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USGS2015

Increased land use by Chukchi Sea polar bears in relation to changing sea ice conditions

Recent observations suggest that polar bears ( Ursus maritimus ) are increasingly using land habitats in some parts of their range, where they have minimal access to their preferred prey, likely in response to loss of their sea ice habitat associated with climatic warming. We used location data from female polar bears fit with satellite radio collars to compare land use patterns in the Chukchi Sea between two periods (1986–1995 and 2008–2013) when substantial summer sea-ice loss occurred. In both time periods, polar bears predominantly occupied sea-ice, although land was used during the summer sea-ice retreat and during the winter for maternal denning. However, the proportion of bears on land for > 7 days between August and October increased between the two periods from 20.0% to 38.9%, and the average duration on land increased by 30 days. The majority of bears that used land in the summer and for denning came to Wrangel and Herald Islands (Russia), highlighting the importance of these northernmost land habitats to Chukchi Sea polar bears. Where bears summered and denned, and how long they spent there, was related to the timing and duration of sea ice retreat. Our results are consistent with other studies supporting increased land use as a common response of polar bears to sea-ice loss. Implications of increased land use for Chukchi Sea polar bears are unclear, because a recent study observed no change in body condition or reproductive indices between the two periods considered here. This result suggests that the ecology of this region may provide a degree of resilience to sea ice loss. However, projections of continued sea ice loss suggest that polar bears in the Chukchi Sea and other parts of the Arctic may increasingly use land habitats in the future, which has the potential to increase nutritional stress and human-polar bear interactions.

Chukchi Sea↗

USGS2016

Rapid environmental change drives increased land use by an Arctic marine predator

In the Arctic Ocean’s southern Beaufort Sea (SB), the length of the sea ice melt season (i.e., period between the onset of sea ice break-up in summer and freeze-up in fall) has increased substantially since the late 1990s. Historically, polar bears ( Ursus maritimus ) of the SB have mostly remained on the sea ice year-round (except for those that came ashore to den), but recent changes in the extent and phenology of sea ice habitat have coincided with evidence that use of terrestrial habitat is increasing. We characterized the spatial behavior of polar bears spending summer and fall on land along Alaska’s north coast to better understand the nexus between rapid environmental change and increased use of terrestrial habitat. We found that the percentage of radiocollared adult females from the SB subpopulation coming ashore has tripled over 15 years. Moreover, we detected trends of earlier arrival on shore, increased length of stay, and later departure back to sea ice, all of which were related to declines in the availability of sea ice habitat over the continental shelf and changes to sea ice phenology. Since the late 1990s, the mean duration of the open-water season in the SB increased by 36 days, and the mean length of stay on shore increased by 31 days. While on shore, the distribution of polar bears was influenced by the availability of scavenge subsidies in the form of subsistence-harvested bowhead whale ( Balaena mysticetus ) remains aggregated at sites along the coast. The declining spatio-temporal availability of sea ice habitat and increased availability of human-provisioned resources are likely to result in increased use of land. Increased residency on land is cause for concern given that, while there, bears may be exposed to a greater array of risk factors including those associated with increased human activities.

Alaska, Yukon↗

USGS2018

Estimating population extinction thresholds with categorical classification trees for Louisiana black bears

Monitoring vulnerable species is critical for their conservation. Thresholds or tipping points are commonly used to indicate when populations become vulnerable to extinction and to trigger changes in conservation actions. However, quantitative methods to determine such thresholds have not been well explored. The Louisiana black bear ( Ursus americanus luteolus ) was removed from the list of threatened and endangered species under the U.S. Endangered Species Act in 2016 and our objectives were to determine the most appropriate parameters and thresholds for monitoring and management action. Capture mark recapture (CMR) data from 2006 to 2012 were used to estimate population parameters and variances. We used stochastic population simulations and conditional classification trees to identify demographic rates for monitoring that would be most indicative of heighted extinction risk. We then identified thresholds that would be reliable predictors of population viability. Conditional classification trees indicated that annual apparent survival rates for adult females averaged over 5 years ( ) was the best predictor of population persistence. Specifically, population persistence was estimated to be ≥95% over 100 years when , suggesting that this statistic can be used as threshold to trigger management intervention. Our evaluation produced monitoring protocols that reliably predicted population persistence and was cost-effective. We conclude that population projections and conditional classification trees can be valuable tools for identifying extinction thresholds used in monitoring programs.

