Genetic structure of North American wolverine (Gulo gulo) populations

Wolverines (Gulo gulo) are found in low densities throughout their circumpolar distribution. They are also potentially susceptible to human-caused population fragmentation (development, recreation and fur harvesting). The combination of these factors has contributed to this species being listed as having either vulnerable or endangered status across much of its current range. The effects of inherently low densities and anthropogenic pressures on the genetic structure and variation of wolverine populations are, as yet, unknown. In this study, 461 individuals were typed at 12 microsatellite loci to investigate the population genetic structure of wolverines from north-western Alaska to eastern Manitoba. Levels of gene flow and population differentiation among the sampled regions were estimated via a genotype assignment test, pairwise F(ST), and two genetic distance measures. Our results suggest that wolverine populations from southernmost regions, in which anthropogenic factors are strongest, revealed more genetic structuring than did northern populations. Furthermore, these results suggest that reductions in this species' range may have led to population fragmentation in the extreme reaches of its southern distribution. The continued reduction of suitable habitat for this species may lead to more populations becoming isolated remnants of a larger distribution of northern wolverines, as documented in other North American carnivore species.     

Sources and Patterns of Wolverine Mortality in Western Montana

ABSTRACT We instrumented 36 wolverines (Gulo gulo) on 2 study areas in western Montana and one study area on the Idaho-Montana (USA) border: 14 (9 M, 5 F) on the Pioneer study area, 19 (11 M, 8 F) on the Glacier study area, and 3 (2 M, 1 F) on the Clearwater study area. During 2002–2005, harvest from licensed trapping accounted for 9 (6 M, 3 F) of 14 mortalities, including individuals from all 3 study areas. Based on Akaike's Information Criterion adjusted for small sample sizes (AIC_c) rankings of 8 a priori models, a trapping model and a trapping-by-sex interaction model were equally supported (ΔAIC_c < 2) in explaining wolverine survival. Estimated annual survival was 0.80 when we did not consider harvest, whereas additive mortality from harvest reduced annual survival to 0.57. Glacier National Park in the Glacier study area provided some refuge as evidenced by an annual survival rate of 0.77 compared to 0.51 for the Pioneer Mountain study area. We incorporated these survival rates into a simple Lefkovitch stage-based model to examine rates of population change. The finite rate of population change (Λ) for the Glacier study area was 1.1, indicating a stable to slightly increasing population, whereas Λ for the Pioneer study area was 0.7, indicating a 30% annual population decrease during our study. Changes in Λ for both study areas were most sensitive to adult survival. In 2004, we used a Lincoln Index to estimate that 12.8 ± 2.9 (95% CI) wolverines resided in the 4 mountain ranges comprising the Pioneer study area, suggesting that small, island ranges in western Montana supported few individuals. Our results suggest that if wolverines are harvested, they should be managed within individual mountain ranges or small groupings of mountain ranges to limit mortality to within biologically defined limits in recognition of the increased vulnerabilities owing to low fecundity and low population numbers in small mountain ranges. We found that refugia, such as Glacier National Park, were important by reducing trap mortality and providing immigrants to the surrounding population, but even large parks were inadequate to provide complete protection to wolverines from trapping as they ranged outside park borders.     

GPS collars have an apparent positive effect on the survival of a large carnivore

Are instrumented animals representative of the population, given the potential bias caused by selective sampling and the influence of capture, handling and wearing bio-loggers? The answer is elusive owing to the challenges of obtaining comparable data from individuals with and without bio-loggers. Using non-invasive genetic data of a large carnivore, the wolverine (Gulo gulo) in Scandinavia, and an open-population spatial capture–recapture model, we found a 16 (credible interval: 4–30) percentage points lower mortality probability for GPS-collared individuals compared with individuals without GPS collars. While the risk of dying from legal culling was comparable for collared and non-collared wolverines, the former experienced lower probability of mortality due to causes other than legal culling. The aforementioned effect was pronounced despite a potentially lower age—and therefore likely higher natural mortality—of collared individuals. Reports of positive effects of bio-loggers on the survival of individuals are uncommon and we argue that GPS collars could shield animals from poaching. Our results highlight the challenges of drawing population-level inferences for populations subjected to poaching when using data from instrumented individuals.     

