Assessment of the Sustainability of Wolverine Harvest in British Columbia,Canada

ABSTRACT Wolverines (Gulo gulo) are distributed across much of northern and western Canada and Alaska, USA, and they extend south into the mountainous western United States. Wolverines occur in most regions of British Columbia, Canada, with the highest population densities occurring in the interior mountainous areas. Wolverine populations in British Columbia have been primarily managed to provide a sustainable harvest for trappers and hunters. We used spatially based population estimates, population vital rate data, and spatially based harvest data to evaluate the sustainability of wolverine harvest (trapping and hunting) from 1985 to 2004. The median annual provincial wolverine harvest from 1985 to 2004 was 172 wolverines per year ( = 174.8), which was less than the median simulated estimate of provincial recruitment (195.9 wolverines/yr; = 209.7). Harvests in individual population units ranged from 0 to 280 over the 20-year period. Spatially, wolverine harvest was likely to have been unsustainable in 15 of the 71 population units with wolverines, and it was likely to have been sustainable in the remaining population units. Harvest in 5 of the other 56 population units was marginally sustainable and thus of potential management concern. To improve harvest management of wolverines in British Columbia, wildlife managers should focus on improved data collection and monitoring at a provincial scale, and they should work with trappers and hunters at regional scales to address issues specific to individual population units. Further research is required to improve the reliability of wolverine vital rate and population data.     

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.     

Grizzly bear selection of avalanche chutes: Testing the effectiveness of forest buffer retention

In mountainous areas with sufficient snowfall, avalanche chutes are an important component of grizzly bear (Ursus arctos) habitat. Therefore, regional land-use plans have recommended retaining adjacent forest buffers to maintain security and thus reduce potential impacts of clearcut forest harvesting. Our objective was to determine if forest buffers affected selection of avalanche chutes by grizzly bears, while accounting for factors such as vegetation composition and other physical attributes. We used radio-location data from 61 grizzly bears collected between 1994 and 2000 in southern British Columbia, mapped a sample of avalanche chutes (1,045), and quantified the amount of forb, shrub, tree, and non-vegetated cover within each chute. We also measured forested buffer width on each side of the chute, solar radiation, chute size, chute frequency (no. of chutes/km), and the area of clearcut logging adjacent to chutes. Each avalanche chute was the sample unit and the number of grizzly bear radiolocations was the dependent variable. We found that natural biophysical attributes were the strongest factors predicting the level of avalanche chute use by bears. Frequency of large chutes (>100 m wide), chute area, forb content, and solar radiation all positively affected use by bears. Larger avalanche chutes had a higher proportion of forb cover than smaller chutes, and more of these large chutes per unit area provided increased forage opportunities. Based on multivariate analyses, forested buffer width or the amount of clearcut logging were not strong factors predicting the level of use. However, a post hoc univariate analysis revealed that clearcut logging reduced the amount of bear use of the best avalanche chutes (large and abundant chutes). Furthermore, because a portion of our study area contained logging but no vehicle traffic, we concluded that it was the removal of tree cover, rather than displacement by vehicles, that caused the observed pattern. Although our multivariate models did not perform well using independent validation in a different geographic area, 4 factors were consistently important (large and abundant chutes, forb content, with a negative but weaker influence of clearcutting), suggesting broad applicability of these factors in mountainous ecosystems. © 2011 The Wildlife Society.     

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.     

Population fragmentation and inter-ecosystem movements of grizzly bears in western Canada and the northern United States

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.     

Grizzly bear population vital rates and trend in the Northern Continental Divide Ecosystem,Montana

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 = 0.300–0.818) and 0.682 (95% CI = 0.258–0.898). Our estimates of subadult and adult female survival were 0.852 (95% CI = 0.628–0.951) and 0.952 (95% CI = 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 = 0.262–0.382) for females with cub litters, our adjusted fecundity estimate (m_x) was 0.367 (95% CI = 0.273–0.461). Using our derived rates, we estimated that the population grew at a mean annual rate of approximately 3% (λ = 1.0306, 95% CI = 0.928–1.102), and 71.5% of 10,000 Monte Carlo simulations produced estimates of λ > 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. © 2011 The Wildlife Society.     

Phylogeography and mitochondrial diversity of extirpated brown bear (Ursus arctos) populations in the contiguous United States and Mexico

The fossil record indicates that the brown bear (Ursus arctos) colonized North America from Asia over 50 000 years ago. The species historically occupied the western United States and northern Mexico but has been extirpated from over 99% of this range in the last two centuries. To evaluate colonization hypotheses, subspecific classifications, and historical patterns and levels of genetic diversity in this region, we sequenced 229 nucleotides of the mitochondrial DNA control region in 108 museum specimens. The work was set in a global context by synthesizing all previous brown bear control region sequences from around the world. In mid-latitude North America a single moderately diverse clade is observed, represented by 23 haplotypes with up to 3.5% divergence. Only eight of 23 haplotypes (35%) are observed in the extensively sampled extant populations suggesting a substantial loss of genetic variability. The restriction of all haplotypes from mid-latitude North America to a single clade suggests that this region was founded by bears with a similar maternal ancestry. However, the levels and distributions of diversity also suggest that the colonizing population was not a small founder event, and that expansion occurred long enough ago for local mutations to accrue. Our data are consistent with recent genetic evidence that brown bears were south of the ice prior to the last glacial maximum. There is no support for previous subspecies designations, although bears of the southwestern United States may have had a distinctive, but recent, pattern of ancestry.     

