Dale's current research program is focused on understanding the habitat requirements of several w... more Dale's current research program is focused on understanding the habitat requirements of several woodland caribou herds. He is also involved in recovery planning for numerous threatened caribou populations. Elena Jones, of Resources North, is a collaborator in all aspects of Dale's current research program.
Land managers face increasing challenges as they try to balance timber harvesting with the habita... more Land managers face increasing challenges as they try to balance timber harvesting with the habitat require- ments of wildlife, including those of woodland caribou in north-central British Columbia. With the aim of conserving caribou by improving forest practices, we employed a hierarchical, scale-explicit approach to study the processes governing movement and distribution of the northern woodland caribou ecotype. Investigations of
The Parsnip River area in British Columbia (BC), Canada, provides important habitat for grizzly b... more The Parsnip River area in British Columbia (BC), Canada, provides important habitat for grizzly bears (Ursus arctos). This area contains 2 adjacent topographic regions: (1) a relatively pristine portion of the Hart Ranges of the Canadian Rocky Mountains, and (2) a plateau on which timber harvests have occurred. Increasing human modification of both landscapes may affect the quality of grizzly
We used genetic identification of individuals and mark-recapture modeling to estimate bear (Ursus... more We used genetic identification of individuals and mark-recapture modeling to estimate bear (Ursus) population size in both a mountainous and interior plateau environment in central eastern British Columbia. We sampled 591 sites within a 9,452-km 2 study area during June and July 2000 and detected grizzly bears (Ursus arctos) at 199 sites and black bears (U. americanus) at 316 sites. We identified 326 different grizzly bears using microsatellite profiling; there were more females than males in the sample (35M:65F). After correcting for the effects of closure bias, we estimated density to be 12 grizzly bears/1,000 km 2 (95% confidence interval [CI] 7-28) on the plateau and 49/1,000 km 2 (CI 43-59) in the mountains. These densities are considerably higher than those predicted by the habitat-based extrapolation method used for grizzly bear management in British Columbia. The incorporation of hunter harvest impacts to population size caused the greatest decline, and therefore discrepancy, in numbers between the habitat-based extrapolation and the DNA-based estimate.
Wild carnivores are often exposed to diseases via contact with peridomestic host species that tra... more Wild carnivores are often exposed to diseases via contact with peridomestic host species that travel through the wildland-urban interfaces. To determine the antibody prevalences and relationships to human activity for two common canid pathogens, we sampled 99 wolves (Canis lupus) from 2000 to 2008 for antibodies to canine parvovirus (CPV) and canine distemper virus (CDV) in Banff and Jasper National Parks and surrounding areas of the Canadian Rockies. This population was the source for wolves reintroduced into the Northern Rockies of the US. Of 99 wolves sampled, 94 had detectable antibody to CPV (95%), 24 were antibody-positive for CDV (24%), and 24 had antibodies to both pathogens (24%). We tested whether antibody prevalences for CPV and CDV were higher closer to human activity (roads, town sites, First Nation reserves) and as a function of sex and age class. Wolves ≥2 yr old were more likely to be have antibodies to CPV. For CDV, male wolves, wolves ≥2 yr, and those closer to Fir...
We collected hair samples from bears and used microsatellite genotyping to identify individual be... more We collected hair samples from bears and used microsatellite genotyping to identify individual bears on three study areas near the Canadian Rocky Mountains. We estimated density of grizzly bears Ursus arctos in eight different ecosystems across five study areas, including the reanalysis of two previously published data sets. We also estimated black bear U. americanus density for two ecosystems in one study area. Grizzly bear density was lowest in boreal and subboreal plateau areas, moderate in the Rocky Mountain east slopes and highest in the Rocky Mountain west slopes. Presumably these gross differences are related to ecosystem productivity. In the Rocky Mountain west slopes, grizzly bear density was lower in populations that were partially isolated from the continuous bear population to the north. Presumably, these differences have more to do with human impacts on habitat and survival than ecosystem productivity, because productivity in partially isolated areas was similar to productivity in adjacent continuous populations. We show that large differences in bear density occur down to the ecoregion scale; broader ecosystem classes such as Banci's (1991) grizzly bear zones, ecoprovinces or ecozones would include areas with major differences in density and are therefore too coarse a scale at which to predict grizzly bear density. There appears to be little movement across ecoregion boundaries further suggesting that this may be an appropriate scale at which to extrapolate density. Differences in density across finerscale ecosystems are likely due to seasonal movements and not population level differences in density. Average bear movements were longer in less productive ecosystems. Female grizzly bears did not appear to leave their home ranges to fish for salmon Oncorhynchus spp., and extra-territorial movements by males appeared to be rare, in both ecosystems which supported spawning salmon.
