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Kyle Joly

Publications and source records attributed to Kyle Joly.

At least 19 recordsLinked to original sources

Validating antler- and movement-based estimates of caribou reproduction using video camera collars

Wildlife agencies invest substantial resources in monitoring ungulate reproductive rates given ongoing concerns about population trends and their implications for management. For barren-ground caribou ( Rangifer tarandus ) in the North American Arctic, data on parturition and neonate survival have traditionally been collected using visual observations of antler retention and calf presence from small, fixed-wing aircraft. However, these surveys are weather-dependent, expensive, and dangerous, motivating the development of movement-based approaches that infer reproductive events from global positioning system (GPS) location data. We had the unique opportunity to use video camera collars to validate both antler- and movement-based methods for estimating reproduction in barren-ground caribou, using data from the Porcupine (2018–2023) and Western Arctic (2021, 2023) herds in Alaska, USA, and Yukon, Canada. For the movement-based approaches, we assessed individual-based and population-based methods, which both inferred parturition following a sharp decline in movement and inferred calf loss following a sharp increase in movement. We investigated these approaches using rarified 1-, 2-, 4-, and 8-hour fix intervals. Based on video collar observations, we found that antler retention at the onset of the calving period was highly accurate for predicting parturition (96% accuracy) but became unreliable as calving progressed, as approximately 80% of parturient females shed their antlers during the calving period. In contrast, both movement methods performed marginally for estimating caribou parturition (~70% accuracy), with estimates from the individual-based method exhibiting greater bias than those from the population-based method. The individual-based method also poorly predicted neonate survival (≤49% accuracy for detecting parturition and calf fate for the Porcupine Herd). Video data revealed that inaccuracies with the movement methods often occurred when caribou bedded during storms, rested after traversing mountainous terrain, lost a calf shortly after birth, or increased movement to evade insects. These results have important implications for management agencies using antler- and movement-based methods to estimate caribou reproduction, particularly when informing assessments of population decline or recovery. Our results demonstrate the utility of video collar data for validating reproductive monitoring approaches and provide insights into how these approaches can be improved.

Alaska, Yukon↗

An energetic tradeoff best explains parturition timing in grizzly bears

Timing of grizzly bear ( Ursus arctos ) parturition during hibernation has been explained by ancestral traits (delayed implantation, altricial young, obligate maternal denning), but the ultimate driver underlying precise timing has not been fully explored. Capitalizing on an observed latitudinal increase in denning duration among four populations in interior North America, we tested two alternative hypotheses. First, that birth timing results from a physiological cue that synchronizes implantation with the onset of hibernation, allowing females to forgo reproduction should they lack adequate fat stores. Alternatively, that parturition is optimally timed relative to den exit to balance an energetic tradeoff between minimizing lactation time to protect the mother and maximizing developmental time to increase cub survival. Using parturition dates previously predicted from accelerometer data (27 Dec–28 Feb), we classified 115 females according to apparent litter survival when first visually observed after den exit: 57% successful (with cubs), 22% unsuccessful (alone), and 21% unknown (not observed). The number of days between birth and den exit showed no association with latitude ( p = 0.29). It averaged 103 days among successful females but only 77 days among unsuccessful females ( p < 0.001) owing to later births and earlier exit. With each increasing degree of latitude, birth date increased by 1.0 and number of days between den entry and birth increased by 2.5 ( p < 0.001). Implantation dates were not centered on den entry dates ( p < 0.001). These results supported the energetic tradeoff hypothesis and suggested natural selection has favored a consistent number of days between parturition and den exit under average body conditions and shifts toward later or earlier births for females with lower or higher levels of bodily stored energy, respectively. This flexible tradeoff may support resilience to climate change and present a possible mechanism explaining reduced natality and cub survival in high-density populations.

