Divide south of Highway 1 and north of Highway 16. In the central region between Highways 1 and 11, we found evidence of natural fragmentation associated with the extensive glaciers and icefields along the Continental Divide. The discontinuities that we identified would form appropriate boundaries for management units. We related sex‐specific movement rates between adjacent areas to several metrics of human use (highway traffic, settlement, and human‐caused mortality) to understand the causes of fragmentation. This analysis used data from 1,508 bears sampled over a 161,500‐km2 area in southeastern BC, western Alberta, northern Idaho, and northern Montana during 1979–2007. This area was bisected by numerous human transportation and settlement corridors of varying intensity and complexity. We used multiple linear regression and ANCOVA to document the responses of female and male bears to disturbance. Males and females both demonstrated reduced movement rates with increasing settlement and traffic. However, females reduced their movement rates dramatically when settlement increased to >20% of the fracture zone. At this same threshold, male movement declined more gradually, in response to increased traffic and further settlement. In highly settled areas (>50%), both sexes had a similar reduction in movements in response to traffic, settlement, and mortality. We documented several small bear populations with male‐only immigration, highlighting the importance of investigating sex‐specific movements. Without female connectivity, small populations are not viable over the long term. The persistence of this regional female fragmented metapopulation likely will require strategic connectivity management. We therefore recommend enhancing female connectivity among fractured areas by securing linkage‐zone habitat appropriate for female dispersal, and ensuring current large source subpopulations remain intact. The fragmentation we documented may also affect other species with similar ecological characteristics: sparse densities, slow reproduction, short male‐biased dispersal, and a susceptibility to human‐caused mortality and habitat degradation. Therefore, regional inter‐jurisdictional efforts to manage broad landscapes for inter‐area movement will likely benefit a broad spectrum of species and natural processes, particularly in light of climate change. © 2011 The Wildlife Society.

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Highways and railroads have come under increasing scrutiny as potential agents of population and habitat fragmentation for many mammalian species, including grizzly bears (Ursus arctos). Using Global Positioning System (GPS) technology and aerial Very High Frequency (VHF) telemetry we evaluated the nature and extent of trans‐highway movements of 42 grizzly bears along the U.S. Highway 2 (US‐2) corridor in northwest Montana, USA, 1998–2001, and we related them to highway and railroad traffic volumes and other corridor attributes. We employed highway and railroad traffic counters to continuously monitor traffic volumes. We found that 52% of the sampled population crossed highways at least once during the study but that crossing frequency was negatively exponentially related to highway traffic volume. We found that grizzly bears strongly avoided areas within 500 m of the highway and that highway crossing locations were clustered at a spatial scale of 1.5 km. Most highway crossings occurred at night when highway traffic volume was lowest but when railroad traffic was highest. Highway crossing locations were flatter, closer to cover in open habitat types, and within grassland or deciduous forest vegetation types. Nighttime traffic volumes were low, averaging about 10 vehicles/hr, allowing bears to cross. However, we project that US‐2 may become a significant barrier to bear movement in ∼30 years if the observed trend of increasing traffic volume continues.

We hypothesized that the relative availability of meat, indicated by contribution to the diet, would be positively related to body size and population productivity of North American brown, or grizzly, bears (Ursus arctos). Dietary contributions of plant matter and meat derived from both terrestrial and marine sources were quantified by stable-isotope analysis (δ13C and δ15N) of hair samples from 13 brown bear populations. Estimates of adult female body mass, mean litter size, and population density were obtained from two field studies of ours and from other published reports. The populations ranged from largely vegetarian to largely carnivorous, and food resources ranged from mostly terrestrial to mostly marine (salmon, Oncorhynchus spp.). The proportion of meat in the diet was significantly correlated with mean adult female body mass (r = 0.87, P < 0.01), mean litter size (r = 0.72, P < 0.01), and mean population density (r = 0.91, P < 0.01). Salmon was the most important source of meat for the largest, most carnivorous bears and most productive populations. We conclude that availability of meat, particularly salmon, greatly influences habitat quality for brown bears at both the individual level and the population level.

We estimated grizzly bear (Ursus arctos) population vital rates and trend for the Northern Continental Divide Ecosystem (NCDE), Montana, between 2004 and 2009 by following radio-collared females and observing their fate and reproductive performance. Our estimates of dependent cub and yearling survival were 0.612 (95% CI ¼ 0.300-0.818) and 0.682 (95% CI ¼ 0.258-0.898). Our estimates of subadult and adult female survival were 0.852 (95% CI ¼ 0.628-0.951) and 0.952 (95% CI ¼ 0.892-0.980). From visual observations, we estimated a mean litter size of 2.00 cubs/litter. Accounting for cub mortality prior to the first observations of litters in spring, our adjusted mean litter size was 2.27 cubs/litter. We estimated the probabilities of females transitioning from one reproductive state to another between years. Using the stable state probability of 0.322 (95% CI ¼ 0.262-0.382) for females with cub litters, our adjusted fecundity estimate (m x ) was 0.367 (95% CI ¼ 0.273-0.461). Using our derived rates, we estimated that the population grew at a mean annual rate of approximately 3% (l ¼ 1.0306, 95% CI ¼ 0.928-1.102), and 71.5% of 10,000 Monte Carlo simulations produced estimates of l > 1.0. Our results indicate an increasing population trend of grizzly bears in the NCDE. Coupled with concurrent studies of population size, we estimate that over 1,000 grizzly bears reside in and adjacent to this recovery area. We suggest that monitoring of population trend and other vital rates using radioed females be continued. ß 2011 The Wildlife Society.

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