The Abundance and Distribution of Wolverines in British Columbia,Canada

ABSTRACT The abundance and distribution of carnivores and their habitat are key information needed for status assessment, conservation planning, population management, and assessment of the effects of human development on their habitat and populations. We developed a habitat quality rating system, using existing wolverine (Gulo gulo) distribution, wolverine food, ecosystem mapping, and human development data. We used this and empirically derived estimates of wolverine density to predict wolverine distribution and abundance at a provincial scale. Density estimates for wolverines in high-quality habitat averaged 6.2 wolverines/1,000 km² (95% CI = 4.2–9.5). We predicted mean densities ranging from 0.3/1,000 km² in rare-quality habitat to 4.1/1,000 km² in moderate-quality habitat. Our predicted population estimate for wolverines in British Columbia was 3,530 (95% CI = 2,700-4,760). We predicted highest densities of wolverines in interior mountainous regions, moderate densities in interior plateau and boreal forest regions, and low densities in mainland coastal regions and drier interior plateaus. We predicted that wolverines would be rare on Vancouver Island, along the outer mainland coast, and in the dry interior forests, and absent from the Queen Charlotte Islands, interior grassland environments, and areas of intensive urban development.     

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.     

Wolverine Occupancy,Spatial Distribution,and Monitoring Design

In the western United States, wolverines (Gulo gulo) typically occupy high-elevation habitats. Because wolverine populations occur in vast, remote areas across multiple states, biologists have an imperfect understanding of this species' current distribution and population status. The historical extirpation of the wolverine, a subsequent period of recovery, and the lack of a coordinated monitoring program in the western United States to determine their current distribution further complicate understanding of their population status. We sought to define the limits to the current distribution, identify potential gaps in distribution, and provide a baseline dataset for future monitoring and analysis of factors contributing to changes in distribution of wolverines across 4 western states. We used remotely triggered camera stations and hair snares to detect wolverines across randomly selected 15-km × 15-km cells in Idaho, Montana, Washington, and Wyoming, USA, during winters 2016 and 2017. We used spatial occupancy models to examine patterns in wolverine distribution. We also examined the influence of proportion of the cell containing predicted wolverine habitat, human-modified land, and green vegetation, and area of the cluster of contiguous sampling cells. We sampled 183 (28.9%) of 633 cells that comprised a suspected wolverine range in these 4 states and we detected wolverines in 59 (32.2%) of these 183 sampled cells. We estimated that 268 cells (42.3%; 95% CI = 182–347) of the 633 cells were used by wolverines. Proportion of the cell containing modeled wolverine habitat was weakly positively correlated with wolverine occupancy, but no other covariates examined were correlated with wolverine occupancy. Occupancy rates (ψ) were highest in the Northern Continental Divide Ecosystem (ψ range = 0.8–1), intermediate in the Cascades and Central Mountains of Idaho (ψ range = 0.4–0.6), and lower in the Greater Yellowstone Ecosystem (ψ range = 0.1–0.3). We provide baseline data for future surveys of wolverine along with a design and protocol to conduct those surveys. © 2020 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

DNA-Based Individual and Sex Identification from Wolverine (Gulo Gulo) Faeces and Urine

Non-invasive genetic analyses are important for studies of species that are rare, sensitive or at risk of extinction. This study investigates the possibility of using faeces and urine to obtain microsatellite genotypes for individual identification of wolverines (Gulo gulo). The reliability of the employed method was assessed by analysing independent amplifications of non-invasive samples (a multiple-tube approach) and by comparing genotypes obtained from faeces to genotypes obtained from blood or tissue of the same individual. Ten microsatellite markers were successfully amplified in 65% of the faecal samples (n = 32) and 40% of the urine samples (n = 22). Allelic dropout was found in 12 and 14% of the amplifications from extracts of faeces and urine, respectively. Nevertheless, all multi-locus genotypes were correct, as judged from comparison to data from tissue or blood samples, after three replicates. These results suggest that a non-invasive approach based on DNA-analysis of faeces can be a powerful tool in population monitoring of wolverines, potentially providing reliable estimates of population size and immigration rate. A second objective of the study was to develop markers for DNA-based sex identification in wolverines using non-invasive samples. We developed two Y-linked markers, one that was specific to wolverine and one that also successfully identified sex in another mustelid. Importantly, none of the markers amplified potential prey species such as reindeer or rodents.     

