Coarse-Scale Distribution Surveys and Occurrence Probability Modeling for Wolverine in Interior Alaska

ABSTRACT We determined wolverine (Gulo gulo) distribution and occurrence probabilities using aerial surveys and hierarchical spatial modeling in a 180,000-km² portion of Interior Alaska, USA. During 8 February-12 March 2006, we surveyed 149 of 180 1,000-km² sample units for wolverine tracks. We observed wolverine tracks in 99 (66.4%) sample units. Wolverine detection probability was ≥69% throughout the survey period. Posterior occurrence probabilities of whether a wolverine track occurred in a sample unit was dependent on survey timing, number of transects flown, number of neighboring sample units with detected tracks, percentage of the sample unit with elevation ≤305 m, and human influences. Our model indicated strong evidence of occurrence (>0.80) in 72% of the 180 survey units, strong evidence of absence (<0.20) in 12%, and weak evidence of occurrence or absence (0.20–0.80) in 16%. Wolverine area of occupancy made up 83% of the study area. Simulations illustrated that 2–4 survey routes were necessary for the survey technique to provide strong evidence of wolverine presence or absence in Interior Alaska if a track was not identified along the first route. The necessary number of survey routes depends on the occurrence probability in a sample unit. We provided managers with a map of wolverine distribution in Interior Alaska and an efficient and lower-cost method to detect coarse-scale changes in wolverine distribution. Our technique was effective in both Interior Alaska and Ontario, Canada, suggesting it would be effective throughout most of the boreal forest range of wolverines where tracks can be readily observed from the air. The technique requires a certain skill level in recognizing tracks; it is essential that tracks are identified correctly and training may be necessary depending on surveyor experience.     

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.     

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.     

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.     

Distribution and Broadscale Habitat Relations of the Wolverine in the Contiguous United States

ABSTRACT Conservation of the wolverine (Gulo gulo) at the southern extent of its North American range requires reliable understandings of past and present distribution patterns and broad-scale habitat relations. We compiled 820 verifiable and documented records of wolverine occurrence (specimens, DNA detections, photos, and accounts of wolverines being killed or captured) in the contiguous United States from museums, the literature, and institutional archives. We spatially referenced 729 records with areal precision ≤1 township (93.2 km²) and temporal precision ≤10 years. Historical records (1827–1960) were located primarily in the western mountains and Great Lakes region. However, our data suggest that the historical distribution of wolverines in the Cascade Range and Sierra Nevada was disjunct, contradicting previous interpretations. Our results indicate that wolverine range in the contiguous United States had contracted substantially by the mid-1900s. Current records (1995–2005) are limited to north-central Washington, northern and central Idaho, western Montana, and northwestern Wyoming. We investigated potential relations between wolverines and alpine vegetation, cold temperatures, and spring snow cover by comparing the distribution of historical wolverine records with Kuchler's potential natural vegetation types, Holdridge's climatic life zones, and EASE snow-cover maps during the latter portion of the wolverine denning period (15 Apr-14 May). In the western mountains, historical wolverine records generally occurred in or near alpine vegetation and climatic conditions, especially at the limits of their distribution in the Cascade Range, Sierra Nevada, and southern Rocky Mountains. However, the only habitat layer that fully accounted for historical distribution patterns was spring snow cover. Causal factors for the extirpation of wolverines from the southern portions of their range in the contiguous United States are unknown, but are likely related to high levels of human-caused mortality and low to nonexistent immigration rates.     

The Sustainability of Wolverine Trapping Mortality in Southern Canada

Range declines, habitat connectivity, and trapping have created conservation concern for wolverines throughout their range in North America. Previous researchers used population models and observed estimates of survival and reproduction to infer that current trapping rates limit population growth, except perhaps in the far north where trapping rates are lower. Assessing the sustainability of trapping requires demographic and abundance data that are expensive to acquire and are therefore usually only achievable for small populations, which makes generalization risky. We surveyed wolverines over a large area of southern British Columbia and Alberta, Canada, used spatial capture-recapture models to estimate density, and calculated trapping kill rates using provincial fur harvest data. Wolverine density averaged 2 wolverines/1,000 km² and was positively related to spring snow cover and negatively related to road density. Observed annual trapping mortality was >8.4%/year. This level of mortality is unlikely to be sustainable except in rare cases where movement rates are high among sub-populations and sizable un-trapped refuges exist. Our results suggest wolverine trapping is not sustainable because our study area was fragmented by human and natural barriers and few large refuges existed. We recommend future wolverine trapping mortality be reduced by ≥50% throughout southern British Columbia and Alberta to promote population recovery. © 2019 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Cumulative effects of climate and landscape change drive spatial distribution of Rocky Mountain wolverine (Gulo gulo L.)