Alaska, British Columbia, Idaho, Montana, Wyoming↗

Detecting occurrence and timing of grizzly bear parturition based on anomalies in accelerometer data

Documenting natality of brown (grizzly) bears ( Ursus arctos ) is an important component of many population monitoring and research programs. Brown bears give birth during hibernation, so observation of litters is generally not feasible until females exit dens, and detection of litters can be compromised by poor observability. Using triaxial accelerometer activity data stored on radiocollars, we developed a technique that predicts births by the presence of brief upsurges in activity likely associated with postnatal maternal behaviors. We developed presence criteria with a training sample of known parturient females ( n = 22) to detect anomalies within time-series of daily motion-counts (i.e., activity readings >0 measured at 10-min intervals) during 25 December–7 March. To test performance, we applied criteria to a blinded sample of activity data obtained from female grizzly bears (captured and collared during 2012–2022) in 4 populations in interior North America ( n = 295). We assigned predicted status and compared assignments to reproductive status at first visual observation. The true positive rate was 91% ( n = 47 females observed with cubs of the year) and the false-positive rate was 17% ( n = 65 females observed with older offspring). Births were predicted for 50% of females with unknown reproductive status ( n = 48), 21% of females observed without offspring ( n = 114), and 10% of subadult females considered too young to reproduce ( n = 21). Dates of predicted births varied from 27 December to 28 February. Our anomaly detection technique was successful at estimating parturition events, and despite some error, indicated that a number of litters were born but not observed, presumably due to mortality in or shortly after den emergence. This technique provides an additional tool for supplementing visual observations for natality estimation and population modeling, provided that potential biases stemming from increased detection rates are considered.

Alaska, British Columbia, Idaho, Montana↗

The influence of human presence and footprint on animal space use in US national parks

Given the importance of protected areas for biodiversity, the growth of visitation to many areas has raised concerns about the effects of humans on wildlife. In 2020, the COVID-19 pandemic led to temporary closure of national parks in the United States, offering a pseudonatural experiment to tease apart the effects of permanent infrastructure and transient human presence on animals. We compiled GPS tracking data from 229 individuals of 10 mammal species in 14 parks and used third-order hierarchical resource selection functions to evaluate the influence of the human footprint on animal space use in 2019 and 2020. Averaged across all parks and species, animals avoided the human footprint, whether the park was open or closed. However, although animals in remote areas showed consistent avoidance, on average those in more developed areas switched from avoidance to selection when protected areas were closed. Findings varied across species: some responded consistently negatively to the footprint (wolves, mountain goats), some positively (mule deer, red fox) and others had a strong exposure-mediated response (elk, mountain lion). Furthermore, some species responded more strongly to the park closure (black bear, moose). This study advances our understanding of complex interactions between recreation and wildlife in protected areas.

western United States↗

Intrinsic and environmental drivers of pairwise cohesion in wild Canis social groups

Animals within social groups respond to costs and benefits of sociality by adjusting the proportion of time they spend in close proximity to other individuals in the group (cohesion). Variation in cohesion between individuals, in turn, shapes important group-level processes such as subgroup formation and fission–fusion dynamics. Although critical to animal sociality, a comprehensive understanding of the factors influencing cohesion remains a gap in our knowledge of cooperative behavior in animals. We tracked 574 individuals from six species within the genus Canis in 15 countries on four continents with GPS telemetry to estimate the time that pairs of individuals within social groups spent in close proximity and test hypotheses regarding drivers of cohesion. Pairs of social canids ( Canis spp.) varied widely in the proportion of time they spent together (5%–100%) during seasonal monitoring periods relative to both intrinsic characteristics and environmental conditions. The majority of our data came from three species of wolves (gray wolves, eastern wolves, and red wolves) and coyotes. For these species, cohesion within social groups was greatest between breeding pairs and varied seasonally as the nature of cooperative activities changed relative to annual life history patterns. Across species, wolves were more cohesive than coyotes. For wolves, pairs were less cohesive in larger groups, and when suitable, small prey was present reflecting the constraints of food resources and intragroup competition on social associations. Pair cohesion in wolves declined with increased anthropogenic modification of the landscape and greater climatic variability, underscoring challenges for conserving social top predators in a changing world. We show that pairwise cohesion in social groups varies strongly both within and across Canis species, as individuals respond to changing ecological context defined by resources, competition, and anthropogenic disturbance. Our work highlights that cohesion is a highly plastic component of animal sociality that holds significant promise for elucidating ecological and evolutionary mechanisms underlying cooperative behavior.