Modelling broad‐scale wolverine occupancy in a remote boreal region using multi‐year aerial survey data

Aim

We used data from aerial surveys of wolverine tracks collected in seven winters over a 10‐year period (2003–2012) within a 574,287 km² study area to evaluate the broad‐scale pattern of wolverine occurrence across a remote northern boreal forest region, identifying areas of high and low occupancy.

Location

Northern Ontario, Canada.

Taxon

Wolverine (Gulo gulo Linnaeus, 1758).

Methods

We collected wolverine tracks and observations in 100‐km² hexagonal survey units, making a total of 6,664 visits to 3,039 units, visiting each 1–9 times. We used hierarchical Bayesian occupancy modelling to model wolverine occurrence, and included covariates with the potential to affect detection and/or occupancy probability of wolverines.

Results

we detected wolverines on 946 visits, 14.2% of total visits. Probability of detecting a wolverine varied among years and between the two ecozones in the study area. Wolverine occupancy was negatively related to two important covariates, the geographical coordinate Easting and thawing degree‐days. A site occupancy probability map indicated that wolverine occupancy probabilities were highest, and standard error lowest, in the western and northern portions of the study area.

Main conclusions

The occupancy framework enabled us to use observation data from tracks of this elusive, wide‐ranging carnivore over a vast, remote area while explicitly considering detectability and spatial autocorrelation, yielding a map of probable wolverine distribution in northern Ontario that would not be possible using other methods of detection across a large region. With resource development pressures increasing in this globally significant region in the face of a changing climate, it is important to monitor changes in distribution of species like wolverines that have low population growth rates, large spatial requirements and sensitivity to human disturbance. This study demonstrates a relatively cost‐effective and non‐invasive alternative to monitoring based on wolverine harvest records, which have not been available since 2009 in Ontario due to changes in the provincial regulatory regime for this threatened species.     

Genetic variation and population structure in Scandinavian wolverine (Gulo gulo) populations

Wolverine (Gulo gulo) numbers in Scandinavia were significantly reduced during the early part of the century as a result of predator removal programmes and hunting. Protective legislation in both Sweden and Norway in the 1960s and 1970s has now resulted in increased wolverine densities in Scandinavia. We report here the development of 15 polymorphic microsatellite markers in wolverine and their use to examine the population sub-structure and genetic variability in free-ranging Scandinavian wolverine populations as well as in a sample of individuals collected before 1970. Significant subdivision between extant populations was discovered, in particular for the small and isolated population of southern Norway, which represents a recent recolonization. Overall genetic variability was found to be lower than previously reported for other mustelids, with only two to five alleles per locus and observed heterozygosities (H(O)) ranging from 0.269 to 0.376 across the examined populations, being lowest in southern Norway. Analysis of the mitochondrial DNA control region revealed no variation throughout the surveyed populations. As the historical sample did not show higher levels of genetic variability, our results are consistent with a reduction in the genetic variation in Scandinavian wolverines that pre-dates the demographic bottleneck observed during the last century. The observed subdivision between populations calls for management caution when issuing harvest quotas, especially for the geographically isolated south Norwegian population.     

Evidence of a large carnivore population recovery: Counting bears in Greece

Reliable population and density estimates are the cornerstone of effective conservation and management planning, as conservation priorities often arise in relation to population numbers. Despite increased public interest and costly conservation programs limited information on brown bear (Ursus arctos, Linnaeus, 1758) abundance and density in Greece exists. We carried out systematic non-invasive genetic sampling using hair traps on power poles, as part of a capture-mark-recapture study design in order to rigorously estimate abundance and density of the Pindos bear population in Greece. From 2007–2010 we identified 211 and estimated a mean of 182.3 individuals in four sampling areas; bear densities ranged from 10.0 to 54 bears/1000 km². These results indicate an important population recovery of this large carnivore in Greece in recent years; a conservative population estimate would place the population size in the entire country >450 individuals. Considering the results of the study and the increased negative interactions between humans and bears recorded currently in Greece, we suggest that systematic genetic monitoring using power poles should continue in order to collect the necessary information that will enable the definition of an effective Action Plan for the long-term conservation of this species.     