Contemporary landscapes are subject to a multitude of human‐derived stressors. Effects of such stressors are increasingly realized by population declines and large‐scale extirpation of taxa worldwide. Most notably, cumulative effects of climate and landscape change can limit species’ local adaptation and dispersal capabilities, thereby reducing realized niche space and range extent. Resolving the cumulative effects of multiple stressors on species persistence is a pressing challenge in ecology, especially for declining species. For example, wolverines (Gulo gulo L.) persist on only 40% of their historic North American range. While climate change has been shown to be a mechanism of range retractions, anthropogenic landscape disturbance has been recently implicated. We hypothesized these two interact to effect declines. We surveyed wolverine occurrence using camera trapping and genetic tagging at 104 sites at the wolverine range edge, spanning a 15,000 km² gradient of climate, topographic, anthropogenic, and biotic variables. We used occupancy and generalized linear models to disentangle the factors explaining wolverine distribution. Persistent spring snow pack—expected to decrease with climate change—was a significant predictor, but so was anthropogenic landscape change. Canid mesocarnivores, which we hypothesize are competitors supported by anthropogenic landscape change, had comparatively weaker effect. Wolverine population declines and range shifts likely result from climate change and landscape change operating in tandem. We contend that similar results are likely for many species and that research that simultaneously examines climate change, landscape change, and the biotic landscape is warranted. Ecology research and species conservation plans that address these interactions are more likely to meet their objectives.     

Spatial ecology of wolverines at the southern periphery of distribution

Wolverines (Gulo gulo) in the conterminous United States have experienced range contraction, are uncommon, and have been designated as warranted for protection under the United States Endangered Species Act. Data from the southern edge of the wolverine's circumpolar distribution is sparse, and development of effective conservation strategies would benefit from a more complete understanding of the species' ecology. We captured and radio-monitored 30 wolverines in the Greater Yellowstone Ecosystem (GYE), tested for seasonal habitat selection by elevation band, and examined a suite of spatial characteristics to clarify our understanding of the wolverine's niche. Wolverines in GYE selected for areas >2,600 m latitude-adjusted elevation (LAE; n = 2,257 wolverine locations [12 F, 6 M]). Wolverines avoided areas <2,150 m LAE, including during winter when the vast majority of ungulates are pushed to these elevations by deep snow. Wolverine home ranges were large relative to body size, averaging 303 km² for adult females and 797 km² for adult males (n = 13 [8 F, 5 M] and 33 wolverine-years). Resident adults fit with Global Positioning System (GPS) collars used an area >75% the size of their multi-year home range in an average of 32 days (n = 7 [5 F, 2 M]). Average movement rates of 1.3 km/2-hr indicated that both sexes move distances equivalent to the diameter of their home range every 2 days or the circumference of their home range in <1 week (n = 1,329 2-hr movements, n = 12 individuals [7 F, 5 M]). This capability for movement, the short time-frame over which home ranges were developed, and a lack of home range overlap by same sex adults ( , 90% CI = 0.0–4.8%, n = 22 pairs) suggested territoriality. We estimated wolverine density to be 3.5/1,000 km² of area >2,150 m LAE (95% CI = 2.8–9.6). Dispersal movements extended to at least 170 km for both sexes (n = 5 F, 2 M). At the southern edge of distribution, where suitable and unsuitable conditions exist in close proximity, wolverines selected high-elevation areas near alpine tree-line where a mix of forest, meadow, and boulder fields were present, deep snow-cover existed during winter, and low temperatures near freezing can occur throughout the year. Persistence in these areas where the growing season is brief requires large home ranges that are regularly patrolled, a social system that provides exclusive access to resources, and low densities. These characteristics, along with low reproductive rates, are prevalent throughout the species range, indicating that wolverines are specialists at exploiting a cold, unproductive niche where interspecific competition is limited. The vulnerability inherent in occupying this unproductive niche was likely influential in previous declines within the conterminous United States and will remain a factor as wolverines encounter modern human influences. Conserving wolverines in the conterminous United States will require collaborative management over a large geographic scale. © 2011 The Wildlife Society.     