Ecology↗

A test of the green wave hypothesis in omnivorous brown bears across North America

Herbivorous animals tend to seek out plants at intermediate phenological states to improve energy intake while minimizing consumption of fibrous material. In some ecosystems, the timing of green-up is heterogeneous and propagates across space in a wave-like pattern, known as the green wave. Tracking the green wave allows individuals to prolong access to higher-quality forage. While there is a plethora of empirical support for such behavior in herbivorous taxa, the green wave hypothesis (GWH) is nuanced based on factors such as body morphometrics and digestive capacity. Furthermore, little is known about whether other taxa, such as omnivores, track the green wave. Our objective was to assess whether the GWH can be extended to explain the movements of omnivores. Using GPS collar data from seven populations (n = 127 individuals) of brown bears Ursus arctos across their entire North American range, we first tested whether bears tracked the green wave. Using conditional resource selection functions (RSFs), we found that variation in proxies of vegetative forage quality better explained movement and habitat selection than proxies of forage biomass in over half of the bears in our study, providing evidence of green wave tracking. Second, we assess factors that explained variation in green wave tracking using linear mixed effects models. Green wave tracking in brown bears was explained by the variation in availability of green-up within spring home ranges, and how green-up transitioned across those home ranges. Our results demonstrate that the GWH can partially explain movement of a non-migratory omnivorous species, extending the generality of the GWH as a broad predictor of animal space use. The green wave is another resource wave brown bears track, and our findings help predict brown bear space use, which can be used to guide conservation and habitat restoration efforts.

Alaska, British Columbia, Idaho, Montana, Wyoming↗

Survey for selected parasites in Alaska brown bears (Ursus arctos)

To assess infection with or exposure to endo- and ectoparasites in Alaska brown bears ( Ursus arctos ), blood and fecal samples were collected during 2013–17 from five locations: Gates of the Arctic National Park and Preserve; Katmai National Park; Lake Clark National Park and Preserve; Yakutat Forelands; and Kodiak Island. Standard fecal centrifugal flotation was used to screen for gastrointestinal parasites, molecular techniques were used to test blood for the presence of Bartonella and Babesia spp., and an ELISA was used to detect antibodies to Sarcoptes scabiei , a species of mite recently associated with mange in American black bears ( Ursus americanus ). From fecal flotations ( n =160), we identified the following helminth eggs: Uncinaria sp. ( n =16, 10.0%), Baylisascaris sp. ( n =5, 3.1%), Dibothriocephalus sp. ( n =2, 1.2%), and taeniid-type eggs ( n =1, 0.6%). Molecular screening for intraerythrocytic parasites ( Babesia spp.) and intracellular bacteria ( Bartonella spp.) was negative for all bears tested. We detected antibodies to S. scabiei in six of 59 (10.2%) individuals. The relatively low level of parasite detection in this study meets expectations for brown bear populations living in large, relatively undisturbed habitats near the northern edge of the range. These results provide a contemporary understanding of parasites in Alaska brown bears and establish baseline levels of parasite presence to monitor for changes over time and relative to ecologic alterations.

Alaska↗

Mapping out a future for ungulate migrations

Migration of ungulates (hooved mammals) is a fundamental ecological process that promotes abundant herds, whose effects cascade up and down terrestrial food webs. Migratory ungulates provide the prey base that maintains large carnivore and scavenger populations and underpins terrestrial biodiversity (fig. S1). When ungulates move in large aggregations, their hooves, feces, and urine create conditions that facilitate distinct biotic communities. The migrations of ungulates have sustained humans for thousands of years, forming tight cultural links among Indigenous people and local communities. Yet ungulate migrations are disappearing at an alarming rate ( 1 ). Efforts by wildlife managers and conservationists are thwarted by a singular challenge: Most ungulate migrations have never been mapped in sufficient detail to guide effective conservation. Without a strategic and collaborative effort, many of the world's great migrations will continue to be truncated, severed, or lost in the coming decades. Fortunately, a combination of animal tracking datasets, historical records, and local and Indigenous knowledge can form the basis for a global atlas of migrations, designed to support conservation action and policy at local, national, and international levels.

Science↗

Ecological insights from three decades of animal movement tracking across a changing Arctic

The Arctic is entering a new ecological state, with alarming consequences for humanity. Animal-borne sensors offer a window into these changes. Although substantial animal tracking data from the Arctic and subarctic exist, most are difficult to discover and access. Here, we present the new Arctic Animal Movement Archive (AAMA), a growing collection of more than 200 standardized terrestrial and marine animal tracking studies from 1991 to the present. The AAMA supports public data discovery, preserves fundamental baseline data for the future, and facilitates efficient, collaborative data analysis. With AAMA-based case studies, we document climatic influences on the migration phenology of eagles, geographic differences in the adaptive response of caribou reproductive phenology to climate change, and species-specific changes in terrestrial mammal movement rates in response to increasing temperature.