Wolverine Conservation and Management

ABSTRACT This Special Section includes 8 peer-reviewed papers on the wolverine (Gulo gulo) in southern North America. These papers provide new information on current and historical distribution, habitat relations at multiple spatial scales, and interactions with humans. In aggregate, these papers substantially increase our knowledge of wolverine ecology and population dynamics in North America, in many cases replacing previous speculations and informed judgments with empirical information. North American wolverines occur primarily in tundra, taiga, and subalpine environments. These environments become increasingly fragmented at southern latitudes, where wolverine populations occur at low densities and are potentially vulnerable to human-caused mortality. The combination of highly fragmented habitat, demographic sensitivity to adult mortality, and low population densities make local wolverine populations difficult to monitor and easy to overharvest. Where populations are fragmented, persistence is critically dependent on dispersal between habitat islands. Although dispersal dynamics are poorly understood, high levels of genetic structure observed in both current and historical populations indicate that dispersal between mountain ranges is limited. Wolverine biology remains poorly understood, and many fundamental issues need additional research.     

Evaluation of Live-Capture Techniques for Free-Ranging Wolverines

Abstract: Wolverines (Gulo gulo) are a rare carnivore and live-capture efforts often comprise a significant component of field research projects. Wolverine studies have used aerial, snowmobile, hand-capture, and live-trap capture techniques. We reviewed existing wolverine live-capture data to evaluate sex-related biases associated with capture technique. We modified round log live traps, developed a new portable wooden live trap, and evaluated effects of live trap type, trap-site selection, and seasonal timing of trapping on wolverine capture success. Aerial capture techniques had a positive bias for capture of male wolverines. Live-capture rates were highest for portable wooden traps and lowest for barrel traps. Trapping success was highest during March when snow conditions were amenable to wolverine travel and temperatures improved bait effectiveness. Traps in corridor habitats were more successful than traps in noncorridor habitats. This difference was more pronounced in environments with rugged topography. We provide guidance for live-trap operation, describe animal handling procedures, and provide detailed instructions for construction of modified round log and portable wooden wolverine live traps. These will benefit future wolverine studies by increasing trap effectiveness and reducing risk of injury and mortality to captured wolverines.     

Noninvasive genetic sampling reveals intrasex territoriality in wolverines

Due to its conspicuous manifestations and its capacity to shape the configuration and dynamics of wild populations, territorial behavior has long intrigued ecologists. Territoriality and other animal interactions in situ have traditionally been studied via direct observations and telemetry. Here, we explore whether noninvasive genetic sampling, which is increasingly supplementing traditional field methods in ecological research, can reveal territorial behavior in an elusive carnivore, the wolverine (Gulo gulo). Using the locations of genotyped wolverine scat samples collected annually over a period of 12 years in central Norway, we test three predictions: (1) male home ranges constructed from noninvasive genetic sampling data are larger than those of females, (2) individuals avoid areas used by other conspecifics of the same sex (intrasexual territoriality), and (3) avoidance of same‐sex territories diminishes or disappears after the territory owner's death. Each of these predictions is substantiated by our results: sex‐specific differences in home range size and intrasexual territoriality in wolverine are patently reflected in the spatial and temporal configuration of noninvasively collected genetic samples. Our study confirms that wildlife monitoring programs can utilize the spatial information in noninvasive genetic sampling data to detect and quantify home ranges and social organization.     

Balancing costs and confidence: volunteer-provided point observations,GPS telemetry and the genetic monitoring of Finland’s wolves

Reliable and updated population estimates are a necessity for the successful conservation and management of endangered animal populations. Citizen science has become increasingly important in wildlife monitoring and is an attractive concept due to its low costs. However, the applicability of citizen science in the monitoring of large carnivore populations is questionable for various reasons, including the difficulties associated with species identification. In Finland, where estimates of the fragmentary wolf (Canis lupus) population have varied between 140 and 280 animals in the last 10 years, population monitoring has been based on volunteer-provided data and telemetry. To compensate for a recent decrease in the proportion of territories with boundaries mapped through telemetry, a non-invasive genetics project was launched in 2016. We evaluated the experiences from this project, in which non-invasive genetic techniques were, for the first time, widely used (n?=?22 territories, 54% of the 41 apparent territories hosted by wolves in March 2017) to determine the post-hunting population estimate in early March 2017, before pack sizes began to decrease due to dispersal by sub-adult wolves. In territories where the non-invasive genetic monitoring was executed in the winter of 2016/2017, the pack sizes resulting from the volunteer-provided observations and the genetic analyses were highly correlated. By using the most typical variation in the proportion of non-residents in the wolf populations (6–20%, Fuller et al. 2003), we derived a population estimate of 150–178 wolves for early March 2016, and, by considering the known mortality during the study period, a minimum estimate of 204–234 wolves for early Aug. 2016. Despite its high costs, we recommend that non-invasive genetic monitoring should cover all known territories supporting Finland’s small and exploited wolf population. This much costlier protocol may be unrealistic in Finland. In any case, there is a need for more genetic sampling to test the quality of volunteer-provided data.     

