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We measured stable carbon and nitrogen isotope ratios in guard hair of 81 populations of grizzly bears (Ursus arctos L., 1758) across North America and used mixing models to assign diet fractions of salmon, meat derived from terrestrial sources, kokanee (Oncorhynchus nerka (Walbaum in Artedi, 1792)), and plants. In addition, we examined the relationship between skull size and diet of bears killed by people in British Columbia. The majority of carbon and nitrogen assimilated by most coastal grizzly bear populations was derived from salmon, while interior populations usually derived a much smaller fraction of their nutrients from salmon, even in areas with relatively large salmon runs. Terrestrial prey was a large part of the diet where ungulates were abundant, with the highest fractions observed in the central Arctic, where caribou (Rangifer tarandus (L., 1758)) were very abundant. Bears in some boreal areas, where moose (Alces alces (L., 1758)) were abundant, also ate a lot of meat. Bears in dryer areas with low snowfall tended to have relatively high meat diet fractions, presumably because ungulates are more abundant in such environments. Kokanee were an important food in central British Columbia. In areas where meat was more than about a third of the diet, males and females had similar meat diet fractions, but where meat was a smaller portion of the diet, males usually had higher meat diet fractions than females. Females reached 95% of their average adult skull length by 5 years of age, while males took 8 years. Skull width of male grizzly bears increased throughout life, while this trend was slight in females. Skull size increased with the amount of salmon in the diet, but the influence of terrestrial meat on size was inconclusive. We suggest that the amount of salmon in the diet is functionally related to fitness in grizzly bears.
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.
Spatially explicit capture-recapture methods, used widely to estimate the abundance of large carnivores, allow for movement within home ranges during sampling. Probability of detection is a decreasing function of distance from the home range center, with one parameter for magnitude and another for spatial scale. Sex-based and other differences in home range size potentially cause heterogeneity in individual detection and bias in estimates of density. The two parameters of detection have hitherto been treated as independent, but we suggest that an inverse relation is expected when detection probability depends on time spent near the detector. Variation in the spatial scale of detection is then compensated by reciprocal variation in the magnitude parameter. We define a net measure of detection ("single-detector sampling area," a(0)), and show by simulation that its coefficient of variation (CV) is a better predictor of bias than the CV of either component or the sum of their squared CVs. In an example using the grizzly bear Ursus arctos, the estimated sex variation in a(0) was small despite large variation in each component. From the simulations, the relative bias of density estimates was generally negligible (< 5%) when CV(a(0)) < 30%. Parameterization of the detection model in terms of a(0) and spatial scale can be more parsimonious and significantly aids the biological interpretation of detection parameters.
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