Science↗

The influence of body size, condition, and age on recruitment of four Alaskan brown bear populations

Recruitment of brown bear ( Ursus arctos ) offspring into a population is the product of initial cub production and subsequent survival and is a critical component of overall population status and trend. We investigated the relationship between maternal body size, body condition, and age (as a surrogate for gained experience) and recruitment of dependent offspring (≥1 yr old) in 4 Alaska, USA (2014–2017), brown bear populations using logistic regression. Body size alone was our top predictor of the presence of offspring and appeared in all top models. Our data suggest that bear size is the primary driver of productivity across all 4 study populations, with larger bears having a greater chance of being observed with offspring. The effect of body condition was likely confounded by the increased energetic costs of supporting cubs through time and had a negative relationship with recruitment. Age (experience) was positively related to recruitment. Understanding the relative importance of body size, body condition, and age on the recruitment of offspring provides insights into life-history trade-offs female bears must manage as they strive to meet the nutritional costs of cub production and rearing, while minimizing risks to themselves and their offspring. Further assessment of long-term longitudinal studies of brown bears that assess the lifetime reproductive output of individuals would be highly informative to further assess the effect of experience on recruitment and to support the management of brown bear populations for recovery, conservation, sustained yield, and ecosystem function.

Alaska↗

Mineral constraints on arctic caribou (Rangifer tarandus): a spatial and phenological perspective

Arctic caribou ( Rangifer tarandus ) have the longest terrestrial migration of any ungulate but little is known about the spatial and seasonal variation of minerals in summer forages and the potential impacts of mineral nutrition on the foraging behavior and nutritional condition of arctic caribou. We investigated the phenology, availability, and mechanistic relationships of calcium, phosphorus, magnesium, sodium, potassium, iron, manganese, copper, and zinc in three species of woody browse, three species of graminoids, and one forb preferred by caribou over two transects bisecting the ranges of the Central Arctic (CAH) and Western Arctic (WAH) caribou herds in Alaska. Transects traversed three ecoregions (Coastal Plain, Arctic Foothills and Brooks Range) along known migration paths in the summer ranges of both herds. Concentrations of mineral in forages were compared to estimated dietary requirements of lactating female caribou. Spatial distribution of the abundance of minerals in caribou forage was associated with interactions of soil pH and mineral content, while temporal variation was related to plant maturity, and thus nitrogen and fiber content of forages. Concentrations of sodium were below caribou requirements in all forage species for most of the summer and adequate only on the Coastal Plain during the second half of summer. Phosphorus declined in plants from emergence to senescence and was below requirements in all forages by mid‐summer, while concentrations of copper declined to marginal concentrations at plant senescence. Interactions of sodium with potassium, calcium with phosphorus, and copper with zinc in forages likely exacerbate the constraints of low concentrations sodium, phosphorus, and copper. Forages on the WAH contained significantly more phosphorus and copper than forages collected on the CAH transect. We suspect that migrations of caribou to the Arctic Coastal Plain may allow parturient females to replenish sodium stores depleted by foraging inland through the long arctic winters, while also extending the availability of adequate phosphorus, if animals are able to selectively track emerging waves of forage.

Alaska↗

Body size and lean mass of brown bears across and within four diverse ecosystems

Variation in body size across populations of brown bears (Ursus arctos) is largely a function of the availability and quality of nutritional resources while plasticity within populations reflects utilized niche width with implications for population resiliency. We assessed skull size, body length, and lean mass of adult female and male brown bears in four Alaskan study areas that differed in climate, primary food resources, population density, and harvest regime. Full body-frame size, as evidenced by asymptotic skull size and body length, was achieved by 8 to 14 years of age across populations and sexes. Lean body mass of both sexes continued to increase throughout their life. Differences between populations existed for all morphological measures in both sexes, bears in ecosystems with abundant salmon were generally larger. Within all populations, broad variation was seen in body size measures of adults with females displaying roughly a 2-fold difference in lean mass and males showing a 3- to 4-fold difference. The high level of intraspecific variation seen across and within populations suggests the presence of multiple life-history strategies and niche variation relative to resource partitioning, risk tolerance or aversion, and competition. Further, this level of variation indicates broad potential to adapt to changes within a given ecosystem and across the species’ range.