PLoS ONE↗

USGS2020

Identifying reliable indicators of fitness in polar bears

Animal structural body size and condition are often measured to evaluate individual health, identify responses to environmental change and food availability, and relate food availability to effects on reproduction and survival. A variety of condition metrics have been developed but relationships between these metrics and vital rates are rarely validated. Identifying an optimal approach to estimate the body condition of polar bears is needed to improve monitoring of their response to decline in sea ice habitat. Therefore, we examined relationships between several commonly used condition indices (CI), body mass, and size with female reproductive success and cub survival among polar bears ( Ursus maritimus ) measured in two subpopulations over three decades. To improve measurement and application of morphometrics and CIs, we also examined whether CIs are independent of age and structural size–an important assumption for monitoring temporal trends—and factors affecting measurement precision and accuracy. Maternal CIs and mass measured the fall prior to denning were related to cub production. Similarly, maternal CIs, mass, and length were related to the mass of cubs or yearlings that accompanied her. However, maternal body mass, but not CIs, measured in the spring was related to cub production and only maternal mass and length were related to the probability of cub survival. These results suggest that CIs may not be better indicators of fitness than body mass in part because CIs remove variation associated with body size that is important in affecting fitness. Further, CIs exhibited variable relationships with age for growing bears and were lower for longer bears despite body length being related to cub survival and female reproductive success. These results are consistent with findings from other species indicating that body mass is a useful metric to link environmental conditions and population dynamics.

PLoS ONE↗

USGS2021

The smell of success: Reproductive success related to rub behavior in brown bears

Several species of bears are known to rub deliberately against trees and other objects, but little is known about why bears rub. Patterns in rubbing behavior of male and female brown bears ( Ursus arctos ) suggest that scent marking via rubbing functions to communicate among potential mates or competitors. Using DNA from bear hairs collected from rub objects in southwestern Alberta from 2011–2014 and existing DNA datasets from Montana and southeastern British Columbia, we determined sex and individual identity of each bear detected. Using these data, we completed a parentage analysis. From the parentage analysis and detection data, we determined the number of offspring, mates, unique rub objects where an individual was detected, and sampling occasions during which an individual was detected for each brown bear identified through our sampling methods. Using a Poisson regression, we found a positive relationship between bear rubbing behavior and reproductive success; both male and female bears with a greater number of mates and a greater number of offspring were detected at more rub objects and during more occasions. Our results suggest a fitness component to bear rubbing, indicate that rubbing is adaptive, and provide insight into a poorly understood behaviour.

Wyoming, Montana, British Columbia, Alberta↗

USGS2021

Age-structured Jolly-Seber model expands inference and improves parameter estimation from capture-recapture data

Understanding the influence of individual attributes on demographic processes is a key objective of wildlife population studies. Capture-recapture and age data are commonly collected to investigate hypotheses about survival, reproduction, and viability. We present a novel age-structured Jolly-Seber model that incorporates age and capture-recapture data to provide comprehensive information on population dynamics, including abundance, age-dependent survival, recruitment, age structure, and population growth rates. We applied our model to a multi-year capture-recapture study of polar bears (Ursus maritimus) in western Hudson Bay, Canada (20122018), where management and conservation require a detailed understanding of how polar bears respond to climate change and other factors. In simulation studies, the age-structured Jolly-Seber model improved precision of survival, recruitment, and annual abundance estimates relative to standard Jolly-Seber models that omit age information. Furthermore, incorporating age information improved precision of population growth rates, increased power to detect trends in abundance, and allowed direct estimation of age-dependent survival and changes in annual age structure. Our case study provided detailed evidence for senescence in polar bear survival. Median survival estimates were lower (<0.95) for individuals aged <5 years, remained high (>0.95) for individuals aged 722 years, and subsequently declined to near zero for individuals >30 years. We also detected cascading effects of large recruitment classes on population age structure, which created major shifts in age structure when these classes entered the population and then again when they reached prime breeding ages (1015 years old). Overall, age-structured Jolly-Seber models provide a flexible means to investigate ecological and evolutionary processes that shape populations (e.g., via senescence, life expectancy, and lifetime reproductive success) while improving our ability to investigate population dynamics and forecast population changes from capture-recapture data.