Much of applied and theoretical ecology is concerned with the interactions of habitat quality, an... more Much of applied and theoretical ecology is concerned with the interactions of habitat quality, animal distribution, and population abundance. We tested a technique that uses resource selection functions (RSF) to scale animal density to the relative probability of selecting a patch of habitat. Following an accurate survey of a reference block, the habitat-based density estimator can be used to predict population abundance for other areas with no or unreliable survey data. We parameterized and tested the technique using multiple years of radiotelemetry locations and survey data collected for woodland caribou across four landscape-level survey blocks. The habitat-based density estimator performed poorly. Predictions were no better than those of a simple area estimator and in some cases deviated from the observed by a factor of 10. We developed a simulation model to investigate factors that might influence prediction success. We experimentally manipulated population density, caribou distribution, ability of animals to track carrying capacity, and precision of the estimation equation. Our simulations suggested that interactions between population density, the size of the reference block, and the pattern of distribution can lead to large discrepancies between observed and predicted population numbers. Over-or undermatching patch carrying capacity and precision of the estimator can influence predictions, but the effect is much less extreme. Although there is some empirical and theoretical evidence to support a relationship between animal abundance and resource selection, our study suggests that a number of factors can seriously confound these relationships. Habitat-based density estimators might be effective where a stable, isolated population at equilibrium is used to generate predictions for areas with similar population parameters and ecological conditions.
In North America, caribou (Rangifer tarandus) experienced diversification in separate refugia bef... more In North America, caribou (Rangifer tarandus) experienced diversification in separate refugia before the last glacial maximum. Geographical isolation produced the barren-ground caribou (Rangifer tarandus groenlandicus) with its distinctive migratory habits, and the woodland caribou (Rangifer tarandus caribou), which has sedentary behaviour and is now in danger of extinction. Herein we report on the phylogenetics, population structure, and migratory habits of caribou in the Canadian Rockies, utilizing molecular and spatial data for 223 individuals. Mitochondrial DNA analyses show the occurrence of two highly diverged lineages; the Beringian-Eurasian and North American lineages, while microsatellite data reveal that present-day Rockies' caribou populations have resulted from interbreeding between these diverged lineages. An ice-free corridor at the end of the last glaciation likely allowed, for the first time, for barren-ground caribou to migrate from the North and overlap with woodland caribou expanding from the South. The lack of correlation between nuclear and mitochondrial data may indicate that different environmental forces, which might also include human-caused habitat loss and fragmentation, are currently reshaping the population structure of this postglacial hybrid swarm. Furthermore, spatial ecological data show evidence of pronounced migratory behaviour within the study area, and suggest that the probability of being migratory may be higher in individual caribou carrying a Beringian-Eurasian haplotype which is mainly associated with the barren-ground subspecies. Overall, our analyses reveal an intriguing example of postglacial mixing of diverged lineages. In a landscape that is changing due to climatic and human-mediated factors, an understanding of these dynamics, both past and present, is essential for management and conservation of these populations.
We used resource selection functions (RSF) to estimate the relative probability of use for grizzl... more 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. 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.