Alaska↗

Plasticity in physiological condition of female brown bears across diverse ecosystems

Variation in life history strategies facilitates the near global distribution of mammals by expanding realized niche width. We investigated physiological plasticity in the spring body composition of adult female brown bears ( Ursus arctos ) across 4 diverse Alaskan ecosystems. Brown bears are a highly intelligent omnivore with a historic range spanning much of North America, Europe, and Asia. We hypothesized that body mass, fat mass, lean mass, and total caloric content would increase across populations with increasing food resource availability. Throughout their range, brown bears enter a period of torpor during winter months, decreasing their metabolic rate as an adaptation to this period of reduced food availability. They also give birth to and nourish offspring during this time. Due to this specific life history strategy, we further hypothesized that proportional body fat and the proportion of total calories derived from fat would be consistent across populations. Our results supported our first hypothesis: body, fat, and lean masses, and caloric content of bears across populations increased with the quality and abundance of available food. However, the proportional body fat content and proportion of calories from fat differed across populations indicating population-specific strategies to meet the demands of reduced seasonal food availability, offspring production and rearing, and climate as well as some plasticity to respond to environmental change or ecosystem perturbations. Investigations of body condition and energetics benefit from combined assessments of absolute, proportional, and caloric metrics to understand the nuances of brown bear physiological dynamics across and within populations.

Alaska↗

Exposure of Alaska brown bears (Ursus arctos) to bacterial, viral, and parasitic agents varies spatiotemporally and may be influenced by age

We collected blood and serum from 155 brown bears ( Ursus arctos ) inhabiting five locations in Alaska during 2013–16 and tested samples for evidence of prior exposure to a suite of bacterial, viral, and parasitic agents. Antibody seroprevalence among Alaska brown bears was estimated to be 15% for Brucella spp., 10% for Francisella tularensis , 7% for Leptospira spp., 18% for canine adenovirus type 1 (CAV-1), 5% for canine distemper virus (CDV), 5% for canine parvovirus, 5% for influenza A virus (IAV), and 44% for Toxoplasma gondii . No samples were seropositive for antibodies to Trichinella spp. Point estimates of prior exposure to pathogens among brown bears at previously unsampled locations generally fell within the range of estimates for previously or contemporaneously sampled bears in Alaska. Statistical support was found for variation in antibody seroprevalence among bears by location or age cohort for CAV-1, CDV, IAV, and Toxoplasma gondii . There was limited concordance in comparisons between our results and previous serosurveys regarding spatial and age-related trends in antibody seroprevalence among Alaska brown bears suggestive of temporal variation. However, we found evidence that the seroprevalence of CAV-1 antibodies is consistently high in bears inhabiting SW Alaska and the cumulative probability of exposure may increase with age. We found evidence for seroconversion or seroreversion to six different infectious agents in one or more bears. Results of this study increase our collective understanding of disease risk to both Alaska brown bear populations and humans that utilize this resource.

Alaska↗

Using gene transcription to assess ecological and anthropological stressors in brown bears

Increasingly, population- and ecosystem-level health assessments are performed using sophisticated molecular tools. Advances in molecular technology enable the identification of synergistic effects of multiple stressors on the individual physiology of different species. Brown bears ( Ursus arctos ) are an apex predator; thus, they are ideal candidates for detecting potentially ecosystem-level systemic perturbations using molecular-based tools. We used gene transcription to analyze 130 brown bear samples from three National Parks and Preserves in Alaska. Although the populations we studied are apparently stable in abundance and exist within protected and intact environments, differences in transcript profiles were noted. The most prevalent differences were among locations. The transcript patterns among groups reflect the influence of environmental factors, such as nutritional status, disease, and xenobiotic exposure. However, these profiles also likely represent baselines for each unique environment by which future measures can be made to identify early indication of population-level changes due to, for example, increasing Arctic temperatures. Some of those environmental changes are predicted to be potentially positive for brown bears, but other effects such as the manifestation of disease or indirect effects of oceanic acidification may produce negative impacts.