PLoS ONE↗

USGS2024

Predicting future grizzly bear habitat use in the Bitterroot Ecosystem under recolonization and reintroduction scenarios

Many conservation actions must be implemented with limited data. This is especially true when planning recovery efforts for extirpated populations, such as grizzly bears ( Ursus arctos ) within the Bitterroot Ecosystem (BE), where strategies for reestablishing a resident population are being evaluated. Here, we applied individual-based movement models developed for a nearby grizzly bear population to predict habitat use in and near the BE, under scenarios of natural recolonization, reintroduction, and a combination. All simulations predicted that habitat use by grizzly bears would be higher in the northern half of the study area. Under the natural recolonization scenario, use was concentrated in Montana, but became more uniform across the northern BE in Idaho over time. Use was more concentrated in east-central Idaho under the reintroduction scenario. Assuming that natural recolonization continues even if bears are reintroduced, use remained widespread across the northern half of the BE and surrounding areas. Predicted habitat maps for the natural recolonization scenario aligned well with outlier and GPS collar data available for grizzly bears in the study area, with Spearman rank correlations of ≥0.93 and mean class values of ≥9.1 (where class 10 was the highest relative predicted use; each class 1–10 represented 10% of the landscape). In total, 52.4% of outlier locations and 79% of GPS collar locations were in class 10 in our predicted habitat maps for natural recolonization. Simulated grizzly bears selected habitats over a much larger landscape than the BE itself under all scenarios, including multiple-use and private lands, similar to existing populations that have expanded beyond recovery zones. This highlights the importance of recognizing and planning for the role of private lands in recovery efforts, including understanding resources needed to prevent and respond to human-grizzly bear conflict and maintain public acceptance of grizzly bears over a large landscape.

Idaho, Montana↗

USGS2025

The importance of method selection when estimating diet composition with quantitative fatty acid signature analysis

Quantitative fatty acid signature analysis (QFASA) is a common method of estimating the composition of prey species in the diets of consumers from polar and temperate ecosystems in which lipids are an important source of energy. A key characteristic of QFASA is that the large number of fatty acids that typically comprise lipids permits the dietary contributions of a correspondingly large number of prey types to be estimated. Several modifications to the original QFASA methods have been suggested in the literature and a significant extension of the original model published in 2017 allows simultaneous estimation of both diet proportions and calibration coefficients, which are metabolic constants in the model whose values must otherwise be estimated in independent feeding experiments. However, comparisons of diet estimates obtained using different estimation options have been limited. QFASA has been used to estimate the diet composition of several polar bear ( Ursus maritimus ) subpopulations, including the Southern Beaufort Sea (SBS) subpopulation. Prior QFASA estimates of SBS polar bear diet composition have most often been obtained using variations of the original QFASA model. We investigated the influence of variations in QFASA analytical methods on diet estimates by re-estimating the diet composition of polar bears from the Alaska portion of the SBS using three different methods and found that differences among the three sets of estimates were substantial. Our results illustrate how important the careful and deliberate selection of QFASA methods can be and we provide some guidance on techniques one might use to evaluate options.

PLoS ONE↗

USGS2025

Landscape associations and population genetics of a generalist carnivore at a range limit

American black bear ( Ursus americanus ) sightings have increased in the Oklahoma Panhandle, an area outside of the species’ historical range, prompting an assessment of bears in the region. We used camera traps and an occupancy modeling framework to identify factors influencing bear detection and space-use patterns. We used noninvasive genetic sampling techniques to evaluate genetic diversity, population structure, and bear abundance in the region. During the summers of 2022–2023, we deployed cameras at 160 sites across western Oklahoma (USA) and detected ≥1 bear at 20 sites. The most-supported model from our single-season single-species analysis indicated that bear detection was positively associated with temperature and precipitation, negatively associated with day of year, and differed between years. The most-supported model indicated that bear space use was negatively associated with elevation (β = −0.013, 85% CI = −0.025, 0.000), and positively associated with slope (β = 0.645, 85% CI = 0.305, 0.984) and coarse woody debris counts (β = 1.539, 85% CI = 0.314, 2.765). We deployed 41 hair snares in Oklahoma resulting in the collection of 153 hair samples and received 69 tissue samples from black bears harvested in northeastern New Mexico. Using 11 microsatellite markers, we identified 21 (12M:9F) bears in western Oklahoma, and 69 (40M:29F) in New Mexico. We found evidence that bears occurring in Oklahoma were an extension of a previously documented population that occurred in northcentral New Mexico. We detected significant population-level heterozygote deficiency ( P = 0.013) compared to expectations under Hardy-Weinberg equilibrium. Using capture with replacement models, we estimated 26 (95% CI = 19–43) bears in western Oklahoma during 2022–2023. Our results provide baseline data on population distribution, abundance, and genetic health of bears in the region and identify factors that may drive human-bear conflicts as the bear population increases in western Oklahoma.