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.     

The diet of breeding female wolverines (Gulo gulo) in two areas of Finland

The wolverine (Gulo gulo) hunts and scavenges for food. We examined how the presence of a semi-domesticated reindeer (Rangifer tarandus tarandus) influences the diet of breeding female wolverines in Finland. We compared the dietary composition of wolverines breeding in the reindeer management area (northern Finland) with that of wolverines breeding outside this area (eastern Finland) by analyzing 421 scats collected from six active dens in northern Finland and four dens in eastern Finland during 2004–2006. The occurrence of food sources in the diet was assessed for both areas using logistic mixed-effects models. Diet breadth per study area was also calculated. As expected, semi-domesticated reindeer was the most important food source (66 %) in northern Finland. Mountain hares (Lepus timidus) were the second most important food item (16 %). Wolverines also utilized grouse (7 %), small rodents (6 %), and moose (Alces alces; 4 %) in their diet. In eastern Finland, the most utilized prey species was moose (55 %), while the diet also comprised mountain hares (13 %), domestic pigs (Sus scrofa; 13 %), grouse (11 %), and small rodents (2 %). The diet of wolverines in eastern Finland included more moose and less semi-domesticated reindeers and small rodents than in northern Finland. The niche breadth did not differ between the study areas. In eastern Finland, the diet of wolverines shifted to 55 % moose and 0 % semi-domesticated reindeer; compared to 4 % moose and 66 % semi-domesticated reindeer in northern Finland. Our study highlights the important role of ungulates in the diet of breeding female wolverines. In areas with a low density of medium-sized ungulates, scavenging for wolf- and human-killed moose and carcasses on feeding sites plays an essential role in food acquisition by wolverines.     

Space use and territoriality of wolverines (<Emphasis Type="Italic">Gulo gulo</Emphasis>) in northern Scandinavia

Space use and territoriality influence population structure and dynamics and is therefore an important aspect in understanding the ecology of animals. We investigated spatial and temporal space use of wolverines (Gulo gulo) in northern Scandinavia. We estimated home ranges of 24 radio-marked individuals (17 females and seven males). Male home ranges (mean 669 km²; SE = 211) were significantly larger than female home ranges (mean 170 km²; Wilcoxon–Mann–Whitney; P = 0.001) and encompassed or included parts of up to five different females. Home range sizes of reproducing (170 km²; SE = 51) and barren (171 km²; SE = 63) adult females did not differ. Wolverines in Scandinavia exhibit intrasexual territoriality, with male home ranges totally exclusive and female home ranges either exclusive or with little home range overlap. Overlap between wolverine territories is most likely explained by intrasexual tolerance and kinship.     

Can Occupancy–Abundance Models Be Used to Monitor Wolf Abundance?

Estimating the abundance of wild carnivores is of foremost importance for conservation and management. However, given their elusive habits, direct observations of these animals are difficult to obtain, so abundance is more commonly estimated from sign surveys or radio-marked individuals. These methods can be costly and difficult, particularly in large areas with heavy forest cover. As an alternative, recent research has suggested that wolf abundance can be estimated from occupancy–abundance curves derived from “virtual” surveys of simulated wolf track networks. Although potentially more cost-effective, the utility of this approach hinges on its robustness to violations of its assumptions. We assessed the sensitivity of the occupancy–abundance approach to four assumptions: variation in wolf movement rates, changes in pack cohesion, presence of lone wolves, and size of survey units. Our simulations showed that occupancy rates and wolf pack abundances were biased high if track surveys were conducted when wolves made long compared to short movements, wolf packs were moving as multiple hunting units as opposed to a cohesive pack, and lone wolves were moving throughout the surveyed landscape. We also found that larger survey units (400 and 576 km²) were more robust to changes in these factors than smaller survey units (36 and 144 km²). However, occupancy rates derived from large survey units rapidly reached an asymptote at 100% occupancy, suggesting that these large units are inappropriate for areas with moderate to high wolf densities (>15 wolves/1,000 km²). Virtually-derived occupancy–abundance relationships can be a useful method for monitoring wolves and other elusive wildlife if applied within certain constraints, in particular biological knowledge of the surveyed species needs to be incorporated into the design of the occupancy surveys. Further, we suggest that the applicability of this method could be extended by directly incorporating some of its assumptions into the modelling framework.     