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.     

Vital rates of two small populations of brown bears in Canada and range‐wide relationship between population size and trend

Identifying mechanisms of population change is fundamental for conserving small and declining populations and determining effective management strategies. Few studies, however, have measured the demographic components of population change for small populations of mammals (<50 individuals). We estimated vital rates and trends in two adjacent but genetically distinct, threatened brown bear (Ursus arctos) populations in British Columbia, Canada, following the cessation of hunting. One population had approximately 45 resident bears but had some genetic and geographic connectivity to neighboring populations, while the other population had <25 individuals and was isolated. We estimated population‐specific vital rates by monitoring survival and reproduction of telemetered female bears and their dependent offspring from 2005 to 2018. In the larger, connected population, independent female survival was 1.00 (95% CI: 0.96–1.00) and the survival of cubs in their first year was 0.85 (95% CI: 0.62–0.95). In the smaller, isolated population, independent female survival was 0.81 (95% CI: 0.64–0.93) and first‐year cub survival was 0.33 (95% CI: 0.11–0.67). Reproductive rates did not differ between populations. The large differences in age‐specific survival estimates resulted in a projected population increase in the larger population (λ = 1.09; 95% CI: 1.04–1.13) and population decrease in the smaller population (λ = 0.84; 95% CI: 0.72–0.95). Low female survival in the smaller population was the result of both continued human‐caused mortality and an unusually high rate of natural mortality. Low cub survival may have been due to inbreeding and the loss of genetic diversity common in small populations, or to limited resources. In a systematic literature review, we compared our population trend estimates with those reported for other small populations (<300 individuals) of brown bears. Results suggest that once brown bear populations become small and isolated, populations rarely increase and, even with intensive management, recovery remains challenging.     

Efficacy of firearms for bear deterrence in Alaska

We compiled, summarized, and reviewed 269 incidents of bear–human conflict involving firearms that occurred in Alaska during 1883–2009. Encounters involving brown bears (Ursus arctos; 218 incidents, 81%), black bears (Ursus americanus; 30 incidents, 11%), polar bears (Ursus maritimus; 6 incidents, 2%), and 15 (6%) unidentified species provided insight into firearms success and failure. A total of 444 people and at least 367 bears were involved in these incidents. We found no significant difference in success rates (i.e., success being when the bear was stopped in its aggressive behavior) associated with long guns (76%) and handguns (84%). Moreover, firearm bearers suffered the same injury rates in close encounters with bears whether they used their firearms or not. Bears were killed in 61% (n = 162) of bear–firearms incidents. Additionally, we identified multiple reasons for firearms failing to stop an aggressive bear. Using logistic regression, the best model for predicting a successful outcome for firearm users included species and cohort of bear, human activity at time of encounter, whether or not the bear charged, and if fish or game meat was present. Firearm variables (e.g., type of gun, number of shots) were not useful in predicting outcomes in bear–firearms incidents. Although firearms have failed to protect some users, they are the only deterrent that can lethally stop an aggressive bear. Where firearms have failed to protect people, we identified contributing causes. Our findings suggest that only those proficient in firearms use should rely on them for protection in bear country. © 2012 The Wildlife Society.     

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.     

Spatiotemporal railway use by grizzly bears in Canada's Rocky Mountains

Railway networks contribute to the direct mortality of wildlife through collisions with trains, which can threaten vulnerable wildlife populations even in protected areas, including grizzly bears (Ursus arctos) in Banff and Yoho National Parks, Canada. Mitigation to reduce bear-train collisions requires information about how grizzly bears use the railway spatially and temporally and how particular types of use might increase collision vulnerability. We used data from 27 grizzly bears fitted with global positioning system (GPS) collars between 2000 and 2016 to relate railway use by bears via resource selection functions to variables that described land cover, human use, and topography. We used the same suite of explanatory variables to distinguish pairs of 4 types of steps, in which 3 successive GPS points (with 2-hr fix rates) included ≥1 within 30 m of the rail (hereafter on) and 2 others that defined locations where bears effectively entered the railway (first fix off rail, next 2 on), crossed it (only the middle fix on the rail), continued along the railway (all 3 fixes on), or exited the railway corridor (first 2 on, last off). We compared both sites of higher use and each of these 4 step types to the relative frequency of bear-train collisions, predicting a positive correlation for continue step types. Relative to available locations, bears were more likely to use the railway close to railroad sidings (sections of twinned track where trains sometimes stop), at intermediate distances from human-use features (e.g., town sites, highways, trails), in areas with lower values of the compound topographic index (a proxy for wetness; within 500 m), and within 90 m of rugged terrain. Seasonally, bears made greater use of the railway in spring and fall. Among 1,515 sequences of 3 steps, crossing locations comprised >50% and were most distinct from continue locations (about 20%), which occurred in areas with more rugged terrain (within 300 m), closer to railway sidings, in spring and fall, and with steps that were 60% shorter. Contrary to our prediction, past reports of bear-train collisions were negatively correlated with continue locations and unrelated to overall use or any other movement type. Our results suggest that railway use by bears increased where it provided increased forage or easier travel, particularly in spring and fall, but more work will be needed to determine the mechanistic basis of bear-train collisions. Meanwhile, mitigation efforts such as habitat alteration or warning systems might target locations where past strikes are concentrated for grizzly bears or other sensitive populations. © 2019 The Wildlife Society.