New Mexico, Oklahoma↗

USGS1993

Human disturbances of denning polar bears in Alaska

Polar bears ( Ursus maritimus ) give birth in dens of snow and ice. The altricial neonates cannot leave the den for >2 months post-partum and are potentially vulnerable to disturbances near dens. The coastal plain (1002) area of Alaska's Arctic National Wildlife Refuge (ANWR) lies in a region of known polar bear denning and also may contain >9 billion barrels of recoverable oil. Polar bears in dens could be affected in many ways by hydrocarbon development, but neither the distribution of dens nor the sensitivity of bears in dens has been known. I documented the distribution of dens on ANWR between 1981 and 1992 and observed responses of bears in dens to various anthropogenic disturbances. Of 44 dens located by radiotelemetry on the mainland coast of Alaska and Canada, 20 (45%) were on ANWR and 15 (34%) were within the 1002 area. Thus, development of ANWR will increase the potential that denning polar bears are disturbed by human activities. However, perturbations resulting from capture, marking, and radiotracking maternal bears did not affect litter sizes or stature of cubs produced. Likewise, 10 of 12 denned polar bears tolerated exposure to exceptional levels of activity. This tolerance and the fact that investment in the denning effort increases through the winter indicated that spatial and temporal restrictions on developments could prevent the potential for many disruptions of denned bears from being realized.

Alaska, Yukon↗

USGS2013

Mapping polar bear maternal denning habitat in the National Petroleum Reserve -- Alaska with an IfSAR digital terrain model

The National Petroleum Reserve–Alaska (NPR-A) in northeastern Alaska provides winter maternal denning habitat for polar bears ( Ursus maritimus ) and also has high potential for recoverable hydrocarbons. Denning polar bears exposed to human activities may abandon their dens before their young are able to survive the severity of Arctic winter weather. To ensure that wintertime petroleum activities do not threaten polar bears, managers need to know the distribution of landscape features in which maternal dens are likely to occur. Here, we present a map of potential denning habitat within the NPR-A. We used a fine-grain digital elevation model derived from Interferometric Synthetic Aperture Radar (IfSAR) to generate a map of putative denning habitat. We then tested the map’s ability to identify polar bear denning habitat on the landscape. Our final map correctly identified 82% of denning habitat estimated to be within the NPR-A. Mapped denning habitat comprised 19.7 km2 (0.1% of the study area) and was widely dispersed. Though mapping denning habitat with IfSAR data was as effective as mapping with the photogrammetric methods used for other regions of the Alaskan Arctic coastal plain, the use of GIS to analyze IfSAR data allowed greater objectivity and flexibility with less manual labor. Analytical advantages and performance equivalent to that of manual cartographic methods suggest that the use of IfSAR data to identify polar bear maternal denning habitat is a better management tool in the NPR-A and wherever such data may be available.

Alaska↗

USGS2019

Use of subsistence-harvested whale carcasses by polar bears in the southern Beaufort Sea

The availability of a food subsidy has the potential to influence the condition, behavior, fitness, and population dynamics of a species. Since the early 2000s, monitoring efforts along the coast of northern Alaska indicated a higher proportion of polar bears (Ursus maritimus) of the southern Beaufort Sea (SB) subpopulation come onshore and feed at subsistence-harvested bowhead whale (Balaena mysticetus) carcasses during the fall and early winter seasons. This increase in onshore usage by polar bears is a consequence of decreasing sea ice coverage. Concurrently, Indigenous communities annually hunt bowhead whale and deposit the unused remains at localized “bone piles” creating the potential for human-bear interactions. Our objective was to determine the annual number of polar bears feeding at the bone pile near Kaktovik, Alaska. Using a hair snag surrounding the bone pile, we collected hair samples to identify individual bears via microsatellite genotypes during 2011-2014. We used capture-mark-recapture data in the POPAN open-population model to estimate the number of bears visiting the bone pile. We estimated that 146 (SE = 21) bears (sexes combined) used the bone pile in 2012, which represents approximately 16% of the SB polar bear subpopulation. Our results indicated that numerous SB polar bears, males and females, visited the bone pile within a given year. Thus, it will be important to monitor the number of bears using the bone pile and subsequent human-bear interactions and conflicts along the northern coast of Alaska. This is particularly important if polar sea ice continues to recede, which will likely result in increasing numbers of polar bears using the bone pile as a food subsidy.