Seasonal Habitat Associations of the Wolverine in Central Idaho

ABSTRACT Although understanding habitat relationships remains fundamental to guiding wildlife management, these basic prerequisites remain vague and largely unstudied for the wolverine. Currently, a study of wolverine ecology conducted in Montana, USA, in the 1970s is the sole source of information on habitat requirements of wolverines in the conterminous United States. The Montana study and studies conducted in Canada and Alaska report varying degrees of seasonal differences in wolverine habitat use. This article provides an empirical assessment of seasonal wolverine habitat use by 15 wolverines (Gulo gulo) radiotracked in central Idaho, USA, in 1992–1996. We controlled for radiotelemetry error by describing the probability of each location being in a habitat cover type, producing a vector of cover type probabilities suited for resource selection analysis within a logistic regression framework. We identified variables that were important to presence of wolverines based on their strength (significance) and consistency (variability in coeff. sign) across all possible logistic regression models containing 9 habitat cover types and 3 topographic variables. We selected seasonal habitat models that incorporated those variables that were strong and consistent, producing a subset of potential models. We then ranked the models in this subset based on Akaike's Information Criterion and goodness-of-fit. Wolverines used modestly higher elevations in summer versus winter, and they shifted use of cover types from whitebark pine (Pinus albicaulis) in summer to lower elevation Douglas fir (Pseudotsuga menziezii) and lodgepole pine (Pinus contorta) communities in winter. Elevation explained use of habitat better than any other variable in both summer and winter. Grass and shrub habitats and slope also had explanatory power. Wolverines preferred northerly aspects, had no attraction to or avoidance of trails during summer, and avoided roads and ungulate winter range. These findings improve our understanding of wolverine presence by demonstrating the importance of high-elevation subalpine habitats to central Idaho wolverines.     

Diet shift of a facultative scavenger, the wolverine, following recolonization of wolves

1. Wolves Canis lupus L. recolonized the boreal forests in the southern part of the Scandinavian peninsula during the late 1990s, but so far there has been little attention to its effect on ecosystem functioning. Wolf predation increases the availability of carcasses of large prey, especially moose Alces alces L., which may lead in turn to a diet switch in facultative scavengers such as the wolverine Gulo gulo L. 2. Using 459 wolverine scats collected during winter-spring 2001-04 for DNA identity and dietary contents, we compared diet inside and outside wolf territories while controlling for potential confounding factors, such as prey density. We tested the hypothesis that wolverine diet shifted towards moose in the presence of wolves, while taking into account possible sexual segregation between the sexes. Occurrence of reindeer, moose and small prey was modelled against explanatory covariates using logistic mixed-effects models. Furthermore, we compared diet composition and breadth among habitats and sexes. 3. Occurrence of reindeer, moose and small prey in the diet varied with prey availability and habitat. As expected, diet contained more moose and less reindeer and small prey in the presence of wolves. Their diet in tundra consisted of 40% reindeer Rangifer tarandus L., 39% moose and 9% rodents. In forest with wolf, their diet shifted to 76% moose, 18% reindeer and 5% rodents; compared to 42% moose, 32% reindeer and 15% rodents in forest without wolf. This diet switch could not be explained by higher moose density in wolf territories. Female diet consisted of more small prey than for males, but there was a tendency for females to use the highly available moose carrion opportunistically and to hunt less on small prey within wolf territories. 4. Our study highlights how wolves increase scavenging opportunities for wolverines, and how sexual differences in diet may also apply to large scavengers. Due to their more restricted home range, female wolverines are forced to rely more on hunting small prey. The relatively high occurrence of wolf kills, however, forms an important food source to wolverines in this area. The recolonization of wolves may therefore have contributed to the consequent recolonization of wolverines into the same area.     