Animal locations collected by Global Positioning System (GPS) collars will represent a biased sam... more Animal locations collected by Global Positioning System (GPS) collars will represent a biased sample of the sites an animal used if some position fixes fail and if those missed locations do not occur randomly. Probability of a GPS receiver obtaining a position fix is known to decline as canopy cover increases, but the impact of forest canopy cover was insufficient to account for the low fix rates we observed for GPS collars on grizzly bears (Ursus arctos). We tested the hypothesis that GPS fix rates were related to the interaction between animal activity (active vs. resting) and canopy cover by evaluating the following predictions: 1) grizzly bear activity should follow a circadian pattern similar to the circadian fix-rate pattern, 2) grizzly bear use of canopy cover should follow a circadian pattern similar to the circadian fix rates, 3) grizzly bear activity should be related to canopy cover (i.e., bears should rest in areas with relatively high canopy covers and feed and move in relatively open areas), and 4) collar orientation and canopy cover should interact to affect the fix rates of test collars. The GPS fix rates traced a bimodal circadian pattern that was directly related to the circadian pattern of grizzly bear activity. Fix rates declined when bears were more likely to be using denser cover, and fix rates of test collars demonstrated that collar orientation interacted with canopy cover, such that fix rates declined much more with increasing canopy cover when the collar was on its side than when the collar was upright. We concluded that inferences made about grizzly bear microhabitat use, based on GPS locations, will underrepresent high canopy cover sites, especially when grizzly bears are resting there.
Mountain caribou are an ecotype of woodland caribou (Rangifer tarandus caribou) that live in suba... more Mountain caribou are an ecotype of woodland caribou (Rangifer tarandus caribou) that live in subalpine forests in southeastern British Columbia, Canada, extending into northern Idaho and Washington, USA. These caribou are listed as Threatened in Canada, Endangered in the United States, and are the subject of recovery planning efforts in both countries. Many areas of mountain caribou winter habitat experience intensive use by recreational snowmobilers. During 4 surveys, we recorded caribou on all 4 census blocks with little or no snowmobile activity (x density ¼ 0.41 caribou/km 2 ), but during 3 of 4 years, we observed no caribou on the census block with intensive snowmobile activity. The year we observed caribou on the snowmobile block, most were using areas inaccessible to snowmobiles. We used a Resource Selection Function (RSF) based on radiotelemetry data for the area to compare habitat quality among the different census blocks. The absence of caribou from the intensive snowmobile area during most years could not be explained by differences in habitat quality. The RSF predicted that the intensive snowmobile area could support 53-96 caribou (95% CI). We conclude that intensive snowmobiling has displaced caribou from an area of suitable habitat. We recommend that snowmobile activity be restricted from all or most high-quality mountain caribou habitat as part of the recovery planning process. (JOURNAL OF WILDLIFE MANAGEMENT 71 :
Dale's current research program is focused on understanding the habitat requirements of several w... more Dale's current research program is focused on understanding the habitat requirements of several woodland caribou herds. He is also involved in recovery planning for numerous threatened caribou populations. Elena Jones, of Resources North, is a collaborator in all aspects of Dale's current research program.
Land managers face increasing challenges as they try to balance timber harvesting with the habita... more Land managers face increasing challenges as they try to balance timber harvesting with the habitat require- ments of wildlife, including those of woodland caribou in north-central British Columbia. With the aim of conserving caribou by improving forest practices, we employed a hierarchical, scale-explicit approach to study the processes governing movement and distribution of the northern woodland caribou ecotype. Investigations of
The Parsnip River area in British Columbia (BC), Canada, provides important habitat for grizzly b... more The Parsnip River area in British Columbia (BC), Canada, provides important habitat for grizzly bears (Ursus arctos). This area contains 2 adjacent topographic regions: (1) a relatively pristine portion of the Hart Ranges of the Canadian Rocky Mountains, and (2) a plateau on which timber harvests have occurred. Increasing human modification of both landscapes may affect the quality of grizzly
We used genetic identification of individuals and mark-recapture modeling to estimate bear (Ursus... more We used genetic identification of individuals and mark-recapture modeling to estimate bear (Ursus) population size in both a mountainous and interior plateau environment in central eastern British Columbia. We sampled 591 sites within a 9,452-km 2 study area during June and July 2000 and detected grizzly bears (Ursus arctos) at 199 sites and black bears (U. americanus) at 316 sites. We identified 326 different grizzly bears using microsatellite profiling; there were more females than males in the sample (35M:65F). After correcting for the effects of closure bias, we estimated density to be 12 grizzly bears/1,000 km 2 (95% confidence interval [CI] 7-28) on the plateau and 49/1,000 km 2 (CI 43-59) in the mountains. These densities are considerably higher than those predicted by the habitat-based extrapolation method used for grizzly bear management in British Columbia. The incorporation of hunter harvest impacts to population size caused the greatest decline, and therefore discrepancy, in numbers between the habitat-based extrapolation and the DNA-based estimate.