Alaska↗

USGS2002

The polar bear management agreement for the southern Beaufort Sea: An evaluation of the first ten years of a unique conservation agreement

Polar bears ( Ursus maritimus ) of the southern Beaufort Sea population, distributed from approximately Icy Cape, west of Point Barrow, to Pearce Point, east of Paulatuk in Canada, are harvested by hunters from both countries. In Canada, quotas to control polar bear hunting have been in place, with periodic modifications, since 1968. In Alaska, passage of the United State Marine Mammal Protection Act (MMPA) of 1972 banned polar bear hunting unless done by Alaska Natives for subsistence. However, the MMPA placed no restrictions on numbers or composition of the subsistence hunt, leaving open the potential for an overharvest with no possible legal management response until the population was declared depleted. Recognizing that as a threat to the conservation of the shared polar bear population, the Inuvialuit Game Council from Canada and the North Slop Borough from Alaska negotiated and signed a user-to-user agreement, the Polar Bear Management Agreement for the Southern Beaufort Sea, in 1988. We reviewed the functioning of the agreement through its first 10 years and concluded that, overall, it has been successful because both the total harvest and the proportion of females in the harvest have been contained within sustainable limits. However, harvest monitoring needs to be improved in Alaska, and awareness of the need to prevent overharvest of females needs to be increased in both countries. This agreement is a useful model for other user-to-user conservation agreements.

Beaufort Sea↗

USGS2001

Moose, caribou, and grizzly bear distribution in relation to road traffic in Denali National Park, Alaska

Park managers are concerned that moose ( Alces alces ), caribou ( Rangifer tarandus ), and grizzly bears ( Ursus arctos ) may be avoiding areas along the 130 km road through Denali National Park as a result of high traffic volume, thus decreasing opportunities for visitors to view wildlife. A wildlife monitoring system was developed in 1996 that used 19 landscape level view sheds, stratified into four sections based on decreasing traffic along the road corridor. Data were collected from 22 samplings of all viewsheds during May-August in 1996 and 1997. In 1997, nine backcountry viewsheds were established in three different areas to determine whether density estimates for each species in the backcountry were higher than those for the same animals in similar road-corridor areas. Densities higher than those in the road corridor were found in one backcountry area for moose and in two backcountry areas for grizzly bears. None of the backcountry areas showed a higher density of caribou. We tested hypotheses that moose, caribou, and grizzly bear distributions were unrelated to the road and traffic. Moose sightings were lower than expected within 300 m of the road. More caribou and grizzly bears than expected occurred between 601 and 900 m from the road, while more moose and fewer caribou than expected occurred between 900 and 1200 m from the road. Bull moose in stratum 1 were distributed farther from the road than bulls and cows in stratum 4; cows in stratum 1 and bul1s in stratum 2 were distributed farther from the road than cows in stratum 4. Grizzly bears in stratum 2 were distributed farther from the road than bears in stratum 3. The distribution of moose sightings suggests traffic avoidance, but the spatial pattern of preferred forage may have had more of an influence. Caribou and grizzly bear distributions indicated no pattern of traffic avoidance.

Alaska↗

USGS2001

Remote identification of polar bear maternal den habitat in northern Alaska

Polar bears ( Ursus maritimus ) give birth in dens of ice and snow to protect their altricial young. During the snow-free season, we visited 25 den sites located previously by radiotelemetry and characterized the den site physiognomy. Seven dens occurred in habitats with minimal relief. Eighteen dens (72%) were in coastal and river banks. These "banks" were identifiable on aerial photographs. We then searched high-resolution aerial photographs (n=3000) for habitats similar to those of the 18 dens. On aerial photos, we mapped 1782 km of bank habitats suitable for denning. Bank habitats comprised 0.18% of our study area between the Colville River and the Tamayariak River in northern Alaska. The final map, which correctly identified 88% of bank denning habitat in this region, will help minimize the potential for disruptions of maternal dens by winter petroleum exploration activities.