Contrasting levels of genetic differentiation among populations of wolverines (<Emphasis Type="Italic">Gulo gulo</Emphasis>) from northern Canada revealed by nuclear and mitochondrial loci

Habitat loss, fragmentation, overharvest, and other anthropogenic factors have resulted in population and distribution declines for North American wolverines (Gulo gulo). Currently, wolverines east of the Hudson Bay are endangered and possibly extinct, whereas the status of wolverines throughout the remaining Holarctic is vulnerable. Three previous studies using nuclear loci have detected little to no significant structuring among wolverines sampled across northern Canada. Based on these results it has been suggested that wolverines in northern Canada represent a single, panmictic population. However, as has been shown in numerous studies, in cases of female site fidelity, it is possible to have demographically autonomous populations even with male-biased gene flow. To better assess the genetic structure of wolverines in northern Canada, we examined nine microsatellite loci and DNA sequence variation from a 200 bp fragment of the mitochondrial (mtDNA) control region for 270 wolverines from nine collecting areas representing three regions of northern Canada. In agreement with previous studies, microsatellite analyses revealed a lack of significant population substructure (F _ST=0.0004). However, analysis of molecular variance, comparisons of pairwise F _ST values and nested-clade analysis of the mtDNA data revealed considerable genetic structuring among samples of wolverines from these three regions of northern Canada. These mitochondrial data provide evidence that wolverines in Canada are genetically structured due to female philopatry. The contrasting patterns of genetic differentiation based on nuclear and mitochondrial data highlight the importance of examining both nuclear and mitochondrial loci when attempting to elucidate patterns of genetic structure.     

新疆貂熊种群初步调查

正貂熊(Gulo gulo)属食肉目,鼬科,貂熊属。此种有两个亚种,分别为古北亚种(G.g.gulo)和新北亚种(G.g.luscus)。新北亚种分布在北美北部地区;古北亚种分布在欧亚大陆,我国分布为此亚种(朴仁珠和张明海,2000)。貂熊主要栖息在高山、苔原、泰加林、北方森林区域、沼泽等环境(Mitchell-Jones et al.,1999)。目前,欧洲种群     

Nonlinear responses of wolverine populations to declining winter snowpack

Understanding the population-level impacts of climate change is critical for effectively managing ecosystems. Predators are important components of many systems because they provide top−down control of community structure. Ecological theory suggests that these species could be particularly susceptible to climate change because they generally occur at low densities and have resource-limited populations. Yet, our understanding of climate-change impacts on predators is hindered by the difficulty in assessing complex, nonlinear dynamics over the large spatial scales necessary to depict a species’ general response to abiotic forcing. Here we use fur-return data to characterize population dynamics of a snow-adapted carnivore, the wolverine, across most of its North American range. Using novel modeling techniques, we simultaneously measured the impact of winter snowpack on wolverine dynamics across critical thresholds in snowpack depth and two domains of population growth. Winter snowpack declined from 1970 to 2004 in nearly the entire region studied, concordant with increases in Northern Hemisphere temperature anomalies. Fur returns have declined in many areas; our models show that snowpack has strong, nonlinear effects on wolverine population dynamics. Importantly, wolverine harvests dropped the fastest in areas where snowpack declined most rapidly and also where snowpack had the greatest effect on population dynamics. Moreover, declining snow cover appears to drive trends in wolverine population synchrony, with important implications for overall persistence. These results illustrate the vulnerability and complex responses of predator populations to climate change. We also suggest that declining snowpack may be an important and hitherto little-analyzed mechanism through which climate change alters high-latitude ecosystems.     