Wild carnivores are often exposed to diseases via contact with peridomestic host species that tra... more Wild carnivores are often exposed to diseases via contact with peridomestic host species that travel through the wildland-urban interfaces. To determine the antibody prevalences and relationships to human activity for two common canid pathogens, we sampled 99 wolves (Canis lupus) from 2000 to 2008 for antibodies to canine parvovirus (CPV) and canine distemper virus (CDV) in Banff and Jasper National Parks and surrounding areas of the Canadian Rockies. This population was the source for wolves reintroduced into the Northern Rockies of the US. Of 99 wolves sampled, 94 had detectable antibody to CPV (95%), 24 were antibody-positive for CDV (24%), and 24 had antibodies to both pathogens (24%). We tested whether antibody prevalences for CPV and CDV were higher closer to human activity (roads, town sites, First Nation reserves) and as a function of sex and age class. Wolves ≥2 yr old were more likely to be have antibodies to CPV. For CDV, male wolves, wolves ≥2 yr, and those closer to Fir...
We collected hair samples from bears and used microsatellite genotyping to identify individual be... more We collected hair samples from bears and used microsatellite genotyping to identify individual bears on three study areas near the Canadian Rocky Mountains. We estimated density of grizzly bears Ursus arctos in eight different ecosystems across five study areas, including the reanalysis of two previously published data sets. We also estimated black bear U. americanus density for two ecosystems in one study area. Grizzly bear density was lowest in boreal and subboreal plateau areas, moderate in the Rocky Mountain east slopes and highest in the Rocky Mountain west slopes. Presumably these gross differences are related to ecosystem productivity. In the Rocky Mountain west slopes, grizzly bear density was lower in populations that were partially isolated from the continuous bear population to the north. Presumably, these differences have more to do with human impacts on habitat and survival than ecosystem productivity, because productivity in partially isolated areas was similar to productivity in adjacent continuous populations. We show that large differences in bear density occur down to the ecoregion scale; broader ecosystem classes such as Banci's (1991) grizzly bear zones, ecoprovinces or ecozones would include areas with major differences in density and are therefore too coarse a scale at which to predict grizzly bear density. There appears to be little movement across ecoregion boundaries further suggesting that this may be an appropriate scale at which to extrapolate density. Differences in density across finerscale ecosystems are likely due to seasonal movements and not population level differences in density. Average bear movements were longer in less productive ecosystems. Female grizzly bears did not appear to leave their home ranges to fish for salmon Oncorhynchus spp., and extra-territorial movements by males appeared to be rare, in both ecosystems which supported spawning salmon.
Much of applied and theoretical ecology is concerned with the interactions of habitat quality, an... more Much of applied and theoretical ecology is concerned with the interactions of habitat quality, animal distribution, and population abundance. We tested a technique that uses resource selection functions (RSF) to scale animal density to the relative probability of selecting a patch of habitat. Following an accurate survey of a reference block, the habitat-based density estimator can be used to predict population abundance for other areas with no or unreliable survey data. We parameterized and tested the technique using multiple years of radiotelemetry locations and survey data collected for woodland caribou across four landscape-level survey blocks. The habitat-based density estimator performed poorly. Predictions were no better than those of a simple area estimator and in some cases deviated from the observed by a factor of 10. We developed a simulation model to investigate factors that might influence prediction success. We experimentally manipulated population density, caribou distribution, ability of animals to track carrying capacity, and precision of the estimation equation. Our simulations suggested that interactions between population density, the size of the reference block, and the pattern of distribution can lead to large discrepancies between observed and predicted population numbers. Over-or undermatching patch carrying capacity and precision of the estimator can influence predictions, but the effect is much less extreme. Although there is some empirical and theoretical evidence to support a relationship between animal abundance and resource selection, our study suggests that a number of factors can seriously confound these relationships. Habitat-based density estimators might be effective where a stable, isolated population at equilibrium is used to generate predictions for areas with similar population parameters and ecological conditions.