Alaska↗

USGS2016

Anti-dog IgG secondary antibody successfully detects IgG in a variety of aquatic mammals

Serological tests play an important role in the detection of wildlife diseases. However, while there are many commercial assays and reagents available for domestic species, there is a need to develop efficient serological assays for wildlife. In recent years, marine mammals have represented a wildlife group with emerging infectious diseases, such as influenza, brucellosis, and leptospirosis. However, with the exception of disease-agent-specific assays or functional assays, few reports describe the use of antibody detection assays in marine mammals. In an indirect enzyme-linked immunoassay (EIA) or an immunofluorescence assay, antibody is detected using an antitarget species secondary conjugated antibody. The sensitivity of the assay depends on the avidity of the binding reaction between the bound antibody and the detection antibody. A commercial polyclonal antidog IgG conjugated antibody was tested in an EIA for its ability to sensitively detect the IgG of seven marine mammals including sea otter ( Enhydra lutris ), polar bear ( Ursus maritimus ), grey seal ( Halichoerus grypus ), harbor seal ( Phoca vitulina ), northern elephant seal ( Mirounga angustirostris ), California sea lion ( Zalophus californianus ), Pacific walrus ( Odobenus rosmarus ) and one freshwater mammal: Asian small-clawed otter ( Aonyx cinerea ). With the exception of Asian small-clawed sea otters, the detection of IgG in these marine mammals either exceeded or was nearly equal to detection of dog IgG. The use of the tested commercial antidog IgG antibody may be a valid approach to the detection of antibody response to disease in sea mammals.

Journal of Zoo and Wildlife Medicine↗

USGS2007

Selection of den sites by black bears in the southern Appalachians

We evaluated selection of den sites by American black bears (Ursus americanus) in the Pisgah Bear Sanctuary, western North Carolina, by comparing characteristics of dens at 53 den sites with availability of habitat characteristics in annual home ranges of bears and in the study area. We also tested whether den-site selection differed by sex, age, and reproductive status of bears. In addition, we evaluated whether the den component of an existing habitat model for black bears predicted where bears would select den sites. We found bears selected den sites far from gravel roads, on steep slopes, and at high elevations relative to what was available in both annual home ranges and in the study area. Den-site selection did not differ by sex or age, but it differed by reproductive status. Adult females with cubs preferred to den in areas that were relatively far from gravel roads, but adult females without cubs did not. The habitat model overestimated the value of areas near gravel roads, underestimated the value of moderately steep areas, and did not include elevation as a predictor variable. Our results highlight the importance of evaluating den selection in terms of both use and availability of den characteristics. ?? 2007 American Society of Mammalogists.

Journal of Mammalogy↗

USGS2012

Foraging optimally for home ranges

Economic models predict behavior of animals based on the presumption that natural selection has shaped behaviors important to an animal's fitness to maximize benefits over costs. Economic analyses have shown that territories of animals are structured by trade-offs between benefits gained from resources and costs of defending them. Intuitively, home ranges should be similarly structured, but trade-offs are difficult to assess because there are no costs of defense, thus economic models of home-range behavior are rare. We present economic models that predict how home ranges can be efficient with respect to spatially distributed resources, discounted for travel costs, under 2 strategies of optimization, resource maximization and area minimization. We show how constraints such as competitors can influence structure of homes ranges through resource depression, ultimately structuring density of animals within a population and their distribution on a landscape. We present simulations based on these models to show how they can be generally predictive of home-range behavior and the mechanisms that structure the spatial distribution of animals. We also show how contiguous home ranges estimated statistically from location data can be misleading for animals that optimize home ranges on landscapes with patchily distributed resources. We conclude with a summary of how we applied our models to nonterritorial black bears (Ursus americanus) living in the mountains of North Carolina, where we found their home ranges were best predicted by an area-minimization strategy constrained by intraspecific competition within a social hierarchy. Economic models can provide strong inference about home-range behavior and the resources that structure home ranges by offering falsifiable, a priori hypotheses that can be tested with field observations.

Journal of Mammalogy↗