Alaska↗

Evaluating potential effects of an industrial road on winter habitat of caribou in North-Central Alaska

Worldwide, some caribou (Rangifer tarandus) populations are experiencing declines due partially to the expansion of industrial development. Caribou can exhibit behavioral avoidance of development, leading to indirect habitat loss, even if the actual footprint is small. Thus, it is important to understand before construction begins how much habitat might be affected by proposed development. In northern Alaska, an industrial road that has been proposed to facilitate mining transects a portion of the Western Arctic caribou herd's winter range. To understand how winter habitat use might be affected by the road, we estimated resource selection patterns during winter for caribou in a study area surrounding the proposed road. We assessed the reductions of habitat value associated with three proposed routes at three distance thresholds for disturbance. High-value winter habitat tended to occur in locally rugged areas that have not burned recently and have a high density of lichen and early dates of spring snowmelt. We found that 1.5% to 8.5% (146-848 km2) of existing high-value winter habitat in our study area might be reduced in quality. The three alternative routes were only marginally different. Our results suggest that the road would have minimal direct effects on high-value winter habitat; however, additional cumulative impacts to caribou (e.g., increased access by recreationists and hunters) should be considered before the full effects of the road can be estimated.

Alaska↗

Can antibrowsing defense regulate the spread of woody vegetation in arctic tundra?

Global climate warming is projected to promote the increase of woody plants, especially shrubs, in arctic tundra. Many factors may affect the extent of this increase, including browsing by mammals. We hypothesize that across the Arctic the effect of browsing will vary because of regional variation in antibrowsing chemical defense. Using birch ( Betula ) as a case study, we propose that browsing is unlikely to retard birch expansion in the region extending eastward from the Lena River in central Siberia across Beringia and the continental tundra of central and eastern Canada where the more effectively defended resin birches predominate. Browsing is more likely to retard birch expansion in tundra west of the Lena to Fennoscandia, Iceland, Greenland and South Baffin Island where the less effectively defended non-resin birches predominate. Evidence from the literature supports this hypothesis. We further suggest that the effect of warming on the supply of plant-available nitrogen will not significantly change either this pan-Arctic pattern of variation in antibrowsing defense or the resultant effect that browsing has on birch expansion in tundra. However, within central and east Beringia warming-caused increases in plant-available nitrogen combined with wildfire could initiate amplifying feedback loops that could accelerate shrubification of tundra by the more effectively defended resin birches. This accelerated shrubification of tundra by resin birch, if extensive, could reduce the food supply of caribou causing population declines. We conclude with a brief discussion of modeling methods that show promise in projecting invasion of tundra by woody plants.

Ecography↗

Fire, grazing history, lichen abundance, and winter distribution of caribou in Alaska's taiga

In the early 1990s the Nelchina Caribou ( Rangifer tarandus ) Herd (NCH) began a dramatic shift to its current winter range, migrating at least an additional 100 km beyond its historic range. We evaluated the impacts of fire and grazing history on lichen abundance and subsequent use and distribution by the NCH. Historic (prior to 1990) and current (2002) winter ranges of the NCH had similar vascular vegetation, lichen cover ( P = 0.491), and fire histories ( P = 0.535), but the former range had significantly less forage lichen biomass as a result of grazing by caribou. Biomass of forage lichens was twice as great overall ( P = 0.031) and 4 times greater in caribou selected sites on the current range than in the historic range, greatly increasing availability to caribou. Caribou on the current range selected for stands with >20% lichen cover ( P < 0.001), greater than 1,250 kg/ha ( P < 0.001) forage lichen biomass and stands older than 80 yr postfire ( P < 0.001). After fires, forage lichen cover and biomass seldom recovered sufficiently to attract caribou grazing until after ≥60 yr, and, as a group, primary forage lichen species did not reach maximum abundance until 180 yr postfire. Recovery following overgrazing can occur much more quickly because lichen cover, albeit mostly fragments, and organic substrates remain present. Our results provide benchmarks for wildlife managers assessing condition of caribou winter range and predicting effects of fires on lichen abundance and caribou distribution. Of our measurements of cover and biomass by species, densities and heights of trees, elevation, slope and aspect, only percentage cover by Cladonia amaurocraea , Cladina rangiferina , Flavocetraria cuculata , and lowbush cranberry ( Vaccinium vitis‐idaea ) were necessary for predicting caribou use of winter range.

Journal of Wildlife Management↗