In North America, caribou (Rangifer tarandus) experienced diversification in separate refugia bef... more In North America, caribou (Rangifer tarandus) experienced diversification in separate refugia before the last glacial maximum. Geographical isolation produced the barren-ground caribou (Rangifer tarandus groenlandicus) with its distinctive migratory habits, and the woodland caribou (Rangifer tarandus caribou), which has sedentary behaviour and is now in danger of extinction. Herein we report on the phylogenetics, population structure, and migratory habits of caribou in the Canadian Rockies, utilizing molecular and spatial data for 223 individuals. Mitochondrial DNA analyses show the occurrence of two highly diverged lineages; the Beringian-Eurasian and North American lineages, while microsatellite data reveal that present-day Rockies' caribou populations have resulted from interbreeding between these diverged lineages. An ice-free corridor at the end of the last glaciation likely allowed, for the first time, for barren-ground caribou to migrate from the North and overlap with woodland caribou expanding from the South. The lack of correlation between nuclear and mitochondrial data may indicate that different environmental forces, which might also include human-caused habitat loss and fragmentation, are currently reshaping the population structure of this postglacial hybrid swarm. Furthermore, spatial ecological data show evidence of pronounced migratory behaviour within the study area, and suggest that the probability of being migratory may be higher in individual caribou carrying a Beringian-Eurasian haplotype which is mainly associated with the barren-ground subspecies. Overall, our analyses reveal an intriguing example of postglacial mixing of diverged lineages. In a landscape that is changing due to climatic and human-mediated factors, an understanding of these dynamics, both past and present, is essential for management and conservation of these populations.
We used resource selection functions (RSF) to estimate the relative probability of use for grizzl... more 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. 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.
Animal locations collected by Global Positioning System (GPS) collars will represent a biased sam... more Animal locations collected by Global Positioning System (GPS) collars will represent a biased sample of the sites an animal used if some position fixes fail and if those missed locations do not occur randomly. Probability of a GPS receiver obtaining a position fix is known to decline as canopy cover increases, but the impact of forest canopy cover was insufficient to account for the low fix rates we observed for GPS collars on grizzly bears (Ursus arctos). We tested the hypothesis that GPS fix rates were related to the interaction between animal activity (active vs. resting) and canopy cover by evaluating the following predictions: 1) grizzly bear activity should follow a circadian pattern similar to the circadian fix-rate pattern, 2) grizzly bear use of canopy cover should follow a circadian pattern similar to the circadian fix rates, 3) grizzly bear activity should be related to canopy cover (i.e., bears should rest in areas with relatively high canopy covers and feed and move in relatively open areas), and 4) collar orientation and canopy cover should interact to affect the fix rates of test collars. The GPS fix rates traced a bimodal circadian pattern that was directly related to the circadian pattern of grizzly bear activity. Fix rates declined when bears were more likely to be using denser cover, and fix rates of test collars demonstrated that collar orientation interacted with canopy cover, such that fix rates declined much more with increasing canopy cover when the collar was on its side than when the collar was upright. We concluded that inferences made about grizzly bear microhabitat use, based on GPS locations, will underrepresent high canopy cover sites, especially when grizzly bears are resting there.
Mountain caribou are an ecotype of woodland caribou (Rangifer tarandus caribou) that live in suba... more Mountain caribou are an ecotype of woodland caribou (Rangifer tarandus caribou) that live in subalpine forests in southeastern British Columbia, Canada, extending into northern Idaho and Washington, USA. These caribou are listed as Threatened in Canada, Endangered in the United States, and are the subject of recovery planning efforts in both countries. Many areas of mountain caribou winter habitat experience intensive use by recreational snowmobilers. During 4 surveys, we recorded caribou on all 4 census blocks with little or no snowmobile activity (x density ¼ 0.41 caribou/km 2 ), but during 3 of 4 years, we observed no caribou on the census block with intensive snowmobile activity. The year we observed caribou on the snowmobile block, most were using areas inaccessible to snowmobiles. We used a Resource Selection Function (RSF) based on radiotelemetry data for the area to compare habitat quality among the different census blocks. The absence of caribou from the intensive snowmobile area during most years could not be explained by differences in habitat quality. The RSF predicted that the intensive snowmobile area could support 53-96 caribou (95% CI). We conclude that intensive snowmobiling has displaced caribou from an area of suitable habitat. We recommend that snowmobile activity be restricted from all or most high-quality mountain caribou habitat as part of the recovery planning process. (JOURNAL OF WILDLIFE MANAGEMENT 71 :
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