Integrating sign surveys and telemetry data for estimating brown bear (Ursus arctos) density in the Romanian Carpathians

Accurate population size estimates are important information for sustainable wildlife management. The Romanian Carpathians harbor the largest brown bear (Ursus arctos) population in Europe, yet current management relies on estimates of density that lack statistical oversight and ignore uncertainty deriving from track surveys. In this study, we investigate an alternative approach to estimate brown bear density using sign surveys along transects within a novel integration of occupancy models and home range methods. We performed repeated surveys along 2‐km segments of forest roads during three distinct seasons: spring 2011, fall‐winter 2011, and spring 2012, within three game management units and a Natura 2000 site. We estimated bears abundances along transects using the number of unique tracks observed per survey occasion via N‐mixture hierarchical models, which account for imperfect detection. To obtain brown bear densities, we combined these abundances with the effective sampling area of the transects, that is, estimated as a function of the median (± bootstrapped SE) of the core home range (5.58 ± 1.08 km²) based on telemetry data from 17 bears tracked for 1‐month periods overlapping our surveys windows. Our analyses yielded average brown bear densities (and 95% confidence intervals) for the three seasons of: 11.5 (7.8–15.3), 11.3 (7.4–15.2), and 12.4 (8.6–16.3) individuals/100 km². Across game management units, mean densities ranged between 7.5 and 14.8 individuals/100 km². Our method incorporates multiple sources of uncertainty (e.g., effective sampling area, imperfect detection) to estimate brown bear density, but the inference fundamentally relies on unmarked individuals only. While useful as a temporary approach to monitor brown bears, we urge implementing DNA capture–recapture methods regionally to inform brown bear management and recommend increasing resources for GPS collars to improve estimates of effective sampling area.     

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.     

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.     

Combining Occurrence and Habitat Suitability Data Improve Conservation Guidance for the Giant Kangaroo Rat

Identifying high-quality habitat (i.e., areas with resources and conditions suitable to support long-term species persistence) is a priority for conservation, but estimating habitat quality is expensive and time consuming. Instead managers often rely on occurrence data or models of habitat suitability, but these data are only proximally related to individual and population persistence on the landscape. In most habitat suitability modeling studies, researchers treat the model as a hypothesis and the occurrence data as the truth. But occurrence does not always correlate with habitat as expected; therefore, occurrence data may be unreliable. We propose that suitability models and occurrence data be given equal weight to highlight areas of disagreement for future demographic study. To highlight this approach, we used the giant kangaroo rat (Dipodomys ingens) as a case study because their distinct burrow mounds allow for remote monitoring of short-term presence and long-term persistence. We conducted trapping, manned aerial surveys, and aerial imagery surveys in the San Joaquin Desert in California, USA, between 2001 and 2017 and compared the results to an existing habitat suitability model to provide estimates of long-term persistence based on the presence of burrow mounds made by giant kangaroo rats. We treated areas of positive agreement as priorities for habitat conservation and areas of negative agreement as areas managers could ignore. Remaining areas should be prioritized for additional occupancy and demographic studies. From an initial area of 17,385 km², we identified 668 km² of currently occupied high-quality habitat. Of this, just 135 km² was on private land and therefore requiring protection. We classified 1,498 km² (8.6%) for additional research. Of that area, 744 km² was flagged for additional occupancy surveys. Our 3 data sets disagreed over 754 km², suggesting a need for further demographic studies to reveal important population-habitat relationships for the species in those areas. This approach can be useful as part of any habitat conservation exercise for prioritizing protection or targeting future demographic studies. © 2021 The Wildlife Society.     

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.     

Integrating motion-detection cameras and hair snags for wolverine identification

We developed an integrated system for photographing a wolverine's (Gulo gulo) ventral pattern while concurrently collecting hair for microsatellite DNA genotyping. Our objectives were to 1) test the system on a wild population of wolverines using an array of camera and hair-snag (C&H) stations in forested habitat where wolverines were known to occur, 2) validate our ability to determine identity (ID) and sex from photographs by comparing photographic data with that from DNA, and 3) encourage researchers and managers to test the system in different wolverine populations and habitats and improve the system design. Of the 18 individuals (10 M, 8 F) for which we obtained genotypes over the 2 years of our study, there was a 100% match between photographs and DNA for both ID and sex. The integrated system made it possible to reduce cost of DNA analysis by >74%. Integrating motion-detection cameras and hair snags provides a cost-effective technique for wildlife managers to monitor wolverine populations in remote habitats and obtain information on important population parameters such as density, survival, productivity, and effective population size. © 2011 The Wildlife Society.