High mitochondrial and nuclear genetic diversity in one of the world’s most endangered seabirds,the Chatham Island Taiko (<Emphasis Type="Italic">Pterodroma magentae</Emphasis>)

Interpreting the levels of genetic diversity in organisms with diverse life and population histories can be difficult. The processes and mechanisms regulating this diversity are complex and still poorly understood. However, endangered species typically have low genetic variation as a consequence of the effects of genetic drift in small populations. In this study we examine genetic variation in the critically endangered Chatham Island Taiko (Tchaik, Pterodroma magentae), one of the world’s rarest seabirds. The Taiko has a very small population size of between 120 and 150 individuals, including just 8–15 breeding pairs. We report surprisingly high mitochondrial and nuclear genetic diversity in this critically endangered long-lived species. We hypothesise that the present Taiko population has retained a significant proportion of its past genetic diversity. However, it is also possible that undiscovered birds are breeding in unknown areas, which could increase the population size estimate. Importantly, from a conservation perspective, we show that the high level of variation is unlikely to be maintained in the future since chicks currently being born have only a limited number of the mitochondrial DNA haplotypes found in adults. Reduced genetic variation will mean that our ability to infer past events and the population history of Taiko using genetics could soon be lost and the power to determine, for example, parentage and other close order relationships will be diminished. Therefore, the maintenance of genetic diversity in future generations is an important consideration for conservation management of the Taiko.     

Inferring Geographic Isolation of Wolverines in California Using Historical DNA

ABSTRACT Delineating a species' geographic range using the spatial distribution of museum specimens or even contemporary detection-non-detection data can be difficult. This is particularly true at the periphery of a species range where species' distributions are often disjunct. Wolverines (Gulo gulo) are wide-ranging mammals with discontinuous and potentially isolated populations at the periphery of their range. One potentially disjunct population occurred in the Sierra Nevada Mountains, California, USA, and appears to have been extirpated by the 1930s. Many early 20th century naturalists believed that this population was connected to other populations occurring in the Cascade Range of northern California, Oregon, and Washington, USA, but a recent analysis of historical records suggests that California wolverines were isolated from other populations in North America. We used DNA extracted from museum specimens to examine whether California wolverines were isolated. Both nuclear and mitochondrial DNA data indicate that California wolverines were genetically distinct from extant populations, suggesting long-term isolation. We identified 2 new control region (mitochondrial DNA) haplotypes located only within California. We used these data and referenced sequences from the Rocky Mountains, USA, to make inferences regarding potential wolverine translocations into California. In addition, we used these genetic data to make inferences about wolverine conservation throughout western North America.     

Multiscale Habitat Use by Wolverines in British Columbia,Canada

ABSTRACT Wolverine (Gulo gulo) distribution in British Columbia, Canada, includes multiple-use lands where human use and resource extraction may influence habitat selection. We evaluated seasonal habitat use by resident adult wolverines using radiotelemetry locations from 2 multiple-use landscapes in British Columbia. Food, predation risk, and human disturbance hypotheses were considered in logistic regression analyses of used and random landscapes. Male wolverine habitat associations were most supported by the food hypothesis in both summer and winter. Moose (Alces alces) winter ranges, valley bottom forests, and avalanche terrain were positively associated with winter male wolverine use. Habitat use by male wolverines in winter was also negatively associated with helicopter skiing areas in the Columbia Mountains. Habitat associations of females were more complex; combinations of variables supporting food, predation risk, or human disturbance hypotheses were included in most supported models from both summer and winter in both study areas. Females were associated with alpine and avalanche environments where hoary marmot (Marmota caligata) and Columbia ground squirrel (Spermophilus columbianus) prey are found in summer. Roaded and recently logged areas were negatively associated with female wolverines in summer. In the Columbia Mountains, where winter recreation was widespread, females were negatively associated with helicopter and backcountry skiing. Moose winter ranges within rugged landscapes were positively associated with females during winter. Our analysis suggests wolverines were negatively responding to human disturbance within occupied habitat. The population consequences of these functional habitat relationships will require additional focused research. Our spatially explicit models can be used to support conservation planning for resource extraction and tourism industries operating in landscapes occupied by wolverines.